ZOOARQUEOLOGÍA - MORFOMETRÍA by G. Lorena L'Heureux
Resumen Este trabajo estudia los tamaños corporales de las poblaciones modernas de Lama gua-nicoe... more Resumen Este trabajo estudia los tamaños corporales de las poblaciones modernas de Lama gua-nicoe de Patagonia continental e insular (Argentina), a partir del análisis osteométrico de cuatro huesos largos delanteros (húmero, radiocúbito, metacarpo y primera falange). Se analizó una muestra de más de 100 individuos procedentes de las provincias de Río Negro (40°S), Santa Cruz (48-52°S) y Tierra del Fuego (53-54.5°S). Las variaciones morfológicas de los tamaños se evaluaron en función de las variables ecoambientales de: temperatura (máxima/mínima), precipitación (estival, invernal y anual), productividad primaria neta aérea (PPNA) e índice de vegetación diferencial normalizado (NDVI). Los datos fueron procesados mediante diferentes análisis uni y multivariados. Los resultados demostraron que, en una escala macroregional, se registra un gradiente clinal de los tamaños corporales de las poblaciones de guanacos (los individuos de Tierra del Fuego son significativamente más grandes que los de Santa Cruz y éstos más grandes que los de Río Negro). Todos los factores ecoambientales que se asocian fuertemente con los tamaños óseos (temperatura máxima, PPNA, NDVI) covarían en la región de estudio con la latitud. El patrón clinal observado es, en principio, consistente con las expectativas derivadas de la Regla de Bergmann. Igualmente las diferentes variables ambientales y biogeográficas que influyen sobre los patrones de los tamaños de los guanacos modernos de Patagonia, exponen que la relación entre el tamaño corporal y el ambiente es de naturaleza compleja y no se puede explicar de forma simple y directa por un solo factor.
Abstract This paper studies the body sizes of modern, continental and insular Patagonian (Argentina) Lama guanicoe populations based on the osteometric analysis of four forelimb bones (humerus, radio-ulnae, metacarpus and first phalanx). A sample of than 100 individuals from the Provinces of Río Negro (40°S), Santa Cruz (48-52°S) and
Tierra del Fuego (53-54.5°S) were analyzed. The morphological size variations were evaluated considering the eco-environmental variables of temperature (maximum/minimum), precipitation (summer, winter and annual), aerial net primary productivity (ANPP) and normalized difference vegetation index (NDVI). The data was analyzed using a number of uni and multivariate statistical models. The results show that, at a macro-regional scale, there was a clinal variation in the body sizes of guanaco populations (with individuals from Tierra del Fuego being significantly bigger than those from Santa Cruz, with the latter in turn being larger than those from Río Negro). All eco-environmental factors strongly associated with bone sizes (maximum temperature, ANPP, NDVI) co-vary, in the study region depending on latitude. The clinal pattern observed was, in principle, consistent with the expectations of Bergmann’s Rule. Also different environmental and biogeographic variables influenced the size patterns of modern Patagonian guanacos, thereby underscoring that the relationship between body size and environment is of a complex nature - and cannot be explained simply and directly by a single factor.
RESUMEN. Este trabajo aborda la identificación taxonómica de dos especies de camélidos, Lama glam... more RESUMEN. Este trabajo aborda la identificación taxonómica de dos especies de camélidos, Lama glama y Lama guanicoe a partir del análisis morfológico basado en técnicas de morfometría geométrica. Se estudia una muestra de primeras falanges delanteras correspondientes a 19 especímenes modernos y adultos de guanacos silvestres y llamas domésticas, con el objetivo de detectar variaciones morfológicas que permitan diferenciar ambas especies y delinear a futuro un sistema que pueda ser aplicado a contextos arqueológicos. Estas dos especies se encuentran representadas en sitios del Noroeste Argentino desde los ca. 4100 AP, y su identificación posee una importancia fundamental, en tanto representan dos estrategias diferentes de subsistencia complementarias y coexistentes (caza y pastoreo). Si bien los análisis morfogeométricos no diferenciaron significativamente las especies estudiadas, el presente trabajo constituye un avance en el abordaje de la variabilidad morfológica y la identificación taxonómica de elementos poscraneales arqueológicos y paleontológicos de los grandes camélidos sudamericanos.
ABSTRACT. This paper addresses the taxonomic identification of two species of camelids, Lama glama and Lama guanicoe, through the morphological analysis based on techniques of geometric morphometrics. A sample of first front phalanges of 19 modern and adult specimens of wild guanaco and domestic llama were studied in order to detect morphological variations that distinguish both species to delineate, in a future, a system that can be applied to archaeological contexts. The identification of these two species found in Argentina's Northwest sites from ca. 4100 BP is fundamentally important, because represent two different, complementary subsistence strategies (hunting and grazing) that coexisted. Even though morphogeometric analysis did not differentiated significantly the species studied, this paper is a step forward in addressing the morphologic variability and the taxonomic identification of archaeological and paleontological postcranial elements of the large South American camelids.
In order to assess the role of pinnipeds in prehistoric subsistence of hunteregatherers from nort... more In order to assess the role of pinnipeds in prehistoric subsistence of hunteregatherers from northern Patagonia, traditional osteometric studies based on modern collections of Arctocephalus australis (Aa) and Otaria flavescens (Of) with known age and sex are being employed with the aim of identifying species, sex and age-classes found in zooarchaeological assemblages. A preliminary research based on a very small sample of Aa from the Uruguayan coast suggested the absence of changes in the skull size of this species during the last five decades. Of skull size reduction through time was observed as a consequence of a density-dependence process in northern and central Patagonia. In this study we discuss the use of modern analogues for the study of zooarchaeological remains of Aa. With this objective in mind, osteometrical data from modern skulls of 30 adult male Aa collected along northern and central Pata-gonia were analysed to assess potential diachronic changes in body size of this species. The results showed that there is no evidence of change in the body size during the last three decades (1990e2013) in the Aa population from northern and central Patagonia. This result supports the relevance of investigations if there are differences in the body size of living individuals that will be employed as analogues , for evaluating the potential of these measurements before using them in the analysis of zooarchaeological remains. The use of modern animals of Aa in osteometric analysis of fur seal assemblages recovered in archaeological sites at northern Patagonia is reasonable.
Lama guanicoe (Mu¨ ller, 1776) is one of the most important South American large mammal species f... more Lama guanicoe (Mu¨ ller, 1776) is one of the most important South American large mammal species for
both biologists, ecologists and archaeologists, and body size of modern and prehistoric populations of
guanaco has been one of the most studied parameters. In this paper, we evaluate the latitudinal
differences in body size of different modern populations of guanaco from Argentinean Patagonia based
on osteometric data of their long bones. The osteological sample is made of 110 fully-fused individuals
drawn from three different guanaco populations from continental and insular Patagonia covering the
latitudinal interval from S408 to S54.58. Quantitative data for the four long bones selected are analyzed
with univariate and multivariate parametric statistical techniques. Apparently consistent with
Bergmann’s rule, results show that guanaco specimens from S53–54.58 are bigger on average than
those from S48–528 which are, in turn, bigger than those from S408. We observe a high and significant
covariation between different environmental variables (maximum temperature, net primary
production, and winter precipitation) and body size of the guanaco populations. Nevertheless, when
the temporal factor is incorporated into the analysis of this geographical gradient of body size the
scenario becomes more complex. We conclude that the current clinal pattern observed in Patagonia is
the result of several microevolutionary and biogeographic processes related to an adjustment of body
size to different environmental and ecological drivers and to the consequences of isolation in Tierra del
Fuego that occurred around the Pleistocene-Holocene transition.
Análisis arqueofaunísticos realizados en la costa norte del Golfo San Matías (Pcia. de Río Negro,... more Análisis arqueofaunísticos realizados en la costa norte del Golfo San Matías (Pcia. de Río Negro,
Argentina) detectaron la presencia de individuos inmaduros de otáridos en varias localidades
arqueológicas. Dada las similitudes existentes a nivel osteológico (postcraneal) entre las dos especies
identificadas en el registro de la costa patagónica (Otaria flavescens y Arctocephalus australis),
desarrollamos un estudio osteométrico sobre materiales modernos (con edad y sexo conocido), con
el objeto de distinguirlas. Los datos generados fueron contrastados con mediciones obtenidas sobre
elementos inmaduros arqueológicos de la costa norpatagónica argentina. Los resultados evidencian
que ésta metodología tiene un elevado potencial para discriminar las especies y, de esta forma, discutir
cuestiones vinculadas con las estrategias de caza de lobos marinos en el pasado.
Palabras clave: Osteometría, Otaria flavescens, Arctocephalus australis, Zooarqueología.
Norpatagonia.
Archaeofaunal analyses carried out in the northern coast of the San Matías Gulf (Río Negro,
Argentina) have detected the presence of immature individuals of otariids in several sites. Due to
the postcranial osteological similarities between the two recognized species (Otaria flavescens and
Arctocephalus australis), we developed an osteometrical study on modern materials with known
age and sex, in order to distinguish between them. The generated database was compared with
measurements obtained on immature elements from northern Patagonia archaeological coastal
sites (Argentina). The results show that this method has a high potential for discriminate species
and discussing strategies of sea lions hunting in the past.
Key words: Osteometry, Otaria flavescens, Arctocephalus australis, Zooarchaeology,Northern Patagonia
Lama guanicoe (guanaco) is the best represented land mammal in continental archaeological hunter-... more Lama guanicoe (guanaco) is the best represented land mammal in continental archaeological hunter-gatherer sites during
the Holocene in Patagonia and the Pampas of South America. The main objective of this paper is to develop a model
to estimate the age of immature guanacos by the length of their long bones, which will help discriminate age in young
bone specimens of this species. The individuals studied belong to a comparative collection of modern guanaco specimens
from the Cinco Chañares locality, Rio Negro province, Northern Patagonia, Argentina. The correlation analyses
show that the association between greatest length (GL) and age was very high for all unfused long bones, with highly
significant probability levels (P < 0.01). This allowed us to develop four predictive models to estimate the age-at-death
for juvenile guanacos using a simple regression analysis. The use of new methodological tools that improve knowledge
of age structure in fossil assemblages of Lama guanicoe significantly contribute to the discussion and interpretation of
prey selection strategies and seasonality of guanaco exploitation by Patagonian and Pampean hunter-gatherer groups during the Holocene.
Zooarchaeological analyses of pinniped remains have been scarce in South America because of lack ... more Zooarchaeological analyses of pinniped remains have been scarce in South America because of lack of specific
studies of species determination and estimation of age and sex. Nonetheless, the presence of small and unfused
bones (assigned as pup remains) of Otaria flavescens (O. flavescens) in the Patagonian archaeological record
has been frequently used as an indicator of summer occupation accomplished by ecological analogues. For this
reason, we carried out a postcranial skeletons osteometric study of modern pups of South American sea lions
(O. flavescens) with known sex and aged 0–12months, allowing a more precise age estimation of archaeological
samples. Then, we revisited the evidence for seasonality in the Cabo Virgenes 6 site at Cabo Virgenes locality, one
of the archaeological localities in Patagonia where the seasonality has been determined only on the basis of the
presence of unfused pup bones. Our metric data study on amodern sample shows that the regression analyses
generated a complementary and dependable age estimation model for unfused appendicular bones of
O. flavescens between 0 and 12 months old that can be applied to samples recovered from archaeological sites.
These osteometric studies of postcranial elements allow us to adjust the age estimation of the animal’s death and
re-examine seasonality of theCabo Virgenes 6 site (SantaCruz, Argentina).Copyright©2013 John Wiley & Sons, Ltd.
Key words: South American sea lions; otariids pups; postcranial osteometry; seasonality indicators; hunter–
gatherer; coastal archaeological sites; Patagonia; Late Holocene
A partir de los patrones morfológicos de una población actual de Lama guanicoe se desarrolla una
... more A partir de los patrones morfológicos de una población actual de Lama guanicoe se desarrolla una
técnica que permite derivar la edad de muerte de guanacos juveniles de conjuntos fósiles en base a las dimensiones
de los huesos largos no fusionados. El uso de esta nueva herramientas metodológica permite ajustar
la información de las estructuras de edad de los conjuntos arqueológicos de guanaco y contribuye con la
discusión de las estrategias de caza, selección de presas y estacionalidad de las explotaciones de la principal
presa terrestre de los cazadores-recolectores de Patagonia durante el Holoceno. Como caso de estudio se analiza la muestra ósea de guanaco procedente del sitio Orejas de Burro 1 y se discute la estacionalidad de las ocupaciones humanas durante el Holoceno tardío en el sector argentino del Campo Volcánico de Pali Aike.
Cabo Virgenes 20 (CV20) is an open air archaeological site located in an erosional beach by the s... more Cabo Virgenes 20 (CV20) is an open air archaeological site located in an erosional beach by the southern Atlantic Ocean,
Patagonia, Argentina. According to previous regional research, terrestrial hunter groups began to use this coastal space
marginally -although purposely- since 2000 years BP. Of all the studied sites in Cabo Virgenes archaeological locality,
CV20 has the highest faunal remains density and high richness of exploited species. Analysis of the bone assemblage shows
emphasis on exploitation of seabirds (Phalacrocorax sp.; Aptenodytes patagonica), in addition to an important presence
of South American sea lions (Otaria flavescens) and carnivores (Pseudalopex culpaeus) with indications of anthropic use.
A high concentration of different taxa of molluscs was recorded, but its human consumption can be sustained only by the
site context. In the regional framework of archaeological records in Cabo Virgenes area, CV20 provides new evidence
about the use of the coast and its resources by hunter-gatherers inland populations at around 800-700 years BP.
The assessment of sexual structure of fossil assemblages provide valuable information for the dis... more The assessment of sexual structure of fossil assemblages provide valuable information for the discussion of paleodemographic aspects and hunting strategies employed by prehistoric human groups. The aim of this study is to evaluate the sexual structure of the skeletal remains of guanaco (Lama guanicoe) recovered from the site Paso Otero 1, which is located in the Paso Otero archaeological locality (Necochea District, Buenos Aires Province). The methodology is based on previous studies conducted on a modern population of guanacos from Río Negro province. These results showed that the major osteological differences related to sexual dimorphism in postcranial skeleton correspond to variations in the size and shape of the entire pelvis and especially of the acetabulum. The fossil assemblage of Paso Otero 1 was analyzed by integrating this methodology with qualitative variables observed in the pelvis. This approach allowed to incorporate a greater number of specimens to the analysis and the results indicate that unlike what was previously known, the sexual structure of the assemblage is dominated by females.
Magallania (Punta Arenas), 2013
Revista Magallania, 2013
This paper presents the study of the archaeofaunal remains recovered at the Cerro León 3 archaeol... more This paper presents the study of the archaeofaunal remains recovered at the Cerro León 3 archaeological site (Santa Cruz, Argentina), with Early to Late Holocene chronology. The measurement of taxonomic and skeletal part abundance, the general patterns of bone frequency and the distribution
as well as pre and postdepositional modifications are assessed. The integration of archaeofaunal analyses with the available technological information for CL3 lithic assemblage suggests an increase in occupation intensity by 4.000 BP, when the highest deposition rate for bones is recorded. Afterwards, by 1.700 BP, archaeofauna shows a shift in patterns of primary prey (Lama guanicoe) exploitation, towards an intensification of its processing. This shift is accompanied by an increment in lithic artifact deposition rate, with assemblages characterized by toolstone intensive-use and an increase in the number of lithic
raw material represented.
KEY WORDS : Patagonia, Holocene sequence, hunter-gatherers, archaeofaunal record, land-use intensity.
Este trabajo pretende incluir la discusión de la variabilidad, tanto interespecífica como intraes... more Este trabajo pretende incluir la discusión de la variabilidad, tanto interespecífica como intraespecífica,
en los esquemas de análisis de diferenciación morfológica de las poblaciones modernas de las especies de camélidos sudamericanos. Como caso de estudio se evalúa, a partir de dos pruebas, la técnica de R. Meadows de identificación de taxa arqueológicos a partir de referentes modernos: a) con un camélido indeterminado (AMNH 14121) seleccionando cuatro estándares actuales (dos llamas y dos guanacos), y b) sobre una población representativa de guanacos modernos de Patagonia
en la que se usaron doce referentes actuales seleccionados dentro del rango intercuartil de la población. Con esta experiencia se observó que la técnica de Meadow resulta dependiente de la variación intraespecífica, lo que reduce su eficiencia como herramienta de identificación taxonómica cuando se utilizan especies con rangos de tamaño semejantes. Se concluye que es indispensable comenzar por entender más acerca de la variación intrapoblacional e intraespecífica para poder conocer la diversidad taxonómica.
En poblaciones vivas, los camélidos silvestres sudamericanos son sexados a partir de la observaci... more En poblaciones vivas, los camélidos silvestres sudamericanos son sexados a partir de la observación de su conducta social y reproductiva. Por lo general, tanto biólogos como arqueólogos asumen la ausencia de caracteres morfológicos
que evidencien un dimorfismo sexual marcado entre individuos adultos de guanaco (Lama guanicoe). Los principales estudios que evalúan las diferencias osteológicas entre machos y hembras en esta especie están basados
en análisis de variables morfométricas craneanas y en caracteres morfológicos de la pelvis. En este trabajo se evalúa la existencia y magnitud de variables osteométricas poscraneales de diferenciación sexual en una población moderna de guanacos procedente de la localidad Cinco Chañares, Provincia de Río Negro, Argentina. Se estudiaron 32 individuos (8 machos y 24 hembras) de más de 30 meses en los que se midieron 51 hemipelvis, 40 húmeros, 39 radioulnas, 40 fémures y 41 tibias. Los resultados mostraron que las principales diferencias osteológicas, vinculadas con el dimorfismo sexual, se relacionan con variaciones en el tamaño y la forma de la pelvis. En los huesos largos, las diferencias osteométricas significativas se registraron sólo en variables aisladas y no se pudieron definir diferencias osteométricas asociadas con el dimorfismo sexual. Igualmente se considera que esto deberá ser reevaluado con mayor profundidad considerando una muestra mayor de individuos machos.
El objetivo de este trabajo es evaluar la variación morfológica temporal de los tamaños de elemen... more El objetivo de este trabajo es evaluar la variación morfológica temporal de los tamaños de elementos apendiculares del guanaco (Lama guanicoe) a partir del análisis cuantitativo
de variables morfológicas de variación continua. Los huesos analizados son los metacarpos distales y proximales y las segundas falanges. En el análisis se utilizaron muestras procedentes de cuatro sitios arqueológicos y uno paleontológico del extremo austral de Patagonia
continental (Argentina y Chile). Las mismas representan dos momentos cronológicos: Pleistoceno final (ca. 12000-10000 años AP.) y momentos posteriores al Holoceno temprano y
Holoceno medio (ca. 8000-6000 y 6000-4000 años AP.).
Los datos fueron evaluados a partir del análisis de varianza para inspeccionar las diferencias estadísticas entre los tamaños de los huesos. Los resultados mostraron huesos más grandes durante el Pleistoceno final y tamaños significativamente menores durante el Holoceno temprano/
medio.
A su vez, se estudió la relación existente entre los tamaños de los huesos de guanaco y la variable climática asociada con la temperatura, la cual se considera una de las potenciales causas de variación morfológica entre los mamíferos. Para esto se utilizaron los valores de dióxido de carbono del núcleo de hielo antártico Domo C. En el análisis conjunto de los patrones morfológicos de tamaño y la variable de temperatura (CO2) se registró una covariación negativa entre el proxy de temperatura y las modificaciones halladas en los tamaños de los huesos postcraneales de guanaco. Estos resultados sugieren que los tamaños fueron estadísticamente más grandes durante períodos de bajas temperaturas y se redujeron concomitantemente con el incremento de
este parámetro climático.
The aim of this paper is to present the results of the archaeofaunal analysis carried out on bone... more The aim of this paper is to present the results of the archaeofaunal analysis carried out on bone materials recovered at the Orejas de Burro archaeological site (OB1), in the Pali Aike lava field (“CVPA”), Santa Cruz, Argentina. Two independently
excavated sectors were considered, one corresponding to a burial area located at the bottom of the cave, and the other one corresponding to a test pit placed next to the burial area. Chronologically, two assemblages were identified, the first one corresponding to the early Late Holocene (ca. 3.500 14C years BP), and the second one to the later Late Holocene (ca. 1.700-500 años 14C AP). The obtained results show statistically significant differences in the manifestation of taphonomic variables and in the frequency and diversity of faunal species both, spatially and temporally. In particular, a more intensive use of the cave at around 3.500 14C years BP is inferred, coincidental with a higher representativeness of coastal taxa like mollusk and sea/coastal birds. Such species may have been exploited during logistic movements to the Magellan strait carried out by hunter gatherers from the inland
Resumen Se presenta una propuesta osteométrica para el estudio de muestras óseas de individuos su... more Resumen Se presenta una propuesta osteométrica para el estudio de muestras óseas de individuos subadultos de lobo marino de un pelo (Otaria flavescens), una de las dos especies frecuentemente halladas en el registro arqueológico costero. En este trabajo se exponen los resultados de una prueba de Error Intraobservador (EIO) realizada sobre una muestra compuesta por 10 cachorros de ambos sexos de entre 1,5 y 12 meses de edad procedente del área Islote Lobos, costa de Río Negro (Argentina). En la prueba de EIO se utilizó una guía de mediciones adaptada a la anatomía de Otaria flavescens (117 variables a relevar en 10 elementos poscraneales no fusionados y en la mandíbula). Los datos se evaluaron a partir del análisis de varianza-ANOVA modelo II, two way-y de la estimación del Coeficiente de Correlación Intraclase-ICC-, que permitieron ajustar la guía osteométrica detectando aquellas variables que mostraron baja consistencia en la repetición de la serie de medidas. Se considera que este trabajo representa un aporte metodológico relevante que permitirá la evaluación del tamaño del esqueleto poscraneal de muestras óseas arqueológicas (y modernas) de lobos marinos de un pelo necesarias para investigar aspectos asociados con diferentes problemas que plantean los especímenes óseos de esta especie recuperados en los sitios arqueológicos costeros sudamericanos, tales como la estimación de la edad y el sexo, la determinación específica, entre otros. Palabras clave: Otaria flavescens, osteometría, error intraobservador.
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ZOOARQUEOLOGÍA - MORFOMETRÍA by G. Lorena L'Heureux
Abstract This paper studies the body sizes of modern, continental and insular Patagonian (Argentina) Lama guanicoe populations based on the osteometric analysis of four forelimb bones (humerus, radio-ulnae, metacarpus and first phalanx). A sample of than 100 individuals from the Provinces of Río Negro (40°S), Santa Cruz (48-52°S) and
Tierra del Fuego (53-54.5°S) were analyzed. The morphological size variations were evaluated considering the eco-environmental variables of temperature (maximum/minimum), precipitation (summer, winter and annual), aerial net primary productivity (ANPP) and normalized difference vegetation index (NDVI). The data was analyzed using a number of uni and multivariate statistical models. The results show that, at a macro-regional scale, there was a clinal variation in the body sizes of guanaco populations (with individuals from Tierra del Fuego being significantly bigger than those from Santa Cruz, with the latter in turn being larger than those from Río Negro). All eco-environmental factors strongly associated with bone sizes (maximum temperature, ANPP, NDVI) co-vary, in the study region depending on latitude. The clinal pattern observed was, in principle, consistent with the expectations of Bergmann’s Rule. Also different environmental and biogeographic variables influenced the size patterns of modern Patagonian guanacos, thereby underscoring that the relationship between body size and environment is of a complex nature - and cannot be explained simply and directly by a single factor.
ABSTRACT. This paper addresses the taxonomic identification of two species of camelids, Lama glama and Lama guanicoe, through the morphological analysis based on techniques of geometric morphometrics. A sample of first front phalanges of 19 modern and adult specimens of wild guanaco and domestic llama were studied in order to detect morphological variations that distinguish both species to delineate, in a future, a system that can be applied to archaeological contexts. The identification of these two species found in Argentina's Northwest sites from ca. 4100 BP is fundamentally important, because represent two different, complementary subsistence strategies (hunting and grazing) that coexisted. Even though morphogeometric analysis did not differentiated significantly the species studied, this paper is a step forward in addressing the morphologic variability and the taxonomic identification of archaeological and paleontological postcranial elements of the large South American camelids.
both biologists, ecologists and archaeologists, and body size of modern and prehistoric populations of
guanaco has been one of the most studied parameters. In this paper, we evaluate the latitudinal
differences in body size of different modern populations of guanaco from Argentinean Patagonia based
on osteometric data of their long bones. The osteological sample is made of 110 fully-fused individuals
drawn from three different guanaco populations from continental and insular Patagonia covering the
latitudinal interval from S408 to S54.58. Quantitative data for the four long bones selected are analyzed
with univariate and multivariate parametric statistical techniques. Apparently consistent with
Bergmann’s rule, results show that guanaco specimens from S53–54.58 are bigger on average than
those from S48–528 which are, in turn, bigger than those from S408. We observe a high and significant
covariation between different environmental variables (maximum temperature, net primary
production, and winter precipitation) and body size of the guanaco populations. Nevertheless, when
the temporal factor is incorporated into the analysis of this geographical gradient of body size the
scenario becomes more complex. We conclude that the current clinal pattern observed in Patagonia is
the result of several microevolutionary and biogeographic processes related to an adjustment of body
size to different environmental and ecological drivers and to the consequences of isolation in Tierra del
Fuego that occurred around the Pleistocene-Holocene transition.
Argentina) detectaron la presencia de individuos inmaduros de otáridos en varias localidades
arqueológicas. Dada las similitudes existentes a nivel osteológico (postcraneal) entre las dos especies
identificadas en el registro de la costa patagónica (Otaria flavescens y Arctocephalus australis),
desarrollamos un estudio osteométrico sobre materiales modernos (con edad y sexo conocido), con
el objeto de distinguirlas. Los datos generados fueron contrastados con mediciones obtenidas sobre
elementos inmaduros arqueológicos de la costa norpatagónica argentina. Los resultados evidencian
que ésta metodología tiene un elevado potencial para discriminar las especies y, de esta forma, discutir
cuestiones vinculadas con las estrategias de caza de lobos marinos en el pasado.
Palabras clave: Osteometría, Otaria flavescens, Arctocephalus australis, Zooarqueología.
Norpatagonia.
Archaeofaunal analyses carried out in the northern coast of the San Matías Gulf (Río Negro,
Argentina) have detected the presence of immature individuals of otariids in several sites. Due to
the postcranial osteological similarities between the two recognized species (Otaria flavescens and
Arctocephalus australis), we developed an osteometrical study on modern materials with known
age and sex, in order to distinguish between them. The generated database was compared with
measurements obtained on immature elements from northern Patagonia archaeological coastal
sites (Argentina). The results show that this method has a high potential for discriminate species
and discussing strategies of sea lions hunting in the past.
Key words: Osteometry, Otaria flavescens, Arctocephalus australis, Zooarchaeology,Northern Patagonia
the Holocene in Patagonia and the Pampas of South America. The main objective of this paper is to develop a model
to estimate the age of immature guanacos by the length of their long bones, which will help discriminate age in young
bone specimens of this species. The individuals studied belong to a comparative collection of modern guanaco specimens
from the Cinco Chañares locality, Rio Negro province, Northern Patagonia, Argentina. The correlation analyses
show that the association between greatest length (GL) and age was very high for all unfused long bones, with highly
significant probability levels (P < 0.01). This allowed us to develop four predictive models to estimate the age-at-death
for juvenile guanacos using a simple regression analysis. The use of new methodological tools that improve knowledge
of age structure in fossil assemblages of Lama guanicoe significantly contribute to the discussion and interpretation of
prey selection strategies and seasonality of guanaco exploitation by Patagonian and Pampean hunter-gatherer groups during the Holocene.
studies of species determination and estimation of age and sex. Nonetheless, the presence of small and unfused
bones (assigned as pup remains) of Otaria flavescens (O. flavescens) in the Patagonian archaeological record
has been frequently used as an indicator of summer occupation accomplished by ecological analogues. For this
reason, we carried out a postcranial skeletons osteometric study of modern pups of South American sea lions
(O. flavescens) with known sex and aged 0–12months, allowing a more precise age estimation of archaeological
samples. Then, we revisited the evidence for seasonality in the Cabo Virgenes 6 site at Cabo Virgenes locality, one
of the archaeological localities in Patagonia where the seasonality has been determined only on the basis of the
presence of unfused pup bones. Our metric data study on amodern sample shows that the regression analyses
generated a complementary and dependable age estimation model for unfused appendicular bones of
O. flavescens between 0 and 12 months old that can be applied to samples recovered from archaeological sites.
These osteometric studies of postcranial elements allow us to adjust the age estimation of the animal’s death and
re-examine seasonality of theCabo Virgenes 6 site (SantaCruz, Argentina).Copyright©2013 John Wiley & Sons, Ltd.
Key words: South American sea lions; otariids pups; postcranial osteometry; seasonality indicators; hunter–
gatherer; coastal archaeological sites; Patagonia; Late Holocene
técnica que permite derivar la edad de muerte de guanacos juveniles de conjuntos fósiles en base a las dimensiones
de los huesos largos no fusionados. El uso de esta nueva herramientas metodológica permite ajustar
la información de las estructuras de edad de los conjuntos arqueológicos de guanaco y contribuye con la
discusión de las estrategias de caza, selección de presas y estacionalidad de las explotaciones de la principal
presa terrestre de los cazadores-recolectores de Patagonia durante el Holoceno. Como caso de estudio se analiza la muestra ósea de guanaco procedente del sitio Orejas de Burro 1 y se discute la estacionalidad de las ocupaciones humanas durante el Holoceno tardío en el sector argentino del Campo Volcánico de Pali Aike.
Patagonia, Argentina. According to previous regional research, terrestrial hunter groups began to use this coastal space
marginally -although purposely- since 2000 years BP. Of all the studied sites in Cabo Virgenes archaeological locality,
CV20 has the highest faunal remains density and high richness of exploited species. Analysis of the bone assemblage shows
emphasis on exploitation of seabirds (Phalacrocorax sp.; Aptenodytes patagonica), in addition to an important presence
of South American sea lions (Otaria flavescens) and carnivores (Pseudalopex culpaeus) with indications of anthropic use.
A high concentration of different taxa of molluscs was recorded, but its human consumption can be sustained only by the
site context. In the regional framework of archaeological records in Cabo Virgenes area, CV20 provides new evidence
about the use of the coast and its resources by hunter-gatherers inland populations at around 800-700 years BP.
as well as pre and postdepositional modifications are assessed. The integration of archaeofaunal analyses with the available technological information for CL3 lithic assemblage suggests an increase in occupation intensity by 4.000 BP, when the highest deposition rate for bones is recorded. Afterwards, by 1.700 BP, archaeofauna shows a shift in patterns of primary prey (Lama guanicoe) exploitation, towards an intensification of its processing. This shift is accompanied by an increment in lithic artifact deposition rate, with assemblages characterized by toolstone intensive-use and an increase in the number of lithic
raw material represented.
KEY WORDS : Patagonia, Holocene sequence, hunter-gatherers, archaeofaunal record, land-use intensity.
en los esquemas de análisis de diferenciación morfológica de las poblaciones modernas de las especies de camélidos sudamericanos. Como caso de estudio se evalúa, a partir de dos pruebas, la técnica de R. Meadows de identificación de taxa arqueológicos a partir de referentes modernos: a) con un camélido indeterminado (AMNH 14121) seleccionando cuatro estándares actuales (dos llamas y dos guanacos), y b) sobre una población representativa de guanacos modernos de Patagonia
en la que se usaron doce referentes actuales seleccionados dentro del rango intercuartil de la población. Con esta experiencia se observó que la técnica de Meadow resulta dependiente de la variación intraespecífica, lo que reduce su eficiencia como herramienta de identificación taxonómica cuando se utilizan especies con rangos de tamaño semejantes. Se concluye que es indispensable comenzar por entender más acerca de la variación intrapoblacional e intraespecífica para poder conocer la diversidad taxonómica.
que evidencien un dimorfismo sexual marcado entre individuos adultos de guanaco (Lama guanicoe). Los principales estudios que evalúan las diferencias osteológicas entre machos y hembras en esta especie están basados
en análisis de variables morfométricas craneanas y en caracteres morfológicos de la pelvis. En este trabajo se evalúa la existencia y magnitud de variables osteométricas poscraneales de diferenciación sexual en una población moderna de guanacos procedente de la localidad Cinco Chañares, Provincia de Río Negro, Argentina. Se estudiaron 32 individuos (8 machos y 24 hembras) de más de 30 meses en los que se midieron 51 hemipelvis, 40 húmeros, 39 radioulnas, 40 fémures y 41 tibias. Los resultados mostraron que las principales diferencias osteológicas, vinculadas con el dimorfismo sexual, se relacionan con variaciones en el tamaño y la forma de la pelvis. En los huesos largos, las diferencias osteométricas significativas se registraron sólo en variables aisladas y no se pudieron definir diferencias osteométricas asociadas con el dimorfismo sexual. Igualmente se considera que esto deberá ser reevaluado con mayor profundidad considerando una muestra mayor de individuos machos.
de variables morfológicas de variación continua. Los huesos analizados son los metacarpos distales y proximales y las segundas falanges. En el análisis se utilizaron muestras procedentes de cuatro sitios arqueológicos y uno paleontológico del extremo austral de Patagonia
continental (Argentina y Chile). Las mismas representan dos momentos cronológicos: Pleistoceno final (ca. 12000-10000 años AP.) y momentos posteriores al Holoceno temprano y
Holoceno medio (ca. 8000-6000 y 6000-4000 años AP.).
Los datos fueron evaluados a partir del análisis de varianza para inspeccionar las diferencias estadísticas entre los tamaños de los huesos. Los resultados mostraron huesos más grandes durante el Pleistoceno final y tamaños significativamente menores durante el Holoceno temprano/
medio.
A su vez, se estudió la relación existente entre los tamaños de los huesos de guanaco y la variable climática asociada con la temperatura, la cual se considera una de las potenciales causas de variación morfológica entre los mamíferos. Para esto se utilizaron los valores de dióxido de carbono del núcleo de hielo antártico Domo C. En el análisis conjunto de los patrones morfológicos de tamaño y la variable de temperatura (CO2) se registró una covariación negativa entre el proxy de temperatura y las modificaciones halladas en los tamaños de los huesos postcraneales de guanaco. Estos resultados sugieren que los tamaños fueron estadísticamente más grandes durante períodos de bajas temperaturas y se redujeron concomitantemente con el incremento de
este parámetro climático.
excavated sectors were considered, one corresponding to a burial area located at the bottom of the cave, and the other one corresponding to a test pit placed next to the burial area. Chronologically, two assemblages were identified, the first one corresponding to the early Late Holocene (ca. 3.500 14C years BP), and the second one to the later Late Holocene (ca. 1.700-500 años 14C AP). The obtained results show statistically significant differences in the manifestation of taphonomic variables and in the frequency and diversity of faunal species both, spatially and temporally. In particular, a more intensive use of the cave at around 3.500 14C years BP is inferred, coincidental with a higher representativeness of coastal taxa like mollusk and sea/coastal birds. Such species may have been exploited during logistic movements to the Magellan strait carried out by hunter gatherers from the inland
Abstract This paper studies the body sizes of modern, continental and insular Patagonian (Argentina) Lama guanicoe populations based on the osteometric analysis of four forelimb bones (humerus, radio-ulnae, metacarpus and first phalanx). A sample of than 100 individuals from the Provinces of Río Negro (40°S), Santa Cruz (48-52°S) and
Tierra del Fuego (53-54.5°S) were analyzed. The morphological size variations were evaluated considering the eco-environmental variables of temperature (maximum/minimum), precipitation (summer, winter and annual), aerial net primary productivity (ANPP) and normalized difference vegetation index (NDVI). The data was analyzed using a number of uni and multivariate statistical models. The results show that, at a macro-regional scale, there was a clinal variation in the body sizes of guanaco populations (with individuals from Tierra del Fuego being significantly bigger than those from Santa Cruz, with the latter in turn being larger than those from Río Negro). All eco-environmental factors strongly associated with bone sizes (maximum temperature, ANPP, NDVI) co-vary, in the study region depending on latitude. The clinal pattern observed was, in principle, consistent with the expectations of Bergmann’s Rule. Also different environmental and biogeographic variables influenced the size patterns of modern Patagonian guanacos, thereby underscoring that the relationship between body size and environment is of a complex nature - and cannot be explained simply and directly by a single factor.
ABSTRACT. This paper addresses the taxonomic identification of two species of camelids, Lama glama and Lama guanicoe, through the morphological analysis based on techniques of geometric morphometrics. A sample of first front phalanges of 19 modern and adult specimens of wild guanaco and domestic llama were studied in order to detect morphological variations that distinguish both species to delineate, in a future, a system that can be applied to archaeological contexts. The identification of these two species found in Argentina's Northwest sites from ca. 4100 BP is fundamentally important, because represent two different, complementary subsistence strategies (hunting and grazing) that coexisted. Even though morphogeometric analysis did not differentiated significantly the species studied, this paper is a step forward in addressing the morphologic variability and the taxonomic identification of archaeological and paleontological postcranial elements of the large South American camelids.
both biologists, ecologists and archaeologists, and body size of modern and prehistoric populations of
guanaco has been one of the most studied parameters. In this paper, we evaluate the latitudinal
differences in body size of different modern populations of guanaco from Argentinean Patagonia based
on osteometric data of their long bones. The osteological sample is made of 110 fully-fused individuals
drawn from three different guanaco populations from continental and insular Patagonia covering the
latitudinal interval from S408 to S54.58. Quantitative data for the four long bones selected are analyzed
with univariate and multivariate parametric statistical techniques. Apparently consistent with
Bergmann’s rule, results show that guanaco specimens from S53–54.58 are bigger on average than
those from S48–528 which are, in turn, bigger than those from S408. We observe a high and significant
covariation between different environmental variables (maximum temperature, net primary
production, and winter precipitation) and body size of the guanaco populations. Nevertheless, when
the temporal factor is incorporated into the analysis of this geographical gradient of body size the
scenario becomes more complex. We conclude that the current clinal pattern observed in Patagonia is
the result of several microevolutionary and biogeographic processes related to an adjustment of body
size to different environmental and ecological drivers and to the consequences of isolation in Tierra del
Fuego that occurred around the Pleistocene-Holocene transition.
Argentina) detectaron la presencia de individuos inmaduros de otáridos en varias localidades
arqueológicas. Dada las similitudes existentes a nivel osteológico (postcraneal) entre las dos especies
identificadas en el registro de la costa patagónica (Otaria flavescens y Arctocephalus australis),
desarrollamos un estudio osteométrico sobre materiales modernos (con edad y sexo conocido), con
el objeto de distinguirlas. Los datos generados fueron contrastados con mediciones obtenidas sobre
elementos inmaduros arqueológicos de la costa norpatagónica argentina. Los resultados evidencian
que ésta metodología tiene un elevado potencial para discriminar las especies y, de esta forma, discutir
cuestiones vinculadas con las estrategias de caza de lobos marinos en el pasado.
Palabras clave: Osteometría, Otaria flavescens, Arctocephalus australis, Zooarqueología.
Norpatagonia.
Archaeofaunal analyses carried out in the northern coast of the San Matías Gulf (Río Negro,
Argentina) have detected the presence of immature individuals of otariids in several sites. Due to
the postcranial osteological similarities between the two recognized species (Otaria flavescens and
Arctocephalus australis), we developed an osteometrical study on modern materials with known
age and sex, in order to distinguish between them. The generated database was compared with
measurements obtained on immature elements from northern Patagonia archaeological coastal
sites (Argentina). The results show that this method has a high potential for discriminate species
and discussing strategies of sea lions hunting in the past.
Key words: Osteometry, Otaria flavescens, Arctocephalus australis, Zooarchaeology,Northern Patagonia
the Holocene in Patagonia and the Pampas of South America. The main objective of this paper is to develop a model
to estimate the age of immature guanacos by the length of their long bones, which will help discriminate age in young
bone specimens of this species. The individuals studied belong to a comparative collection of modern guanaco specimens
from the Cinco Chañares locality, Rio Negro province, Northern Patagonia, Argentina. The correlation analyses
show that the association between greatest length (GL) and age was very high for all unfused long bones, with highly
significant probability levels (P < 0.01). This allowed us to develop four predictive models to estimate the age-at-death
for juvenile guanacos using a simple regression analysis. The use of new methodological tools that improve knowledge
of age structure in fossil assemblages of Lama guanicoe significantly contribute to the discussion and interpretation of
prey selection strategies and seasonality of guanaco exploitation by Patagonian and Pampean hunter-gatherer groups during the Holocene.
studies of species determination and estimation of age and sex. Nonetheless, the presence of small and unfused
bones (assigned as pup remains) of Otaria flavescens (O. flavescens) in the Patagonian archaeological record
has been frequently used as an indicator of summer occupation accomplished by ecological analogues. For this
reason, we carried out a postcranial skeletons osteometric study of modern pups of South American sea lions
(O. flavescens) with known sex and aged 0–12months, allowing a more precise age estimation of archaeological
samples. Then, we revisited the evidence for seasonality in the Cabo Virgenes 6 site at Cabo Virgenes locality, one
of the archaeological localities in Patagonia where the seasonality has been determined only on the basis of the
presence of unfused pup bones. Our metric data study on amodern sample shows that the regression analyses
generated a complementary and dependable age estimation model for unfused appendicular bones of
O. flavescens between 0 and 12 months old that can be applied to samples recovered from archaeological sites.
These osteometric studies of postcranial elements allow us to adjust the age estimation of the animal’s death and
re-examine seasonality of theCabo Virgenes 6 site (SantaCruz, Argentina).Copyright©2013 John Wiley & Sons, Ltd.
Key words: South American sea lions; otariids pups; postcranial osteometry; seasonality indicators; hunter–
gatherer; coastal archaeological sites; Patagonia; Late Holocene
técnica que permite derivar la edad de muerte de guanacos juveniles de conjuntos fósiles en base a las dimensiones
de los huesos largos no fusionados. El uso de esta nueva herramientas metodológica permite ajustar
la información de las estructuras de edad de los conjuntos arqueológicos de guanaco y contribuye con la
discusión de las estrategias de caza, selección de presas y estacionalidad de las explotaciones de la principal
presa terrestre de los cazadores-recolectores de Patagonia durante el Holoceno. Como caso de estudio se analiza la muestra ósea de guanaco procedente del sitio Orejas de Burro 1 y se discute la estacionalidad de las ocupaciones humanas durante el Holoceno tardío en el sector argentino del Campo Volcánico de Pali Aike.
Patagonia, Argentina. According to previous regional research, terrestrial hunter groups began to use this coastal space
marginally -although purposely- since 2000 years BP. Of all the studied sites in Cabo Virgenes archaeological locality,
CV20 has the highest faunal remains density and high richness of exploited species. Analysis of the bone assemblage shows
emphasis on exploitation of seabirds (Phalacrocorax sp.; Aptenodytes patagonica), in addition to an important presence
of South American sea lions (Otaria flavescens) and carnivores (Pseudalopex culpaeus) with indications of anthropic use.
A high concentration of different taxa of molluscs was recorded, but its human consumption can be sustained only by the
site context. In the regional framework of archaeological records in Cabo Virgenes area, CV20 provides new evidence
about the use of the coast and its resources by hunter-gatherers inland populations at around 800-700 years BP.
as well as pre and postdepositional modifications are assessed. The integration of archaeofaunal analyses with the available technological information for CL3 lithic assemblage suggests an increase in occupation intensity by 4.000 BP, when the highest deposition rate for bones is recorded. Afterwards, by 1.700 BP, archaeofauna shows a shift in patterns of primary prey (Lama guanicoe) exploitation, towards an intensification of its processing. This shift is accompanied by an increment in lithic artifact deposition rate, with assemblages characterized by toolstone intensive-use and an increase in the number of lithic
raw material represented.
KEY WORDS : Patagonia, Holocene sequence, hunter-gatherers, archaeofaunal record, land-use intensity.
en los esquemas de análisis de diferenciación morfológica de las poblaciones modernas de las especies de camélidos sudamericanos. Como caso de estudio se evalúa, a partir de dos pruebas, la técnica de R. Meadows de identificación de taxa arqueológicos a partir de referentes modernos: a) con un camélido indeterminado (AMNH 14121) seleccionando cuatro estándares actuales (dos llamas y dos guanacos), y b) sobre una población representativa de guanacos modernos de Patagonia
en la que se usaron doce referentes actuales seleccionados dentro del rango intercuartil de la población. Con esta experiencia se observó que la técnica de Meadow resulta dependiente de la variación intraespecífica, lo que reduce su eficiencia como herramienta de identificación taxonómica cuando se utilizan especies con rangos de tamaño semejantes. Se concluye que es indispensable comenzar por entender más acerca de la variación intrapoblacional e intraespecífica para poder conocer la diversidad taxonómica.
que evidencien un dimorfismo sexual marcado entre individuos adultos de guanaco (Lama guanicoe). Los principales estudios que evalúan las diferencias osteológicas entre machos y hembras en esta especie están basados
en análisis de variables morfométricas craneanas y en caracteres morfológicos de la pelvis. En este trabajo se evalúa la existencia y magnitud de variables osteométricas poscraneales de diferenciación sexual en una población moderna de guanacos procedente de la localidad Cinco Chañares, Provincia de Río Negro, Argentina. Se estudiaron 32 individuos (8 machos y 24 hembras) de más de 30 meses en los que se midieron 51 hemipelvis, 40 húmeros, 39 radioulnas, 40 fémures y 41 tibias. Los resultados mostraron que las principales diferencias osteológicas, vinculadas con el dimorfismo sexual, se relacionan con variaciones en el tamaño y la forma de la pelvis. En los huesos largos, las diferencias osteométricas significativas se registraron sólo en variables aisladas y no se pudieron definir diferencias osteométricas asociadas con el dimorfismo sexual. Igualmente se considera que esto deberá ser reevaluado con mayor profundidad considerando una muestra mayor de individuos machos.
de variables morfológicas de variación continua. Los huesos analizados son los metacarpos distales y proximales y las segundas falanges. En el análisis se utilizaron muestras procedentes de cuatro sitios arqueológicos y uno paleontológico del extremo austral de Patagonia
continental (Argentina y Chile). Las mismas representan dos momentos cronológicos: Pleistoceno final (ca. 12000-10000 años AP.) y momentos posteriores al Holoceno temprano y
Holoceno medio (ca. 8000-6000 y 6000-4000 años AP.).
Los datos fueron evaluados a partir del análisis de varianza para inspeccionar las diferencias estadísticas entre los tamaños de los huesos. Los resultados mostraron huesos más grandes durante el Pleistoceno final y tamaños significativamente menores durante el Holoceno temprano/
medio.
A su vez, se estudió la relación existente entre los tamaños de los huesos de guanaco y la variable climática asociada con la temperatura, la cual se considera una de las potenciales causas de variación morfológica entre los mamíferos. Para esto se utilizaron los valores de dióxido de carbono del núcleo de hielo antártico Domo C. En el análisis conjunto de los patrones morfológicos de tamaño y la variable de temperatura (CO2) se registró una covariación negativa entre el proxy de temperatura y las modificaciones halladas en los tamaños de los huesos postcraneales de guanaco. Estos resultados sugieren que los tamaños fueron estadísticamente más grandes durante períodos de bajas temperaturas y se redujeron concomitantemente con el incremento de
este parámetro climático.
excavated sectors were considered, one corresponding to a burial area located at the bottom of the cave, and the other one corresponding to a test pit placed next to the burial area. Chronologically, two assemblages were identified, the first one corresponding to the early Late Holocene (ca. 3.500 14C years BP), and the second one to the later Late Holocene (ca. 1.700-500 años 14C AP). The obtained results show statistically significant differences in the manifestation of taphonomic variables and in the frequency and diversity of faunal species both, spatially and temporally. In particular, a more intensive use of the cave at around 3.500 14C years BP is inferred, coincidental with a higher representativeness of coastal taxa like mollusk and sea/coastal birds. Such species may have been exploited during logistic movements to the Magellan strait carried out by hunter gatherers from the inland
The focus was on the exploitation of Lama guanicoe and, occasionally, forest resources (Hippocamelus bisulcus), and European fauna (Equus caballus). The evidence reinforces the interpretation of Laguna Condor as logistically, redundantly and intensively exploited place during short periods ca. 200 years AP.