Human predatory behavior and the social implications of communal hunting based on evidence from the TD10.2 bison bone bed at Gran Dolina (Atapuerca, Spain)
Introduction
There is strong evidence for hunting as the regular form of accessing animal carcasses from the early Pleistocene (Bunn, 1981, Bunn, 2001, Bunn and Kroll, 1986, Domínguez-Rodrigo et al., 2005, Domínguez-Rodrigo et al., 2007, Domínguez-Rodrigo et al., 2009a, Domínguez-Rodrigo et al., 2010, Domínguez-Rodrigo et al., 2014, Domínguez-Rodrigo and Barba, 2006, Pickering et al., 2007, Sahnouni et al., 2013). It can be assumed from this that all subsequent hominins had the ability to be effective hunters. In fact, archeological evidence from recent decades has further confirmed the hunting capabilities of Middle Paleolithic/Middle Stone Age (MP/MSA) hominins (Speth and Tchernov, 2001, Yeshurun et al., 2007, Clark and Kandel, 2013, Clark and Speth, 2013, Yravedra and Cobo-Sánchez, 2015) in addition to their aptitudes for the planning, anticipation, coordination, and communication used in hunting (e.g., White et al., 2016). In the words of Stiner, MP hominins “were expert hunters of large game animals wherever they lived” and exhibited great flexibility in the “large game species that they hunted follow[ing] regional variation in animal community composition” (Stiner, 2013:289). This adaptive capacity is reflected in the great variety of strategies employed to obtain prey from megafauna to small game (Thompson, 2010, Thompson and Henshilwood, 2011, Cochard et al., 2012, Yravedra et al., 2012, Yravedra et al., 2014, Smith, 2015). By contrast, there is less information concerning hunting behavior and its social implications during the Lower Paleolithic (LP), especially within the European context, although several sites, such as Schöningen, suggest the existence of complex dynamics (Thieme, 1999, Voormolen, 2008, Conard et al., 2015, Van Kolfschoten et al., 2015).
Among the many strategies for the procurement of prey, communal hunting has been proposed as part of the subsistence repertoire of Mousterian groups at the end of the European MP, especially during Marine Isotope Stage (MIS) 5 (Jaubert et al., 1990, Jaubert et al., 2005, Farizy et al., 1994, Brugal, 1995, Gaudzinski, 1995, Gaudzinski, 1996, Gaudzinski, 2005, Costamagno et al., 2006, Rendu et al., 2009, Rendu et al., 2012, Discamps et al., 2011, White et al., 2016). This strategy has also been proposed for some assemblages of the MSA in South Africa (Klein, 1978, Klein, 1999, Klein and Cruz-Uribe, 1996, Weaver et al., 2011a).
Following Driver (1995), the social organization of hunting parties, the form of predation (number and rate of animals slaughtered), and the technology used (tactics and tools) must be taken into account to identify and classify hunting methods in prehistory. In this work, communal hunting is considered as a technique that requires the participation of several people, including those that usually do not participate in hunting parties, to kill several prey animals in a single event, often seasonally (Driver, 1990, Driver, 1995, Steele and Baker, 1993). In this sense, zooarcheological testing of the remains resulting from this hunting practice provides valuable information concerning cognitive development, social integration, cooperation among group members, and other aspects of behavior beyond the strictly economical.
Ethnographic data indicate that communal hunting occurs for different economic, social, cultural, and symbolic reasons (Forbis, 1978, Speth, 1983, Speth, 1997, Speth, 2013, Driver, 1995). When the goal is to obtain a large quantity of meat to store (Binford, 1978, Driver, 1990) or be consumed in a place of aggregation (Frison and Todd, 1987), a communal hunt (1) exhibits large numbers of slaughtered individuals of the same species (Driver, 1995, Speth, 1997, Frison, 2004, Lubinski, 2013), (2) presents catastrophic mortality profiles, usually with a marked seasonality in deaths (Frison and Reher, 1970, Reher, 1970), and (3) displays a pattern of selective exploitation of carcasses and the systematic transportation of elements of greater nutritional value to the camps (David and Enloe, 1993, Costamagno, 1999).
The deep knowledge of environments, prey behavior, and seasonal biological cycles of the prey, necessary to perform successful communal kills, is strongly linked with anticipation capacity, social complexity, and the development of cognitive tools, such as articulated language, that are not fully recognized in Neanderthals and their relatives (Binford, 1982, Binford, 1989, Straus, 1997, Morin, 2004). In fact, traditionally it has been thought that communal hunting was exclusively a modern human behavior that was developed during the Upper Paleolithic as part of the “human revolution” (Binford, 1982, Binford, 1985, Binford, 1989, Mellars, 1996, Mellars, 2004). However, as mentioned above, European Mousterian sites associated with convincing evidence of communal hunting are common (Farizy et al., 1994, Brugal, 1995, Grayson and Delpech, 1998, Jaubert et al., 2005, Gaudzinski and Niven, 2009, Niven et al., 2012, Rendu et al., 2012, White et al., 2016), suggesting that the skills and the cognitive capacities for the development of complex hunting techniques of MP hominins were similar to those observed among other “modern” communal hunters.
Some of the technological, anatomical, and behavioral features of MP hominins emerge in transitional moments between the LP and MP (Roberts and Parffit, 1999, Thieme, 1999, White and Ashton, 2003, Hublin, 2009, Stiner et al., 2009, Stiner et al., 2011, Moncel et al., 2011, Moncel et al., 2012, Fontana et al., 2013, Stiner, 2013, Arsuaga et al., 2014). Among those that are linked with social behavior, the Middle Pleistocene hominin record of Sima de los Huesos (Atapuerca, Spain) has offered evidence of conspecific care (Gracia et al., 2009), communicative capacities and possible symbolic behavior (Carbonell and Mosquera, 2006, Martínez et al., 2013, Sala et al., 2015) around 430 ka (Arsuaga et al., 2014). The zooarcheological analysis of the broadly contemporary Gran Dolina TD10.2 sub-unit allows for the evaluation of the economic and social behavior of pre-Neanderthal populations of Atapuerca through the study of a faunal assemblage heavily dominated by a single species of large ungulate. The evidence allows for a discussion of the emergence of communal hunting as a paleoeconomic strategy and its implications for LP social behavior.
Section snippets
Gran Dolina TD10.2
Gran Dolina cave is one of the many karstic formations located in the Sierra de Atapuerca in the northern section of the Iberian Peninsula (Fig. 1a). The cave is of phreatic origin and more than 20 m deep with “keyhole” section morphology. Internal and external deposits fill the cavity, which was discovered in the early 20th century after being cut through during the construction of a railway. Gran Dolina has now collapsed, and upon first glance, far too little remains of the walls and roof in
Materials and methods
During the excavation of the Gran Dolina TD10.2 sub-unit, all faunal remains longer than 2 cm and all identifiable remains (e.g., individual teeth) have been recovered and coordinated in three-dimensional space using a 3-Coor system (Canals, 2008). The excavation protocol at Gran Dolina does not typically piece-plot mesovertebrates (e.g., leporids and small birds). These remains were placed in bone collection bags each day, recording square and depth, together with non-identifiable macromammal
Results
In this work, 24,216 faunal remains (NSP) belonging to a wide variety of taxa, including ungulates, carnivores, large rodents, leporids, birds, and reptiles have been analyzed. Despite this taxonomic diversity, as the name of the bone bed indicates, the assemblage is dominated by bison remains (22,532 or 98.4% NISP) (Table 1). More importantly, in understanding the taxonomic features of the assemblage, the other 17 taxa are represented by less than 60 specimens, each below 0.3% NISP. Only 0.1%
Discussion
The large concentration of archeological remains in the TD10.2 bison bone bed represents a thin, discrete archeostratigraphic layer in which no significant post-depositional processes have occurred. It is an in situ layer where hominins performed tasks related to subsistence, tasks that have been interpreted in this work through examination of the faunal record. In the same stratigraphic context, the upper part of TD10 sequence (named TD10.1) has previously been studied from a taphonomical
Conclusions
The zooarcheological analysis of the faunal assemblage of the Gran Dolina TD10.2 bison bone bed presented here shows that the cave was used as the kill-butchering site for several seasonal events of mass communal hunting in which herds of bison were slaughtered and exploited intensively by the hominins that occupied the cave. The main contribution of this research is the convincing demonstration that humans at Atapuerca were communally driving and killing bison at least 400,000 years ago.
Acknowledgments
We want to express our gratitude to our colleagues in the Atapuerca Research Team, especially those involved in the excavations at Gran Dolina for their hard work. This paper has benefited from discussions held with many of them and with our colleagues Charles Egeland, Manuel Domínguez-Rodrigo, Philippe Fosse, José Yravedra, Anne-Marie Moigne, María Soto, Asier Gómez-Olivencia, Jennifer Parkinson, Eric Delson, Britt Starkovich and Nick Conard. We thank, in particular, John Speth. The exchange
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