Neanderthal foraging in freshwater ecosystems: A reappraisal of the Middle Paleolithic archaeological fish record from continental Western Europe

Highlights

• This paper explores the evidence of Neanderthal fishing.

• If such fishing was a casual, opportunistic activity or a systematic practice.

• Review of fish remains from 11 Middle Palaeolithic contextualised freshwater fish deposits.

• Large (i.e. > 1kg) fishes in anthropic deposits and absence of animal digestive traces and gnawing marks.

• Neanderthals could have played a role in the accumulation of part of these remains.

Abstract

The prevalence of large game found in association with Middle Paleolithic tools has traditionally biased our ideas of Neanderthal subsistence practices. Studies document the exploitation of small mammals, birds, and plants by Neanderthals, whereas data on aquatic resources are still scarce and data on fish are almost non-existent. This article presents a review of fish remains from 11 Middle Palaeolithic fish bone assemblages from well contextualized sites in Belgium, France and Spain. It explores the nature of the evidence in order to determine whether Neanderthal fished and if so, whether fishing was a casual, opportunistic activity or a systematic practice. The first issue to address is whether archaeological fish remains at any given site represent human activity or not. Our study tests that assertion while enhancing our understanding of the diversity of food alternatives available to Neanderthals at any given site, and their ability to adapt to them. Methodological protocols include quantification, body mass and length estimations, and, whenever possible, spatial distribution of fish remains, taphonomic analyses and inference of the season of death. This methodology constitutes an analytical protocol to assess the contribution of fish to the human diet during the Paleolithic and set apart human-generated fish deposits from those generated by alternative fish accumulators. The evidence gathered so far points essentially to circumstantial fishing by Neanderthals, and the question must necessarily remain open for the moment. Nevertheless, some of the evidence, in particular the presence of large (>1 kg) fish in anthropogenic deposits and the absence of animal digestive traces and gnawing marks on fish bones in such deposits, seems compelling and suggests that Neanderthals could have played a role in the accumulation of some of these remains.

Introduction

Western Europe constitutes a key area for investigating Neanderthal subsistence practices due to the quantity and quality of sites available for study. The most recent investigations in this region have significantly expanded our knowledge on the chronology (e.g., Richard et al., 2015; Guérin et al., 2017), environmental conditions (e.g., Britton et al., 2011; Moncel et al., 2015; Discamps and Royer, 2017), and subsistence strategies of Neanderthals (e.g., Delagnes and Rendu, 2011; Cortés-Sánchez et al., 2011; 2019; Niven et al., 2012; Daujeard et al., 2012; 2019; Gaudzinski-Windheuser et al., 2018). These data have contributed to establishing a reliable framework for interpreting the profound biological and cultural changes that occurred from Marine Isotopic Stages (MIS) 11–9 to 3 (Moncel et al., 2016). Traditionally, Neanderthals have been considered as big-game hunters (i.e., top-level carnivores; Bocherens, 2009; Gaudzinski-Windheuser and Niven, 2009; Richards and Trinkaus, 2009; Jaouen et al., 2019). However, over the past decade, studies have evidenced a diversified spectrum of subsistence strategies, with a significant contribution of small game (e.g., Blasco and Fernández Peris, 2012; Cochard et al., 2012; Morin et al., 2019). Archaeozoological and archaeobotanical analyses provide evidence of the exploitation of small mammals, birds, shellfish and plants (Laroulandie, 2000; Cortés-Sánchez et al., 2011, 2019; Hardy et al., 2012, 2013; Laroulandie et al., 2016; Estalrrich et al., 2017), yet the role played by aquatic-vertebrate resources in Neanderthal diets has barely been documented, and Neanderthal fishing remains a controversial issue (Le Gall, 1992, 2000; Roselló-Izquierdo and Morales-Muñiz, 2005). Evidence exists that hominins practiced river or lake fishing during the Lower Paleolithic (Braun et al., 2010; Stewart, 1994) and the Middle Paleolithic in Africa, where specialized technologies such as barbed bone points have been developed by ca. 90 000 BP (Brooks et al., 1995; Yellen et al., 1995). However, in Western Europe, there is currently no evidence that the exploitation of aquatic vertebrates was significant before the Upper Paleolithic period, especially the Magdalenian period (Cleyet-Merle, 1990). For most of the Middle Paleolithic, the sea level was lower than it is today, except during MIS 11 and 5e when sea levels were about 6–13 m above present levels (Dutton et al., 2015). For this reason, the vast majority of coastal archaeological sites, where hominins may have left traces of fishing activities, are currently submerged, often at great depths (Cleyet-Merle, 1990). Although a Neanderthal coastal adaptation, including fishing activities, that would have increased their dietary breadth is possible (see Brown et al., 2011; Zilhão et al., 2020), no such evidence has been found for the exploitation of continental freshwater ecosystems. Hence, the study of inland fishing activities, close to rivers or lakes, remains one of the only possibilities for studying Neanderthal fishing.

Middle Paleolithic fish bones are occasionally documented in inland caves close to rivers, often mixed with stone tools and faunal remains. These remains can only be considered as evidence of fishing by humans when other fish predators, in particular cave-dwelling carnivores and raptors, are ruled out as alternative accumulators. At the time of writing, few studies have been published on how to reliably diagnose fish deposits produced by most of these predators and set them apart from those produced by humans. In Europe, most of these papers focus on the Eurasian eagle-owl (Bubo bubo) and the Eurasian otter (Lutra lutra; Nicholson, 1991, 2000; Le Gall, 1999; Russ and Jones, 2011, Guillaud et al., 2017a, 2018).

This paper explores the role that Western European Neanderthals played in accumulating fish remains. The question “were Neanderthals freshwater fishers?” will also aim to determine whether fishing was an opportunistic or a systematic activity for this hominin. The issue has far-reaching consequences since shifts in prey selection by Neanderthals and anatomically modern humans (AMH) are hotly debated issues. Small terrestrial faunas contributed to the Neanderthal diet as a fallback resource and as staples, yet small game became more frequent in the European Upper Paleolithic (UP), and this suggests that a non-overlap of niches between the two hominins might have existed (Stiner, 1994; Patou-Mathis, 2000; Costamagno et al., 2005; Blasco and Fernández-Peris, 2012; Rufá et al., 2016; Balter and Simon, 2009; Hardy et al., 2013; Wiβing et al., 2019). Fish and shellfish also appear in European Middle Paleolithic (MP) sites, but data on the use of aquatic vertebrates by Neanderthals are still scarce and aquatic faunas are rarely taken into account in studies of Neanderthal consumption patterns (Stringer et al., 2008; Cortés-Sánchez et al., 2011). This scarcity may simply reflect a time-mediated taphonomic bias; fish bones are unlikely to be preserved in archaeological archives because their chemical composition makes them much more susceptible to degradation (Szpak, 2011). This would explain why the earliest exploitation of marine mollusks by Neanderthals, dated to 166 ka cal BP (Bajondillo Cave, Málaga, Spain), is in keeping with that documented for AMH at Pinnacle Point in South Africa (Marean et al., 2007; Cortés-Sánchez et al., 2011, 2019), or on the Atlantic coast of North Africa (Campmas et al., 2016; Chakroun et al., 2017). Later MP deposits from Vanguard (ca. 41.8 ka cal BP) and Gorham’s Cave (ca. 30–32 ka cal BP) in Gibraltar document dolphins, seals and fish, in addition to marine mollusks (Stringer et al., 2008). Additional lines of evidence for the exploitation of aquatic resources by Neanderthals include residue analyses on stone tools, tooth use wear, and anthropological studies (Puech and Puech, 1993; Hardy et al., 2013; Trinkaus et al., 2019). Another potential line of inquiry is isotopic analyses. No isotopic studies have yet focused on the consumption of aquatic resources by Neanderthals, perhaps because if these did not make up a significant fraction of their diet, they may not be discernible using current analytical procedures (Patou-Mathis, 2000; Drucker and Bocherens, 2004; Drucker et al., 2005; Stringer et al., 2008). For this reason, the development of sulfur or zinc isotope analyses seems promising for directly assessing fish consumption on Neanderthal bones (Privat et al., 2007; Nehlich et al., 2010; Jaouen et al., 2018).

Evidence of the use of aquatic resources helps to explain why Neanderthals settled in marine and freshwater areas and lends weight to the proposal that fishing was a well-established Neanderthal practice or an occasional, opportunistic activity. To explore this issue, our paper analyzes and reviews a series of MP fish collections from Belgium, France and Spain that span from MIS 11 to MIS 3 by applying a battery of protocols in order to define a methodology for assessing the contribution of fish to the human diet during prehistoric times.