Being-with other predators: Cultural negotiations of Neanderthal-carnivore relationships in Late Pleistocene Europe of Anthropological Archaeology

Late Pleistocene hominins co-evolved with non-analogue assemblages of carnivores and carnivorous omnivores. Although previous work has carefully examined the ecological and adaptive significance of living in such carnivore-saturated environments, surprisingly little attention has been paid to the social and cultural conse- quences of being-with, and adapting to, other charismatic predators and keystone carnivores. Focusing on Neanderthal populations in Western Eurasia, this paper draws together mounting archaeological evidence that suggests that some Late Pleistocene hominins devised specific behavioral strategies to negotiate their place within the vibrant carnivore guilds of their time. We build on integrative multispecies theory and broader re- conceptualizations of human-nature relations to argue that otherwise puzzling evidence for purported ‘symbolic ’ behavior among Neanderthals can compellingly be re-synthesized with their ecology, settlement organi- zation and lifeworld phenomenology. This re-framing of Neanderthal lifeways in the larger context of startling carnivore environments reveals that these hominins likely developed intimate, culturally mediated, and hence varied, bonds with raptor, hyena and bear others, rather than merely competing with them for resources, space and survival. This redressing of human-carnivore relations in the Middle Paleolithic yields important challenges for current narratives on evolving multispecies systems in the Late Pleistocene, complicating our understanding of Late Quaternary megafaunal extinctions and the roles of hominins in these processes.


Introduction
Both historically and cross-culturally, relationships between humans and other carnivores are often contentious, ambivalent and tensionridden affairs (Kellert et al. 1996;Campmans 2017;Lamb et al. 2020), yet they similarly rank among the most culturally vigorous and materially generative human-animal interfaces documented by ecologists (Monsarrat and Kerley 2018;Sommerville 2021;Clark et al. 2021), historians (Gade 2006;Zehnle 2015;Widlok 2018), anthropologists (Barsh and Marlor 2003;Rival 2012; Baynes-Rock 2015a) and archaeologists alike (Hahn 1986;Mithen 1999;Clottes and Azéma 2005;Germonpré and Hämäläinen 2007;Betts et al. 2015;Poole 2015;Porr 2015;Hussain and Floss 2015;Costamagno et al. 2018a). Carnivore assemblages and ecological communities of the deep past were extremely diverse and probably considerably enriched in terms of species count and biomass contribution when compared to modern-day carnivore-bearing environments (Turner 1990(Turner , 1995; Van Valkenburgh et al. 2016;Galetti et al. 2018;Faith et al. 2019;Stanton et al. 2020). In many parts of the world including Europe, carnivore assemblages were strongly impacted by Late Quaternary extinction trajectories (Stuart and Lister 2007;Malhi et al. 2016;Galetti et al. 2018), putting a final nail to the coffin of most large-bodied carnivores and socalled mega-carnivores that are a hallmark of Pleistocene animal ecologies (Faith et al. 2019). The Late Pleistocene in particular has seen substantial turnover in carnivore community dynamics and composition and documents the extinction of some notable carnivore others (Turner 1985), such as various hyena species with whom hominins in Europe and elsewhere have co-evolved and shared the environment for many millennia (Turner et al. 2008). Some scholars have argued that Eurasian Neanderthals in particular are to be understood as an integral part of situated carnivore ecologies and their associated predator-prey interlinkages, whose gradual disappearance in the second part of the Late Pleistocene may have consequently precipitated the eventual demise of the Neanderthal phenotype at the end of MIS 3 (Estévez 2004;Stewart 2005Stewart , 2007. Previous work has tried to address the importance of Pleistocene hominin-carnivore interactions from the perspective of subsistence behavior (Kindler 2012;Wojtal et al. 2020), competitive exclusion/ niche partitioning (Stiner 2002(Stiner , 2004(Stiner , 2012Dusseldorp 2011Dusseldorp , 2013Enloe 2012;Churchill 2014;Orbach and Yeshurun 2019), and energetics or interspecies conflict (Shipman 1984;Binford et al. 1988;Blumenshine 1987;Selvaggio 1994;Kruuk 2002;Turner et al. 2013;Camarós et al. 2016;Domínguez-Rodrigo and Pickering 2017), mostly within the framework of 'hard' ecology approaches, including Human Behavioral Ecology and generalized Neo-Darwinian evolutionary theory (see also García and Virgós 2007). Even though some of this scholarship acknowledges the potential social significance of the carnivore environment for Neanderthals and other Late Pleistocene hominins (e.g., Wojtal et al. 2020), there is hardly any specific, let alone substantial, engagement with the cultural and material consequences of coinhabiting environments with 'charismatic' carnivore others (sensu Lorimer 2007;Ducarme et al. 2013), for example in terms of interspecies sympatry (but see Brantingham 1998; White and Pettitt 2011: 51;Jimenez et al. 2021), landscape learning and sharing, home-making/ neighboring, attuned cognition and perception, or extended, 'morethan-human' kinshipcommon axes of investigation in multispecies studies. Foregoing scholarship on hominin-carnivore relations, in other words, has overwhelmingly focused on questions of survival, niche coordination and environmental adaptation, largely ignoring, or glossing over, the phenomenological implications of living-with and engagingwith powerful carnivore others. At least in part, the focus on die-hard issues of subsistence, and especially competition with other carnivores, remains a living legacy of Binford's (1989) New Archaeology-founding critique on Neanderthal hunting prowess and the ensuing highly influential 'scavenging debate ' (cf. Uthmeier 2006: 19), whilst ecologists increasingly stress the importance of apex predators in promoting biodiversity and species coexistence (Wallach et al. 2015). Recent years have seen a resurge of archaeological attention to carnivores, especially in the Late Pleistocene, due to a series of new discoveries revealing unusual hominin behaviors in relation to various carnivore taxa, both mammals and birds, including but not limited to the special treatment of whole skeletons or some selected body parts (e.g., Peresani et al. 2011;Morin and Laroulandie 2012;Walker et al. 2012;Romandini et al. 2014;Radovčic et al. 2015;Sanchis et al. 2015;Cueto et al. 2016;Sanz et al. 2017;Hussain 2019;Rodríguez-Hidalgo et al. 2019;Wojtal et al. 2020;Val et al. 2020;Hallett et al. 2021). This accumulating deep-time evidence clearly suggests that carnivores were more than mere evolutionary antagonists and may have contributed significantly, and in diverse contingent ways, to the human story over many millennia. Relatedly, Jimenez et al. (2021) have recently suggested to revisit the predatory sympatry of hominins and hyenas in Late Pleistocene Eurasia by rejoining palaeontology and archaeology and paying closer attention to how changing behaviors of key predators 'orchestrated' the behavior of their interlocutorsboth prey and competitorand in this way modulated broader co-evolutionary dynamics on ecosystem scales.
The present paper joins this broader call but proposes that a conceptual reframing of the study of Pleistocene hominin-carnivore relationships is required in order to fully explore the vital importance of nonhuman carnivores in the broader context of early human biocultural evolution. Carnivores need to come into view not only as evolutionary competitors, but equally so as co-constituents, perhaps even protagonists (Hussain in press) of the phenomenal and cognitive worlds in which past hominins operated, so that precarious hominin-carnivore communities and more-or-less stable neighborhoods can be investigated, detailed and contrasted from the perspective of integrated ecocultural analysis. This opening up of the conceptual space for examining hominin-carnivore intersections and their situated variability in deep prehistory provides new possibilities for integrating multispecies theory and the achievements of the widely celebrated 'animal turn' across the humanities and social sciences (Ritvo 2007;Haraway 2007;Kirksey and Helmreich 2010;Wolfe 2011;DeMello 2012;Ogden et al. 2013;Grusin 2015;Kalof 2017;Pilaar Birch 2018a). Combining these bodies of theory with more traditional palaeoecological and archaeological analysis and insight, whilst nurturing knowledge from general forager anthropology, discloses fresh perspectives on the interdigitation of hominin and animal lives in the Pleistocene past, and the centrality of changing hominincarnivore interstices in the evolution of hominin lifeways and ecocultural adaptations.
The paper first outlines some basic multispecies concepts to facilitate the study of human-carnivore interactions, in particular its generative cultural and material dimensions. We then turn to the archaeology of Marine Isotope Stage (MIS) 5-3 Neanderthals in different parts of Europe as a case in point and explore their complex ecocultural entanglements with raptors, hyenas and bears. This exploration indicates that at least some instances of Neanderthal material culture previously interpreted as 'symbolic' can be framed as 'emergent materialities' of cohabitation and persistent friction between hominins and significant carnivore others in the landscape. We map these instances vis-à-vis situated interspecies immediacies following the seminal work of Bird-David (1990, 2017a. Finally, we examine the broader implications of this redressing of hominin-carnivore relationships, especially with regard to understandings of 'nature' in the deep past and problematic yet persisting narratives of accumulating hominin 'modernity' during the Late Pleistocene (cf., Nowell 2013;Ames et al. 2013;Porr and Matthews 2017;Kissel and Fuentes 2020). As an alternative reading 'against the grain', we suggest that much of the palaeoarchaeological evidence routinely mustered to bolster modernity narratives and related views in the field may in fact express diverse hominin-world engagements, i.e., the shifting roles of different nonhumans, including carnivore others, in various hominin affairs and the evolving modes and efforts of distributed multispecies 'world-making' (cf. e.g., Goodman 1978;Anderson and Harrison 2010;Descola 2010aDescola , 2014Stewart 2010). We propose there is a large, yet hitherto uncharted field of investigation where evolved carnivore legacies can fruitfully be exposed by investigating the coevolutionary matrix of human-carnivore interactionand its consequences for and sociomaterial imports to multiple trajectories of becoming human.

Multispecies theory
At the core of multispecies thinking and cognate bodies of theorizing is a growing epistemological unease with the Western legacy of centering the human species in cultural and evolutionary processes (e.g., Kohn 2013;Nance 2015;Locke and Münster 2015;Boyd 2017;Armstrong Oma 2018;Swanson et al. 2018). Critiqued are especially those variants of anthropocentric analysis in which radical domain autonomy is assumed and explicitly defended (Livingston and Puar 2011), to the extent that human affairs are portrayed and explained in complete isolation from the so-called phenomena of 'nature' (Anderson and Perrin 2018). Notwithstanding, multispecies theorists are equally adamant in their fundamental critique of undifferentiated, homogenizing and overexpansive views of nature and what is then symptomatically framed simply as the 'external' environment (Descola and Pálsson 1996;Descola 2005), more often than not falling back into naïve ecological determinisms or perpetuating global or even universal heteronomy presumptions. Yet humans, their behaviors, cultures and adaptations can hardly be analyzed or understood in separation from the diverse natures in which they partake (Hastrup 2014). To the contrary, human behaviors, material cultures and sociocultural horizons are always enfolded in, and to this effect partly a consequence of, variegated hybrid assemblages of humans and nonhumans and their situated internal dynamics, feedbacks, short and long-term histories, path dependencies and relational logics (Descola 2010b;Arluke and Sanders 2008).
Following Haraway (1991Haraway ( , 2016, the 'multispecies condition' of human life on Earth eclipses equalizing tendencies of 'naturalization' precisely because humans do not evolve in contradistinction but in connection and community with the heterogeneous repertoire of nonhumans, including other animals, with whom they share their surroundings (cf. Lestel and Hollis 2013;Brittain and Overton 2013;Armstrong Oma and Goldhahn 2020). Due to this foundational antiessentialist inclination, multispecies thinking promotes, and often joins forces with, relational approaches to human-nonhuman configurations in the past and their often far-reaching behavioral and sociocultural consequences (Watts 2013a;Hill 2013; Harrison-Buck and Hendon 2018) -sometimes flagged as the 'relational view' in archaeology (Herva and Lahelma 2020;Wisher and Kay 2020). What Shipman (2010) has termed the 'animal connection' in human evolution is then much more than mere addendum to a list of evolved, supposedly quintessential human traits. Rather, as Tsing (2012) has poignantly noted, 'being human [, then,] is an interspecies relationship' (emphasis added).
In what follows, we delineate a number of key concepts derived from this expanding corpus of multispecies theory in archaeology, anthropology and relevant neighboring fields, especially from what is now routinely labeled 'animal studies', 'anthrozoology', 'multispecies ethnography' and, increasingly, 'multispecies archaeology'. This exercise is necessary to prepare the stage for our subsequent attempt of redressing hominin-carnivore relationships in human deep prehistory.

Coevolution
The multispecies critique has important ramifications for how we think about human evolution and what place we secure for humananimal relationships in evolutionary dynamics. The agents of nonhuman nature, especially sentient animal others, draw attention not only as fuel, resources and raw materials of evolution or as fierce competitors, they come into view as potent and consequential media of evolution, altering the conditions and modes of evolutionary dynamics themselves (Hussain in press). Animals are not merely acted upon by humans or other towering, anonymous evolutionary forces (Naiman 1988), they can emerge as direction-giving protagonists within such processes (e.g., Jones et al. 1997). The often-underrated agency of animal others (pace Sultan 2015; Walsh 2015) comes for example to the fore when animals enter co-evolutionary relationships with hominins and play a role in fashioning local environments or creating behavioral opportunities and affordances for their hominin neighbors (e.g., Tchernov 1984;O'Connor 2013;Pierotti and Fogg 2017;Kost and Hussain 2019;Hjørungdal 2019). Hominins may then adapt to particular animal ecologies or specific animal species and the same may occur vice versa: nonhuman animals attend and adapt to particular environments, behaviors and behavioral possibilities catered by hominins (Haynes 2006;Clucas and Marzluff 2011;Boivin et al. 2016;Bocherens 2018). When both conditions are met, we can speak of reciprocal coevolutionor mutual 'facilitation' (Bruno et al. 2003) -which frequently produces intimate multispecies entanglements molding the long-time histories of those involved (cf., Fuentes and Wolfe 2002;Albarella et al. 2007;Sykes et al. 2020). Some of these strands of cobecoming can be framed as processes of 'interspecies niche construction' (Candea 2010;Fuentes 2010) in which humans and nonhumans modify and redress each other's environments, and in this way alter the ecology-behavior nexus in larger, co-evolving multispecies communities (Pilaar Birch 2018b; Kost and Hussain 2019).
Importantly, such intimate co-evolutionary interlacings of hominins and other animals are premised to result in tangible and often unique cultural consequences, affecting for example how hominins move in their landscapes and organize their settlement systems, how environments are materially and cognitively appropriated, how material culture and hominin technology mediate encounters with various nonhuman others, and how hominins perceive, emplace, enstory and conceptualize the worlds they inhabit. Co-evolutionary linkages with animal others have thus the potential to reconfigure the totality of hominin behavior and cognition and this should be reflected in the various ways Pleistocene hominins have made themselves at home in specific animal landscapes and ecologies. Haraway (2016: 58) reminds us that no living beings fully make themselves, and the same principle applies to human evolutionary processesstipulated as 'autopoietic' or self-making, human evolution becomes a hollow, tractionless concept. Childe's (1936) credo 'man makes himself' is thus at best misleading insofar as the human story hardly marks out a self-sufficient development, but rather bespeaks of complicated, distributed, multi-responsive, co-adaptive and situated trajectories, in which heterogeneous forces and agents, including hominins and other animals, engage with one another and probe into the emerging possibilities of co-existence and 'being-with'. This constant tangling and cross-pollination of variegated evolutionary subjects that, in various ways and to varying extent, make each other is captured by Haraway's (2007) now classic formulation of the 'humanimal' -the inescapable and historically significant co-making of hominins and animals, arguably exploration-prone especially in the thickets of deep time (sensu Shryock et al. 2011).
In this formulation, evolution is fundamentally a 'sympoietic' process (Haraway 2016) -change is predicated on 'making-with' -and sympoiesis becomes a key mechanism and driving force of hominin evolution writ large. Becoming or evolving is then almost always a process of 'becoming-with' (cf., Kirksey and Helmreich 2010: 546), so that the creation of 'mutual ecologies' (Fuentes 2010) and the articulation of hybrid 'contact zones' (sensu Pratt 1991Pratt , 2008cf., Aisher and Damodaran 2016;Isaacs and Otruba 2019) in hominin evolution become privileged loci of investigation, drawing attention to the delicate coordination of multiple lifeworlds and livelihoods that steer the directionalities of long-term change. Coevolution thus tables a range of useful concepts to incorporate more-than-human thinking into emerging contemporary approaches to human evolutionfor example in the spirit of Fuentes' (2015) call for an 'integrative anthropology of the human niche'. In general, sympoiesis can be especially powerful, and may even become an overwriting process, when humans and other animals encounter each other as key ingredients and perceptual touchstones of their respective lifeworlds, and hence when animal others emerge as 'companion species' (sensu Haraway 2007). These salient animals are prone to co-configure the emotional, psychological and empathic horizons of their hominin interlocutors (cf., Hussain and Breyer 2017) and to foster particular emotive responsese.g., respect, awe, curiosity or fear that derive from the ethology of the animals in question and the nature of specific human-animal engagements, and in this way become paramount in shaping the experiential regimes and emotive topologies of the lifeworlds they share and co-make with hominins.

Interspecies immediacy and proximity
Over the last decades, animal studies and anthrozoology have begun to describe and map out variegated forms of interspecies relationship that arguably are human nature, and try to reframe archaeologists' and anthropologists' traditional concern with animals as 'good to use', 'eat' (Harris 1985) or 'think with' (Sax 2001) in favor of the historically situated conditions of shared multispecies life (Haraway 2007;Kirksey and Helmreich 2010: 552;Lestel and Hollis 2013;Brittain and Overton 2013;Locke 2014;Hussain 2019, in press). This emphasis on shifting constellations of living together opens up an entire set of animal-related research questions: e.g., how do hominins and animals overlap (or not) in geographic space? When and how do their behaviors intersect? What is the nature of their encounter and is the latter framed by particular places or mediated by specific materials and technologies? What are the cultural and biological rhythms that hominins and other animals attend to, and how are these related? What are the affordances provided by the respective other? How salient and regular are encounters and interactions between the involved agents? What cultural and material resources are activated and what strategies are deployed (if any) to ensure auspicious coordination, co-habitation and mutual respect? The relationship between 'living-with' and 'becoming-with' hereby recapitulates the intersection between ecology and evolution in biology, but the accent is put on different ways of relating and engaging at shifting multispecies interfaces, in turn born out of the dynamics generated by historically-specific assemblages containing hominins, other animals and larger biophyscial landscapes. Such multispecies interfaces are never entirely stable but mutable and open to reorganization (Livingston and Puar 2011). Their characterization thus requires careful analysis of how different life-processes and behaviors interconnect and resonate with one another, and what the specific ecological, sociocultural and material consequences as well as creative potentialities are.
This life-oriented rendering of hominin-animal interaction draws attention to the topologies of interspecies immediacy (cf., Locke 2014; Hussain and Breyer 2017;Bird-David 2017a) and motivates their analysis and comparison. Hominins never lived in exclusionary enclaves of conspecifics, but had regular and often intimate contact with nonhuman others when operating in their environments (cf., Hussain 2019). There is a non-arbitrary relationship between the significance and nature of these contacts and the cultural and social technologies devised to respond to, curate, and process them. A brief ethnographic intervention is required to elaborate this point. As Bird-David (1994, 2017a, 2017b and others (e.g., Meillassoux 1973;Price 1975) have lucidly unpacked in a number of writings, many small-scale foraging peopleespecially those attuned to a 'band-like' form of social organizationpursue their livelihoods in a general condition of immediacy. Despite the pronounced diversity of hunter-gatherer lifestyles, forms of social organization and intracommunity relations especially in the past (cf., Finlayson and Warren 2017), there is often little spatial and temporal segregation of the social world, i.e., hardly any boundaries of interaction and communication (Bird-David 1994: 591). Moreover, many smallscale societies tend to resist sociodemographic upscaling (cf. esp., Bird-David 2017band their 'tiny-scale' is not just a matter of numbers and effective population sizes or demography structures, yet is linked to the cultivation of culturally-specific scaling practices, or the premediated lack thereof (cf., Fowles 2018), which in turn feed into regulative ideas of world-making and sociality, fundamentally regulating how these foragers perceive and attend to their world (Bird-David 2017b). The small size of these communities, often bound to an ethos of 'total sharing', has particular consequences for their modes of beingwith conspecifics but also with nonhuman others with whom the local environment is co-inhabited and shared. Small-scale societies, especially those adhering to 'immediate return' modes of production (Woodburn 1980(Woodburn , 1982, thus tend to inhabit intricate 'nanoscale' worlds (Bird-David 2017a) where social and ecological distance is considerably reduced and 'immediacy' often actively sought-for, created and behaviorally facilitated. Forager life in these socioeconomic contexts is characterized by an intrinsic quality of co-presence (pace Widlok 2020) -or by what Bird-David (2017a has called 'pluripresence' -and this quality readily extends beyond the limits of the human, also encompassing multispecies relationships and understandings. Conditions of pluripresence, which are commonly tied to closeknitted and non-exclusionary communities of practice, interaction and routine encounter, often with deep knowledge of local ecologies and heightened levels of social transparency, easily give way to expansive 'we understandings' and 'designations' (Bird-David 2017b). These can accommodate a significant number of nonhuman members, especially if they constitute phenomenological cornerstones and key agencies of the immediate environments in questioni.e., animal others who are frequently witnessed, recurrently encountered and those who typify or markedly shape these local surroundings. The commonality, and in some cases near-banality, of their being-with humans creates a sense of sharing, belonging and tangible companionship that goes hand in hand with sociocultural adjacency and proximity. As Bird-David (2017a, 2017b points out, the central trope and experience in these contexts is phenomenological diversity and not homogenization, and such basal diversity crosscuts the Western human-animal boundary. The key point is that nonhuman others are capable of entering the wider social field of these forager societies (Hussain 2013;Langley 2013;Hussain and Breyer 2017: 216-218) and 'society' can become a near-synonym for the social bonds that permeate human-nature systems as a whole (Descola , 2005Bird-David 1999;2017a;Bird-David and Naveh 2008;cf., Bawaka Country et al. 2015). Just like other members of society, socially recognized nonhumans co-configure the specific 'horizons of concern' (Bird-David 2017b) that anchor the everyday activities and occupations of their forager neighbors.
Life in phenomenologically 'small worlds' therefore lays the foundations for real and 'imagined' multispecies communities (pace Anderson 1983) and can afford more-than-human belonging and kinship (pace Haraway 2016), even if the modes of 'being many' might differ across time and space. Under conditions of pluripresence, there is reduced social pressure to address the already-present, yet multispecies neighborhoods are nonetheless culturally cherished and often require material and cultural negotiation and coordination, especially if lived or experienced as precarious. These consequential more-than-human configurations can for example be examined by confronting human and nonhuman geographies (Bennett 1960;Wolch and Emel 1998;Ogden et al. 2013;Baynes-Rock 2013;Rutherford and Wilcox 2018), affective multispecies landscapes and places (Ogden 2011;Overton and Hamilakis 2013;Lorimer et al. 2019) and the distribution of more and less mobile elements such as artefacts in them, including their fragmentation, circulation and deposition. Some tiny-scale forager societies also seem to engage in behaviors that register as 'distance-reduction' strategies, and hence increase proximity between humans and nonhumans, sometimes even extending to the inanimate, for instance when stones and other 'natural' objects are brought to occupational spaces in order to 'live with' and accompany people (Bird-David and Naveh 2008;Bird-David 2018: 312).
Indigenous thinking similarly highlights the pluralistic nature of society-making based on the 'circulation of agency through human and nonhuman worlds' (Watts 2013b) as well as the maintenance and careful curation of foundational social relationships with other-thanhuman entities such as animals (TallBear 2011;Todd 2014;Musharbash 2017;Maynard 2022). What counts as 'society' not only depends on human practice and thought then, but also critically relies on how both are enfolded in larger multispecies landscapes populated by stones, rivers, plants and other-than-human animals (Watts 2013b). This agential 'heterotopology' of space is anchored in lived experience of cobeing and interspecies proximity and directly bespeaks of the mutable architecture of situated multispecies lifeworlds, hence emerging as a precondition rather than a consequence of significations of 'place' and how the land ('Country' in many Indigenous conceptions, cf. Bawka Country et al. 2015) is navigated, storied and enculturated (Todd 2017). Recognizing society as a species-transcending process of multi-agential polyphonic participation, in which human-animal boundaries are encated as fluid and continuously re-negotiated, rather than cast as categorically invariant, is part of broader decolonization efforts in anthropology and archaeology and as such tables new resources to develop and probe into alternative ways of exploring and understanding the Pleistocene past (Steeves 2021).
Indigenous thinking further draws attention to the structural connection between human-nonhuman co-becoming, interspecies coordination and the importance of cultivating attentiveness and cultures of respect, care and responsibility in the face of nonhuman others (Bawaka Country et al. 2015). Indigenous perspectives highlight the centrality of 'more-than-human geographies of care' that begin rather than end with a 'social ontology of connection, foregrounding social relationships of mutuality and trust' (Lawson 2007: 3). This is not to say that these relationships are always bound to be harmonious or positively sanctioned, to the contrary, it emphasizes their porosity and the resulting need to continuously belabor them and their success for the sake of physical, social, emotional and cosmological sustenance. Connection, respect and co-responsibility are not abstract theoretical notions, then, but are retrieved from lived experience and situated, embodied interactions in the concrete context of a lifeworld stained in proximity, where humans and nonhumans inexorably 'share[] time, space, resources, and experiences of both good times and the bad' (Musharbash 2017: 109, emphasis added).

Naturecultures
Multispecies considerations collapse the fundamental distinction between nature and society, and thus undermine the tenets of dualist theory (Haraway 2007(Haraway , 2016Kirksey and Helmreich 2010;Baynes-Rock 2013. Multispecies being-with and becoming-with cannot properly be elucidated when adhering to an analytical separation between ecological and sociological orders, such that the 'former is primarily an extension of the order found everywhere in nature, whereas the latter is exclusively, or at least almost, a distinctly human phenomenon', often with an explicit coupling of ecology and competition vs. social organization and communication (Hollingshead 1940: 358). This classic Western dualist view of human-nature relationsinscribing either into 'orientalist' or 'paternalist' paradigms (sensu Pálsson et al. 1996) can be contrasted with a paradigm of 'communalism' emphasizing that notions of 'nature' and 'society' are at best products of or nodal points within a continuous, dialogical process of co-making among heterogeneous actors (Pálsson 1996: 72;cf. Descola 1994cf. Descola , 1996. This focus on dialogue not only enables to take into account that hunting-gathering people and animal others are known to engage in the 'mutual production of each other['s realities]' (Tapper 1988: 52), resisting the 'Othering' of nature so central to the Modernist project (Pálsson 1996: 64), but also, and perhaps more importantly, to recognize the possible alterity and non-analogicity of human-nature constellations in comparative deep-time perspective, while remaining inclusive and sensitive to Indigenous knowing.
The rejection of orientalism and paternalism and their problematic discontinuous framing of the human-nature nexus in favor of communalism centered on human-nature continuity as well as contingent, situated and participatory forms of generalized, although often nonbalanced reciprocity (cf., Pálsson 1996: 66-67, 72) provides a productive vision of variously textured 'naturecultures' (sensu Haraway 2003: 1-5; cf., Potter and Hawkins 2009; Malone and Ovenden 2017) available for interdisciplinary investigation and confrontation. The material residues, artefacts and signatures recovered by archaeologists are then, on a basic level, to be considered first and foremost 'emergent materialities' of such shifting nature-culture interlacings, rather than 'expressing' or 'representing' mysteriously pre-assembled natural or cultural capacities; they are both the means and outcomes of changing articulations of humans and nonhumans and their mutable modes of attending, comaking and sympatry. Distinct naturecultures and their specific sociomaterial productions and generative relations are likely bound up with different 'styles of worlding' (Descola 2014), and as such help to illuminate, or at least draw near, the ontological contours of past hominin lifeways in unique multispecies worlds.

The sympatric Neanderthal
In an early review of human-animal interactions in hominin prehistory, Mithen (1999: 198) concluded that 'Neanderthals did not develop any relationships with animals beyond those of predator/prey as had characterized human ancestors for the previous million years', and that 'the role animals played in Neanderthal society was [therefore] limited'. More recently, Shipman (2010Shipman ( , 2015a has suggested that the establishment of consequential and permanent social ties with animal others, primarily through early forms of domestication or self/autodomestication, remains a milestone achievement of Homo sapiens populations, and thus possibly contributed to the competitive disadvantage of Neanderthals and other Late Pleistocene hominins in Europe and elsewhere (esp. Shipman 2015b). Mithen (1996Mithen ( , 2013 has further weaved his view into a 'cathedral model' of the mind, explicitly denying Neanderthals any domain fluidity and thereby reinforcing problematic modernist nature-culture polarities. In a similar spirit, Langley (2013), albeit not explicitly discussing Middle Paleolithic human-animal interfaces, has suggested that Neanderthals had a limited capacity to 'socialize' their landscapes, and developed material culture such as personal ornaments primarily to mediate interpersonal relationships rather than to negotiate hominin-animal-place linkages. Symptomatically, animal historians also tend to start their exploration of animal contributions to human history with opulent H. sapiens cave art and the manipulation, management and domestication of nonhuman animals at the Pleistocene-Holocene threshold (e.g., Kalof 2007).
We suggest that this received view reflects a fundamental misunderstanding of Neanderthal society promoted by a narrow discourse on hominin modernity and so-called symbolic capacities plaguing the archaeological expert literature for many years now. Neanderthals and other Late Pleistocene hominins were deeply attuned to their local animal ecologies and developed varied forms of interspecies sympatry with so far overlooked consequences for hominin life and materiality. Rethinking Neanderthal-nature relations in the light of multispecies theory requires to take seriously that animal ecologies and interspecies relations may have been basic cultural resources for these hominins. This necessiates moving past binary disciplinary discourses, which continuously re-cast Neanderthals in terms of a 'us' vs. 'them' logic (Speth 2004), either overemphasizing difference or similarity (cf. e.g., Villa and Roebroeks 2014). This conceptual deadlock and the resulting imaginaries of Neanderthal behavior have trickled down to and fundamentally biased the study of Middle Paleolithic hominin-animal interactions, which continue to be examined almost exclusively in ecological and economic terms (Priestley 2020). What is more, recovering the purportedly 'symbolic' dimensions of Neanderthal behavior (e. g., Finlayson 2019) may only worsen the situation since symbolism is rarely unpacked and placed in its deep-historical context, but more often than not serves as a privileged lighthouse of affirming hominin 'capacities' and cognitive dispositions. A phenomenological approach helps to lay bare and explore the specificity and historicity of Neanderthal engagements with the nonhuman world without retreating to normative juxtapositions of Middle and Upper Paleolithic hominins.
Phenomenology also matters when the social dimension and immediacy of Neanderthal being-in-the-world is taken into account. Neanderthal societies were tiny-scaleas indicated by the scattered nature of their material traces (e.g., Jöris 2014Jöris : 1724Jöris -1726Locht et al. 2016), the size and palimpsest character of their sites, living spaces, shelters and settlement structures (Kolen 1999;Uthmeier 2006: 25;Hayden 2012) as well as the latest palaeodemographic reconstructions and interpretations (Stiner and Kuhn 2006;Bocquet-Appel and Degioanni 2013;Mafessoni and Prüfer 2017;French 2021). Their socioeconomic strategies, furthermore, presently lack substantial evidence for the storage and delayed consumption of food, labor-intensive trapping and catching devices or supports such as nets and boats, the organized manipulation of wild plants and/or animals, and differential access to some of these resources within their communities (pace Woodburn 1982: 432-433). Taken as a whole, this signals their broad operative reliance on 'immediate return' foraging systems (in contrast to 'delayed return' systems; sensu Woodburn 1980), albeit likely without direct historical or contemporary analogues, and despite increasing evidence for general niche construction capacities (Roebroeks et al. 2021;Nikulina et al. 2022). Strikingly, recent eco-cultural niche modeling of Middle Paleolithic Western Europe, in particular France, suggests that Neanderthal groups displayed pronounced territorial integrity and region-fidelity (Banks et al. 2021), which, together with their high mobility profile (Jöris 2014), would have amplified their ecological intimacy and exposure to particular animal-ripe 'giving environments' (pace Bird-David 1990; see above).
The archaeological record of Neanderthals in Europe includes evidence for practices of social care (Spikins et al. 2018) and plant-based medical knowledge (Hardy et al. 2012) -and thus potential 'social technologies' of delayed returnbut there is no unambiguous subsistence-related evidence satisfying the original definition of delayed-return foragers (sensu Woodburn 1982;cf., Uthmeier 2006: 24). That said, some researchers have argued that the apparent spoil surplus implied by some Neanderthal kill and processing sites may indicate meat storage and tactics of 'deferred' consumption with low archaeological visibility Speth 2017;Wragg Sykes 2020: 165). Further, Neanderthal stone working and tool-use technology is certainly more complex and presuppositional than often acknowledged (Çep et al. 2021). Middle Paleolithic lithic technology is frequently anchored in coexisting débitage and façonnage systems including more-or-less specialized operational sequences and tool concepts showcasing considerable technical planning-depth as well as operative delayedreturn rationales (e.g., Soriano 2000;Soressi and Hays 2003;Delagnes and Rendu 2011;Turq et al. 2013;Weiss 2020). Yet overall, economic intensification remains to be convincingly demonstrated at least for European Neanderthals (see Speth and Clark 2006 for possible Levantine evidence) and the small number of pit and 'scoop' features known from Middle Paleolithic contexts, for example at the classic site of La Quina in Western France, have mostly been interpreted as non-alimentary structures (cf., Debénath and Jelinek, 1998: 37;Pettitt 2002: 18;but see Speth 2017: 62).
Furthermore, the Western Eurasian Neanderthal record and its human-animal relations are not as homogeneous as they have long been pictured (Mithen 2006: 230-231;de la Torre et al. 2013;Kuhn 2013;Angheliu 2014). There is now increasing evidence for varied material and subsistence practices and sociomaterial entanglements with various animals and ecological assemblages in different spatiotemporal contexts (Stringer et al. 2008;Nowell 2013;Churchill 2014: 185;Roebroeks and Soressi 2016;Shipley and Kindscher 2016;Morin et al. 2016;Villa et al. 2020), pointing to a heterogeneous and bafflingly complex chronogeography of Neanderthal naturecultures and ecocultural regimes (cf., Wragg Sykes 2020 for a similar global argument). This diversity has previously been explained by the ecological flexibility of Neanderthal populations and their ability to successfully adapt to local conditions (Churchill 2014), but a closer look at the intersection of local multispecies assemblages, animal agencies, Neanderthal behaviors and material productions reveals the limitations and lack of context-specificity of such explanations, glossing over the phenomenological ramifications and challenges of Neanderthal life in disparate multispecies arenas (cf., Wragg Sykes 2020: 221-225). Similar concerns have led Chang and Nowell (2020) to advocate an 'ethnographic' approach (in opposition to 'ethological') to the Neanderthal record in order to account for the diverse ecological, social and cultural lives of these Late Pleistocene hominins, and, one should add, their hominin-animal intersections. Their call resonates with Lestel's (2006: 150) efforts to pave the ground for a vigorous synthesis of ethology and ethnography in the behavioral sciences more broadly, arguing that the 'evolution of social complexity needs to be rethought' in light of hybrid human-animal societies 'sharing meaning, interests and affects' and which are 'more often the norm rather than the exception.'.
Given that 'nature' and 'culture' are distinct categories of Western modernity and often have only limited value for understanding smallscale foraging societies (Viveiros de Castro 1996;Pálsson 1996;Bird-David 1999;Descola 2005, 2010b, Kohn 2013, the search for explanations should also not prematurely stop at the fringes of subsistence strategies, Neanderthal resource management and behavioral ecology. Instead, Neanderthal scholarship must actively explore the multifaceted significance of varied hominin engagements with the non-human world, including non-animate materials such as stone, ivory and bone (cf. esp., Conneller 2011) as well as living organisms such as plants and animals. Current dualist theorythe default point of departure for most Pleistocene archaeologiststoo often suggests, or simply takes for granted, that hominin behaviors and material signatures not directly tied to the fulfilment of the basic requirements of survival and ecological adaptation are the only residuals of culture, and as such earmark milestone cognitive and sociopsychological achievements in hominin evolution (e. g., Renfrew and Morley 2009;Henshilwood and d'Errico 2011;Hoffecker 2017). Those behaviors supposedly intelligible by the lower rungs of the Hawkesian ladder of inference (pace Hawkes 1954) are in turn considered to be confined to 'lower' domains, with no obvious or knowable implications for 'higher' domains such as culture and cosmology. This has reinforced the tendency in Middle Paleolithic research to cut off the discussion of hominin culture from hominin subsistence, even though hunting is a key locus of culture and sociality in huntergatherer societies (cf., Ingold 2000;Nadasdy 2007;Bawaka Country et al. 2015, see discussion below), and thus to refrain from more-thaneconomic, integrative and holistic approaches to Neanderthal-animal interaction.
The classic example for this broader logic of research enveloping other-than-H. sapiens prehistory is the handling of archaeological evidence for the dislocation of so-called 'unusual' or 'exotic' objects, mostly unaltered but sometimes materially manipulated and incised (cf., Marshack 1991: 47-48;Peresani et al. 2013Peresani et al. , 2014Leder et al. 2021), such as fossils, shells, minerals and prominently colored or textured stones, and their incorporation into hominin sites (Bolus and Schmitz 2006: 139;Radovčić et al. 2015;Wilkins et al. 2021). The received dualist narrative identifies these alleged 'curiosities' as signposts of a dawning 'sense of aesthetics' and, relatedy, as emerging 'proto-symbolic capacities' in the hominin lineage (Bednarik 1994;Mellars 1996: 371;Lorblanchet 1999;Frayer et al. 2020). Lorblanchet and Bahn (2017: 55) for example confidently assert that 'the first humans had a sense of aesthetics, and […] they were even endowed with a certain delicacy of touch. Their behaviour seems very close to our own, as we continue to pick up shells, fossils and other natural curiosities'. The manifold examples mobilized in support of this narrative occupy a salient place especially in Neanderthal archaeology (e.g., Frayer 2019), where they are recurrently evoked to bolster the popular idea that these hominins were in many ways 'like us', thereby exposing problematic H. sapiens benchmark assumptions which at times even develop into different forms of sapienscentrism and thus once again revealing the blatant universalist lens that is too frequently deployed in this debate. In point of fact, it is often unclear what exactly eruditions of the above-cited type are supposed to disclose: given broad comparative evidence for multisensuous experience, emotion, affect and even 'aesthetics' in many nonhuman animals (Bekoff 2002;Menninghaus 2011;Watanabe 2012), it seems neither surprising that affective factors played a role in shaping Neanderthal lifeways, nor is it entirely clear how the alleged archaeological recovery of such generalized qualities can assist in understanding the specificity of Neanderthal being and becoming beyond a broad traitregister logic.
From a multispecies and communalist perspective, psychological and proto-symbolic ascriptions detract attention from concrete lifeworld situations and sociodemographic contexts in which these objects make their archaeological appearance. Given that many small-scale forager societies consistently gather stones and other natural objects and for example also bring back seemingly abandoned animal puppies from their local surroundings in order to 'raise them' and/or 'live with them' (cf., Serpell 1989; Bird-David 2017b; Bird-David and Naveh 2008; Simonova 2018; cf., Shipman 2010 on 'cross-species alloparenting') at least indicates that deep-historical 'curiosities' and 'manuports' may similarly inform us about practiced socioecological relations, rather than being mere expressions of so-called 'modern' human behavior and cognition. Often-cited 'non-utilitarian' objects might thus derive from hominin-environment relationships characterized by acute and pervasive immediacy, rather than being the externalizations of hominin 'essences' and inborn qualities, and are then foremost to be regarded as varied forms of material engagement with the inhabited and continuously re-made more-than-human world. In this way, these objects can be acknowledged as potent pointers to situated hominin-ecology intersections and may revealsignifications and perpetual negotiations of extended small-world societies under conditions of pluripresence (pace Bird-David 2017a).
Instead of invoking the detached contemplative capacities and invariant aesthetic sensibilities of Neanderthals and other contemporary non-H. sapiens hominins, such 'special' objects may give testimony to shared and entangled multispecies life in which nature is not merely appropriated but integrated to, and as such part and parcel of, diverse efforts of past society-assembly, place-making and world-building. The respective objects, in this view, are drawn into hominin places and incorporated into occupational deposits not primarily because of their evocative materiality, but as a means of promoting relatedness and 'being-as-many' in more-than-human landscapes (Bird-David 2017a), and thus to affirm social immediacy. This yields an argument from 'horizontality' according to which human beings do not 'walk […] on top of an ontological floor, but attending to and being attended by the bodies surrounding [them], some [] animated and some not, some solid and some liquid and gaseous' (de Carvalho Cabral 2021). Approached from this perspective, Middle Paleolithic curiosities provide evidence for distributed, nonhuman-focused 'horizons of concern' and related Neanderthal efforts to facilitate forms of mutual habitation that are both the 'continent and the content of life' (de Carvalho Cabral 2021) in a multispecies world. The concept of aesthetics that can be derived here closely resembles what pragmatists have rendered as 'aesthetics of the everyday' (Sartwell 2009) or 'social aesthetics' (Coleman et al. 2013)understandings that insist on the embedded nature as well as the interaction, experience and lifeworld-dependency of aesthetic certification (cf., Skeates 2017), hence undermining the dominant cognitivist interpretations in archaeology that emphasize inert pre-social and/or pre-lifeworld capacities with traceable biological, lineage-specific origins. As Welsch (2004) puts it, aesthetics, instead, emerges from mingled human-nonhuman layers of existence and is thus a derived property of more-than-human worlds and their sensous conditions.
In what follows in the subsequent sections, we extend this perspective into the domain of nonhuman carnivores and explore the sympatric ecology of European Neanderthals with regard to raptor, hyena, and ursid others sharing the landscape with these hominins. By synthesizing the available evidence on Neanderthal engagement with these animals and putting the available data into multispecies and integrative anthropological perspective, we hope to show that valuable insights can be gleaned on Middle Palaeolithic lifestyles, adaptations and distinct naturecultures within and between different deep-historical contexts, and that sharing the landscape with other animals and attending to them in a regular manner often had decisive, long-lasting consequences for hominin behavior, society and material culture which archaeologists should begin to examine more systematically, in particular outside of the traditional confines of H. sapiens prehistory.

Raptors: Familiars who soar high
We suggest that understanding Neanderthal sensitivities towards, and engagement with, raptor others can benefit from paying more attention to the co-constructed naturecultures of Late Pleistocene hominins, vultures and other large birds of prey, instead of foregrounding, and thereby prematurely narrowing, the debate in terms of the alleged 'symbolic' proficiency of raptor-derived items of material culture (Finlayson 2019). From a multispecies and dedicated communalist point of view, the recurrent interest of Neanderthals in the talons, and perhaps wings and feathers , of different large raptor species, especially Golden and White-tailed eagles (Negro 2018;Rodríguez-Hidalgo et al. 2019;Laroulandie et al. 2020), points to the significant intersection and emergent socioecological proximity between these hominins and raptor others. Given the predatory profile of Neanderthals at the top of the trophic pyramid of their ecosystems (Gaudzinski-Windheuser and Kindler 2012;Jöris 2014;Smith 2015; Wragg Sykes 2020) and the known structural impact of such social carnivores on their ecologies (Wallach et al. 2015), obligatory and facultative scavengers such as vultures and some large eagles, including the focal species documented in the Neanderthal record (Table 1), would have been drawn to these carcass-accumulating predatory hominins in order to take advantage of hominin-mediated feeding affordances (cf., Moléon et al. 2014). These opportunities can be direct or indirect (e.g., flushing of smaller and medium-sized animals when encircling larger prey, etc.), and are closely linked to the scheduling and nature of hominin behavior in the wider landscape (see Sick 1997 for a nonhuman example; cf., Galetti et al. 2018). Anthropogenic facilitation and opening up of attractive feeding niches for raptors, even though heterogeneous and varying in productivity and impact, is well-demonstrated in presentday contexts (e.g., Bird et al. 1996;Lambertucci et al. 2009) and there is no reason to assume that Pleistocene raptors would refrain from adapting to anthropic landscapes of opportunity. White et al. (2016) have shown that Neanderthals, especially in interglacial environments, were often selective in their ungulate carcass processing behavior, and when taking advantage of favorable ambushalleviating settings, frequently exploited only a subset of the killed animals in a comprehensive mannerespecially at largely monospecific killsites documenting short-term mass death scenarios of animal prey such as the MIS 5/4 or MIS 4/3 site of Salzgitter-Lebenstedt (Gaudzinski and Roebroeks 2003;Gaudzinski-Windheuser 2006cf. Pastoors 2001). Neanderthal subsistence behavior, in other words, would have created a carrion surplus in the wider landscape, which likely acted as a significant hominin ecosystem service mediating broader Middle Paleolithic carrion ecologies (pace Barton et al. 2013). Moreover, bodypart transport patterns and inferred decision-making rationales have certified Neanderthals as 'optimal foragers' focusing primarily on the meat-heavy and nutritionally valuable portions of dead animals as well as importing and/or exporting mainly the respective gourmet parts (Costamagno et al. 2006;Gaudzinski-Windheuser and Kindler 2012;Rendu et al. 2012;Castel et al. 2017;Daujeard et al. 2019). While this behavior can justifiably be described as 'optimal' from an energetic perspective, it similarly implies that carcasses would often have been exploited only partially and in a targeted fashion, providing recurrent scavenging opportunities for other animal co-inhabitants in the same landscape (cf., Moléon et al. 2014: 397). As apex predators in their environments, Neanderthals, through their situated hunting behaviors, would have thus actively contributed to the construction of potent coadaptive interfaces with other carnivores including vultures and occasionally scavenging eagles, hence amplifying the lifeworld association and intersection of hominins and raptors during the Middle Paleolithic.
'Wasteful' hunting has been discussed both for interglacial and glacial Neanderthals (Wragg Sykes 2020: 222, 230) and is evident at the high-resolution forested Eemian lakeshore environment of Neumark-Nord in Eastern Germany, where the close-quarter ambush of two prime-aged stags is linked to low intensity butchering and even the most heavily processed deer carcasses were filleted but not jointed (Gaudzinski-Windheuser et al. 2018;cf., Wragg Sykes 2020: 220-221). 'Wasteful' butchering has similarly been suggested for elephant carcass exploitation at Eemian Lehringen and Gröbern (cf., Thieme and Veil 1985;Mania et al. 1990). As Priestley (2020) has rightly pointed out, however, not only is it problematic to impose a Western conception of 'waste' onto otherwise highly strategic and ecologically perceptive Neanderthals (Uthmeier 2006), non-comprehensive carcass exploitation is actually fairly common among ethnographically-documented northern and circumpolar foragers. For the Yukaghir of Northeastern Siberia, for example, leaving parts of hunted animals at the killsite and being selective about what is taken back to human camp sites is a significant social and cosmological practice (Willerslev 2007: 34-35;Cleghorn 2017;Priestley 2020: 144-145). In a liminal world where bodies are never stable and contained, hunting easily becomes a 'generative' enterprise safeguarding the cycle of life and the sustenance of life processes (Jordan 2003) by submitting part of what has been taken to the land, and therefore literally sharing carcasses with other organisms.
For Neanderthals, hunting of large, meat-heavy ungulates and making 'gourmet' cuts as well as taking only highly praised pieces such as the tongue may thus have played a more-than-economic role by reproducing and reinforcing the intersection with prime animal scavengers, and in this way negotiating the hominin place as predators and hunters in a more-than-human world. The striking selectivity of Neanderthal bone utilization, perceptively noted by Wragg Sykes (2020: 149, 229), which seems to indicate particular cultural norms and meaningful, non-arbitrary associations between particular animal species, specific bones and tool manufacturing decisions and biographies (e.g., Martisius et al. 2020) is consistent with this interpretation and further illustrates the part-focused logic of Neanderthal prey processing and animal body transformation. As argued by Zahara and Hird (2015), drawing on Indigenous Inuit communities' practices in the Eastern Canadian arctic, food leftovers and anthropogenic dumps are 'waste' only from a Western, phenomenologically-detached point of view and in other contexts often promote proximity between humans and nonhumans and hence facilitate the creation and perpetuation of shared multispecies communities and livelihoods.
It is notable that the guild of large avian scavengers was also much larger in the Late Pleistocene than it is today, encompassing vulture species of considerably larger body-size and biomass recycling capacity than their surviving Holocene sister taxa (Hertel 1994;Galetti et al. 2018). Neanderthals and avian scavengers thus participated in broader ecological dynamics rendering the Plio-Pleistocene epochs particularly diverse in scavenging species and niches, both terrestrial and airborne, and in the latter case also comprising other birds such as the highly flexible Corvidae (Galetti et al. 2018). On palaeoecological grounds alone, many of these species were most likely co-adapted and were thus dependent on their respective trophic interaction(s), including emerging forms of sympatry and commensalism, as recently already proposed by Gómez-Olivencia et al. (2018) for eagles and Iberian Neanderthals. The close association between Pleistocene megafauna, involving Neanderthals and other hominins, and facultative or expert avian scavengers , must therefore be regarded not only as evidence for their landscape sharing and heightened ecological overlap, but similarly as a testimony of interspecies co-becoming, promotion and reliance.
The co-occurrence of Neanderthal, raptor and corvid signatures across large parts of Pleistocene Eurasia  would then appear to primarily denote a multispecies assemblage rooted in significant, lived interspecies ties, encounters and consistent associations rather than indexing 'advanced cognitive capacities' or the 'ritual', 'symbolic' or 'religious' status of the birds in question (sensu Finlayson et al. 2019). As Wragg Sykes (2020: 259) has noted, eagles and 'early adopter' commensals such as crows and ravens (pace O'Connor 2013) participated with particular roles in the intra-scavenging guild dynamics of their time (Moléon et al. 2014) and were probably 'familiars' at Neanderthal killsites, even though birds such as choughs were likely encountered not only in the open landscape but also in their cave roost sites (Blanco et al. 2021). There is therefore certainly a striking congruence between the archaeological co-occurrence of Neanderthals, raptors and eagles and their attendance in a shared ecological microcosm of post-kill relationships and biosemiotics ties. Eagles have for example been shown to often rely on clues from corvids to detect carcasses (Grubb et al. 2018;Orr et al. 2019) and eagles, in turn, alert and thus 'call-in' vultures (Kane et al. 2014), giving rise to distinct post-kill biographies of Neanderthal killsites fundamentally centered on these avian others.
The accumulating evidence for Middle and Early Upper Palaeolithic, likely Neanderthal-associated, material engagement with raptors and corvids must therefore arguably be reframed as an emergent consequence of living and intersecting with these nonhuman others in the wider landscape and the resulting situational cross-referencing and interspecies attunement (Fig. 1). The extraction of raptor and corvid feathers (Peresani et al. 2011;Finlayson et al. 2012;Blasco et al. 2016;Romandini et al. 2016), the manipulation of raptor talons and their cooptation as bodily ornaments (Morin and Laroulandie 2012;Romandini et al. 2014Romandini et al. , 2016Radovčic et al. 2015;Rodríguez-Hidalgo et al. 2019;Laroulandie et al. 2020; cf., Table 1) as well as the modification of corvid bones (Majkić et al. 2017) can then be redressed as 'emergent materialities' of being and becoming Neanderthal in the carnivore and scavenger-invested animal ecologies of the early and middle Weichselianand as such as an inextricable part of evolving configurations of millennial-scale interspecies interaction and coordination. Ironically, this reframing of avian-related material culture in the Neanderthal world draws attention to the importance of acknowledging and qualifying the uniquely derived intersection between different hominins and their animal surroundings, including the implied specificity of phenomenological references enveloped by the involved material culture, rather than perpetuating received interpretations construing the significance of these items in terms of cognitive capacity or Neanderthal 'behavioral modernity' (e.g., Peresani et al. 2011;Fiore et al. 2016;Frayer et al. 2020). Finlayson et al.'s (2019) recent attempt to re-cast Neanderthal-raptor relationships as a 'cult of the Sun Bird' with strong culture-historical continuity claims based on direct parallels with practices of ethnohistoric groups in North America and elsewhere illustrates the pitfalls of such sapienscentrism. Finlayson et al. (2019) enthusiastically conclude that 'the long Human-Golden Eagle relationship may end up as the key example of the transmission of a symbolic-cultural behaviour from Neanderthals to Modern Humans'. While we agree with the assertion that hominin-raptor relationships were most likely culturally significant, we strongly caution against overly eager continuity propositions based on decontextualized analogies, naïve evolutionist and diffusionist premises as well as problematic understandings of culture and society as antipoles to 'nature', especially in the Pleistocene past where varying hominin biocultural dispositions are at play (cf., French 2019). Moving forward in the field of Pleistocene interspecies studies instead requires the careful calibration of archaeological interpretations with current multispecies theory and the comparatively assessable specificities and situated behaviors of hominins and animal others in their broader ecological and material contexts.

Hyenas: Neighbors who administer bodies
Pleistocene hyenas, as pointed out by Baynes-Rock (2015b), have customarily been studied in isolation from hominins (Kuhn 2011;Diedrich 2012Diedrich , 2014Diedrich , 2015, and their behavioral domains and ecosystem impacts are deliberately re-cast as separate of the human, thus ironically prefiguring modes of behavioral hominin-hyena rapprochement primarily under the register of antagonism, conflict and competition (Brugal and Fosse 2004;Stiner 2004;Diedrich and Žá k 2006;Villa et al. 2010;Enloe 2012;Dusseldorp 2011Dusseldorp , 2013Discamps 2014). This is astonishing giving the well-documented, close ecogeographic association of early hominins and various hyena taxa throughout the Lower and Middle Pleistocene, especially in Africa (Turner 1990;Werdelin and Lewis 2005;Treves and Palmquist 2007). Even though the involved taxonomic groups change over time (Turner et al. 2008), Hominidae and Hyaenidae share an exceptionally long evolutionary history (Brantingham 1998;Stiner 2012), and either of the two is often a strong proxy for the presence of the respective other, as reflected in early expansions of faunal assemblages with hominins and hyenas from Africa into the Eastern Mediterranean and Western Europe (Arribas and Palmqvist 1999;Lewis and Werdelin 2010;Brugal et al. 2020), for example documented at key sites such as Dmanisi (Coil 2016), Pakefield (Parfitt et al. 2005;Iannucci et al. 2021) and Atapuerca (García and Arsuaga 2001).
Yet, this co-occurrence, and possible history of co-dispersal, has rarely been examined from a sympatric and sociocultural point of viewthat is, in terms of coevolution in the above outlined sensebut mainly as a phenomenon of niche segregation and differentiation and the eventual competitive exclusion of the last hyenas in Late Pleistocene Europe (but not in Africa; cf., Stiner 2012; Galetti et al. 2018). Decentering of mutualities and more-than-hostile interstices between hominins and hyenas reflects the Western conviction of portraying biological organisms as ecologically 'authentic' as possible, which in the classic rendering is taken to call for a study of organisms 'as they are by themselves' (Baynes-Rock 2015a, 2015b), independently of other interfering organisms (especially humans) and the adaptive 'noise' they produce. We suggest that the archaeological record of Late Pleistocene hominins other-than H. sapiens, in particular Neanderthals in Western Eurasia, therefore offers a useful model case to flip this received vision of the past on its head and begin exposing the evolved, intricate socially and materially productive, co-adaptive interfaces between hominins and their contemporary hyena neighbors in changing spatiotemporal contexts.
The ancestral cousins of today's spotted hyenas (Crocuta crocuta) arrived in Europe shortly before 300 kya Turner et al. 2008), broadly falling into the second half of the Middle Pleistocene and thus predating 'classic' fossil Neanderthal phenotypes which only appear around MIS 5 (Serangeli and Bolus 2008). This timing bears evolutionary significance because what we recognize as Neanderthal biocultural characteristics developed gradually between ca. 350 and 300 kya, and thus in hyena-bearing ecosystems. Hyenas can even be considered to have been 'superabundant' in some of the ecologies mediating this process of 'Neanderthalization' (Turner et al. 2008;Galetti et al. 2018), with a possible palaeodemographic 'Hyena event' at the end of MIS 3 in Western Europe (Discamps 2009(Discamps , 2014; see below). The combined archaeological and paleontological records clearly show that MIS 5-3 Neanderthals, especially in Western, Northwestern, Southern and Central Europe, consistently shared the landscape with these opportunistic and highly successful carnivores over many millennia (Fig. 2;Brugal and Fosse 2004;Arsuaga et al. 2011;Brugal et al. 2020). Neanderthals and hyenas also regularly used or at least visited the same localities, especially caves and other karstic landscape features, in temporal succession or not (e.g., Straus 1982;Brugal and Jaubert 1991;Villa and Soressi 2000;Villa et al. 2004; Beauval et al.  Table 3 (site numbers correspond to site IDs in  Table 3; Zaskalnaya VI and Matuzka are located outside of the shown map section). Colors indicate inferred site category and shape of symbols represent period/ techno-complex contexts (MP = Middle Paleolithic; LRJ = Lincombian-Ranisian-Jerzmanovician). Animal outlines from http://phylopic.org/ via Public Domain. 2005;Miracle 2005;Brugal, 2010;Sala et al. 2012;Discamps et al. 2012;Turner et al. 2013;Morley et al. 2019). The quality of some of these sites as 'persistent places' (sensu Shaw et al. 2016;Maher 2021), furthermore, often derives from the space-altering activities of both agents and is to this effect 'multispecies' in nature. There are several reasons to believe that this notable spatiotemporal entanglement of Neanderthals and hyenas was not just a matter of self-confined geographic coexistence and fierce competition, but also implied heightened levels of interspecies immediacy and coordination with significant sociomaterial import, eventually leading to the development of 'mutual ecologies' (sensu Fuentes 2010; see above) which co-shaped the evolutionary trajectories of both hominins and hyenas.
First, the breadth of currently available evidence on hyena behavior in the present suggests that these animals are hyperflexible in their local adaptability and benefit from their pronounced ecological plasticity to make a successful living in varying environments and under shifting ecological regimes (Boydston et al. 2003;Abay et al. 2011;Mhlanga et al. 2017). After all, hyenas appear to be to a lesser extent adapted to specific conditions than to the exploitation of feeding opportunities that present themselves in different ecosystems, supported by their adept 'cooperative problem solving' (Drea and Carter 2009;Benson-Amram and Holekamp 2012) and mediated by multilevel, typically femaledominated clan-societies Silk 2019;Ilany et al. 2021). In places where hyenas have survived until today, for example in the Horn of Africa, they have, in part because of their flexibility, almost always developed intricate relationships with humans and adapted to human-shaped environments (Gade 2006;Abay et al. 2011), to the point where hyena lifeways have become entirely dependent on their human neighborhoods (Baynes-Rock 2013, 2015a; Yirga et al. 2012).
Humans, in turn, have similarly adapted to their hyena neighbors, developing particular modes of relating and integrating hyenas into the cultural fabric of everyday life (Baynes-Rock 2015a). In the Harar region of Eastern Ethiopia, hyenas are for example bound up with human khat (Catha edulis) consumption and circulation, which the animals are thought to promote through their protective presence and oftenoverlooked pathogen clearing service (Fuentes and Baynes-Rock 2017). Hyenas are not only socially meaningful neighbors, who are involved in local identity formation and endow human stories and gossipingi.e., so-called 'hyena talk' -as well as cultural understandings of landscape, they are sometimes deliberately fed close to or in human settlements (Baynes-Rock 2015a: 37) and as 'caretakes' of shrines and other important places receive special gifts to reaffirm human-hyena 'commons' (Baynes-Rock 2013), explicitly conceived as a form of 'social contract' (Baynes-Rock 2015a: 58-72). As Baynes-Rock (2016: 288) notes, the Harar data on human-hyena relations highlight that 'people's conceptual worlds are not so much constructed and structured within their minds as they are unfolded with Others in spatiotemporal, historically informed, phenomenal worlds.'.
It is notable that human-hyena interstices seem to be particularly stable and culturally productive where humans rely to a large degree on animal products, especially meat, from medium to large-bodied ungulates (Baynes-Rock 2013: 215). Hyenas have plenty of feeding opportunities in these human-shaped contexts and at the same time provide valuable ecosystem services by taking care of carcasses, food scraps and residual bones. Even in settings with a less developed human fingerprint such as in protected areas, natural reserves or even national parks, hyenas adjust and synchronize their behavior in various ways with regard to human activities and exploit the few anthropogenic feeding opportunities that arise in these landscapes (Kolowski and Holekamp 2007), often at their outer fringes. Humans are therefore always an integral part of hyena ecologies, and this is probably also because hyenas have evolved in multispecies landscapes shared with hominins over thousands of years (cf., Moléon et al. 2014;Baynes-Rock 2015b). The mutualities of human-hyena relations and their proximate co-ecologies must consequently be re-considered also in their deep-time contexts.
The Neanderthal-hyena association in the European Late Pleistocene seems hardly coincidental then: Neanderthals were themselves powerful ecological keystone species and exhibited an elevated carnivorous profile (Richards et al. 2000;Bocherens 2011;Gaudzinski-Windheuser et al. 2018;Jaouen et al. 2019), even though plants and other food resources unquestionably also played a role in their diets (e.g., Hardy and Moncel 2011). Yet Neanderthals have been shown to consistently occupy the trophic apex of their ecological communities and to be capable big game hunters systematically targeting large-bodied mammals of up to 800-1000 kg (Uthmeier 2006), especially ungulates, even though their species preferences varied in space and time (Gaudzinski and Roebroeks 2001;Niven et al. 2012;Yravedra and Cobo 2015;Morin et al. 2016).
The key point is that Neanderthals were among the landscape-scale carcass accumulators par excellence of their time, and Neanderthal presence certainly ensured the availability of predictable and specific types of carrion in the wider landscape. In contrast to later H. sapiens populations, Neanderthals, as noted before, also do not seem to have habitually exploited the entirety of animal carcasses and often selected only some of the available animal materials for further transformation (Patou-Mathis 2000: 393;Richter 2018: 202), often leaving behind for example the axial elements of the animals they killed (e.g., Valensi and Psathi 2004;Gaudzinski and Niven 2009) and in this way promoting a broad feeding niche especially for bone-cracking, osteophagic hyenas.
Based on their critical review of MIS 5-3 Neanderthal kill sites including Mauran, La Borde, Taubach, Zwolén and Salzgitter-Lebenstedt, White et al. (2016) have furthermore shown that these hominins recurrently took advantage of opportune hunting situations and favorable topographic conditions, and often killed more ungulates than they excessively exploited (see also Daujeard et al. 2019) -they appear to have been fairly 'discerning' diners adding to the image of selective carcass processors and transporters discussed before. This behavioral proclivity would have facilitated hyena scavenging and simultaneously drawn hyenas and Neanderthals closer together in the landscape. Neanderthal sites with bone accumulations and evidence for secondary hyena interference are generally consistent with such a coadaptive scenario (e.g., Majkić et al. 2018;Sanchis et al. 2019;Daschek and Mester 2020), now also supported by ZooMs findings, for example from Denisova cave , showing the chronostratigraphic importance of hyena activity, material accumumation and secondary bone modification and/or fragmentation at many Neanderthal sites across Eurasia. At Denisova, Morley et al. (2019) document near-continuos hyena occupation with a pervasive coprolite record and observe that '[s]pecific areas of the site might have been designated as waste dumps for lithic debitage and food detritus, […], which in turn attracted scavengers such as the cave hyena when hominins were absent from the site.' Secondly, broader MIS 5-3 Neanderthal land-use systems appear to have considerably overlapped with hyena core territories and den localities (Fosse 1997;Armand 1998;Stiner 2004;Dusseldorp 2013;Crezzini et al. 2016;Jimenez 2017), and Crocuta places and hyenainvested locales seem to have regularly been integrated, although in complex and not always easy-to-decipher ways, into Neanderthal mobility patterns and land-use systems (Enloe et al. 2000;Villa and Soressi 2000;Villa et al. 2004Villa et al. , 2010Huguet et al. 2010;Beauval and Morin 2010;White and Pettitt 2011;Discamps et al. 2012;Picin et al. 2020). Given the already noted territorial stability of Neanderthal groups inferred from eco-cultural niche models (Banks et al. 2021) and other lines of evidence indicating that Neanderthals frequently returned to the same places and sometimes valleys (cf., Wragg Sykes 2020: 216-218), we suggest that this interweaving of Neanderthal and Crocuta geographies and the resulting mixing of hominin and hyena materialities (e.g., bones, artefacts) in particular landscape contexts, and often in recurrent fashion, reflects socioecological and landscape proximity between the two agents (cf., Jimenez et al. 2021: 2), thus pointing to situated modes of engaging with hyena others that were an integral part of Neanderthal society and culture. Hyenas co-defined the tracks, pathways and rhythms typifying and binding together Neanderthal landscapes (pace Gamble 1998) and thus arguably co-shaped both the temporality and spatiality of hominin life in many parts of the Middle Palaeolithic.
Examples of such hybrid more-than-human landscapes and their interlaced Neanderthal-hyena geographies are not difficult to pinpoint in the European Middle Palaeolithic. In the Bohemian karst and the area surrounding Prague, for example, a dense lineup of caves and palaeoravines with evidence for pronounced hyena activity (Diedrich and Žá k 2006) meets broadly penecontemporaneous but relatively sporadic hominin presence with only a few sites such as Bečov documenting more extensive Neanderthal occupation, while many other Middle Palaeolithic sites have yielded only a handful of lithic artefacts (Valoch 2013a), either associated with hyena contexts or not. The Sauerland karst region in Western Germany similarly documents microregional overlay of major Neanderthal occupation sites such as Balve cave (Kindler 2012) and Volkringhauser Höhle (Tafelmaier 2011) and intense hyena denning activity, for example well-attested at Perick caves and Wilhelms cave (Diedrich 2009(Diedrich , 2011. Another example is Southwestern France where diversified Neanderthal settlement occurs within rich hyena landscapes (Discamps 2011), including a large number of penecontemporaneous den sites such as Plumettes, Camiac and La Chauverie, some of which have yielded small numbers of lithic artefacts and human-modified bones in dense hyena-accumulated deposits (Discamps et al. 2012;Airvaux et al. 2012). A final example is provided by the Belgian Ardennes where a cluster of important Late Middle Palaeolithic sites including Spy, Scladina, Walou and Goyet, in which the majority of human-accumulated faunal assemblages were also affected by subsequent hyena activity (Daujeard et al. 2016), are found in direct spatial proximity to the recently identified long-term hyena birthing den of Caverne Marie-Jeanne, dated to the same narrow MIS 3 timeframe as the adjacent Neanderthal sites (Jimenez et al. 2021). Interestingly, cementum analysis of teeth from hyeans and their prey from the latter region has detected a marked absence of seasonal differences in occupation, suggesting that hyena populations where firmly established in, and regionally attached to, the Belgian Ardennes (Jimenez 2017). All of this suggests that hyenas were familiar co-inhabitants and spatial 'companions' (pace Haraway 2007) as well as key configurators of Neanderthal lived experience in various European regions.
A critical review of the European Neanderthal record further indicates that there are at least three different types of archaeological sites which can be discriminated in order to elucidate the dynamics of hominin-carnivore engagement (Table 2), and all three site types appear to have been a common component of Neanderthal land-use systems, spatial understanding and 'landscaping' (pace Arroyo-Kalin 2016), even though not all of them were necessarily of equal importance for different Neanderthal groups at different times and in different regions (cf., Fig. 2): • 'Type one' sites encompass archaeological occurrences where material culture is abundant and hominins appear to have been the primary agents of site formation and material accumulation but subordinate carnivore activity is sometimes also attested. • 'Type two' sites comprise archaeological occurrences in which Neanderthal material culture is frequent but carnivore behaviors and remains have also contributed substantially to the formation and nature of the deposits; these sites often document alternate frequentation of hominins and other carnivores, including Pleistocene hyenas. • 'Type three' sites are characterized by the anecdotal presence of hominin material culture, often only a few selected items such as finished and curated lithic tools, and carnivores are the primary formative agents; these sites sometimes bear partial hominin skeletal remains or isolated bones and teeth (e.g., Tournepiche 1994;Stiner 1991Stiner , 1994Beauval et al. 2005;Enloe 2012;Diedrich 2014; see discussion below) and frequently correspond to carnivore habitation sites such as hyena dens.
The three site-types are distinguished by a differential contribution and mixture of behavioral and material signatures of hominins and hyenas. This interweaving of hominin and hyena spatialities again reassures us of the geosocial proximity between the two and that they were differentially engaging with each other not only in the open landscape but in different landscape situations and with reference to different kinds of places with varying degrees of hominin and hyena 'authorship', even though time averaging, taphonomic filtering, occupational histories and the details of site formation need to be carefully considered on a case-to-case basis. This complex and spatially differentiated pattern of hominin-hyena exposition involved hominin visitations of hyena occupation sites (cf., Airvaux et al. 2012;Rodríguez-Hidalgo 2010;Discamps et al. 2012;Picin et al. 2020: 3), and the other way around, for example when hyenas had secondary access to otherwise anthropic faunal accumulations (e.g., Costamagno et al. 2005;Daschek and Mester 2020) and possibly ravaged Neanderthal deposits and fossil remains (Diedrich 2014). Direct evidence of two butchered hyena remains within the otherwise nonhuman-accumulated deposits from the depths of Maltravieso cave in Southwestern Spain broadly dating to MIS 5 indicates that the exploration and potential scouting of hyena-infused places has an extended history at least among some Neanderthal groups (Rodríguez-Hidalgo 2010).
All of this supports a potent co-adaptive nexus of Neanderthals and hyena others and highlights that the associated interspecies neighborhoods mattered a great deal, steering and variously co-configuring hominin behaviors and 'horizons of concern' (pace Bird-David 2017a). The affirmation and reproduction of 'mutual ecologies' (pace Fuentes 2010) was thus perhaps a more important factor in shaping Neanderthal spatial organization and landscaping than previously acknowledged. The mixing of Neanderthal and hyena materialities in different locations in the landscape may accordingly not only tell us something about the nature and scheduling of subsistence-related tasks, but also reflect ongoing negotiations of place and the significance of sharing the landscape with nonhuman carnivores. Given that movements and material Table 2 Three-tiered categorization of archaeological sites with implications for understanding carnivore neighborhoods and hominin-carnivore intersections in different palaeolandscapes.

Signature(s)
1 Hominin material culture is abundant and operational sequences tend to be complete; important role of local raw materials and their on-site transformation/consumption; artefact-bearing layers are primarily anthropogenic in nature; reduced evidence of carnivore bone interference; primary access of carcasses by hominins; pronounced spatial organization of hominin activities and evidence for repeated or longer-duration stays 2 Hominin material culture is frequent but operational sequences are fragmented; carnivore remains are frequent and carnivores have substantially contributed to bone accumulation and archaeological site-formation; non-local raw materials and the import of finished and/or curated tools or cores play more important roles; elevated frequencies of gnawing and bite marks; often but not always alternate frequentation of hominins and carnivores; ephemeral/short-term hominin occupations 3 Hominin material culture is anecdotal or extremely rare and only specific items (e.g. bifacial tools) or fragmented/isolated hominin fossils are found; incomplete operation sequences with emphasis on the distal segments; evidence for carnivore habitation/denning/ hibernation, e.g. abundant coprolites; elevated ratio of juvenile individuals among bones and teeth (including presence of cubs), >10 % gnawing mark frequency; evidence for primary access of carnivores to at least some animal carcasses; burrows or other animal habitation, hibernation and feeding structures; ephemeral/shortterm hominin occupation or 'visitation' signatures traces in the landscape can be viewed not only as physical concretizations of past behaviors but equally as residues or flows of knowledge (McBryde 2000; Rockman 2013), Neanderthals were apparently concerned with acquiring hyena-related landscape knowledge, to learn about hyena places and to familiarize themselves with these proximate carnivores, so that hyena spatialites would figure prominently in Neanderthal social memory. The deliberately incised fragment of a hyena femur from Les Pradelles in Southern France (d' Errico et al. 2018) -a 'type one/two' site dating to MIS 4 -provides an intriguing example of material engagement with hyenas as important landscape co-dwellers, and can be reinterpreted as a form of 'emergent materiality' within socioecologically meaningful, yet nevertheless precarious hominin-carnivore neighborhoods. The incised hyena femur makes its appearance in layer 2a of Les Pradelles, which has been interpreted as a specialized Quina Mousterian reindeer hunting and processing station with considerable hyena input (Costamagno et al. 2005(Costamagno et al. , 2006. The site also yielded an unusual assortment of intensively processed Neanderthal body parts, originally belonging to at least 9 individuals (Wragg Sykes 2020: 303). Interestingly, the condition of the hominin bones is much more heterogeneous than the associated primary anthropogenic ungulate fauna, and a major hominin concern was body and bone fragmentation, and perhaps export, as indicated but the unequal representation of hominin body parts (Mussini 2011), also documented in other Middle Palaeolithic sites with excessive hominin-oriented body manipulation such as Krapina, Moula-Guercy and El Sidrón (cf., Wragg Sykes 2020: 300-307). Not only does this treatment structurally mimic the carcass ravaging and bone-cracking behavior of hyenas, there is also evidence for hyena consumption of hominin teeth as well as considerable hyena-induced damage on hominin bones themselves .
Even though it is not entirely clear whether the marked hyena femur is of local origin or was imported to the site, both hyenas and Neanderthals had some kind of access to each other's bodies, and the broader context of layer 2a evokes the fluidity and permeability of homininhyena interstices involving material exchange and mutual body transformation, which not only created a mingled multispecies assemblage but also links the biography of hominin remains to place-bound hyena agency, highlighting the world-making capacity of these nonhuman carnivores. Moreover, Les Pradelles is one of the few Middle Palaeolithic sites where hominin bones were selected and co-opted as retouchers and the respective bones are robust femur shaft fragments (Rougier et al. 2016;Costamagno et al. 2018b), and thus, analogically speaking, reference the same body part as the marked hyena femur found at the site. The incised hyena bone from Les Pradelles, we would argue, has to be understood against the larger background of this complex reticulation of human and nonhuman materialities and associations. As Wragg Sykes (2020: 303) has pointed out, there is good reason to conceive of the involved behaviors as 'acts of intimacy, not violence' and this point can be nuanced by taking stock of the broader hominin-animal-material interrelationships and material exchanges and transformations implicated at Les Pradelles. Interpreting the marking of the hyena femur as 'emergent materiality' of specific place-bound Neanderthal-hyena intersections has the advantage of recognizing the situational context of human and nonhuman agencies and references as a key motivation for the observed material engagement, without overrating (or underrating) the idiosyncrasy of the object in question.
In addition, Les Pradelles draws attention to larger ecosystem dynamics during MIS 4 -the heyday of the Quina Mousterian (Jaubert et al. 2011;Faivre et al. 2017) -when the total herbivore biomass in Southwestern France seems to have decreased (Discamps 2014) and large ungulate herds would have been focal landscape agents and carnivore attractors. The apparent specialization of Quina Neanderthals on reindeer groups and their strategic interception resulting in extensive, spatially dense bone dumps would have thus acted as particularly strong magnets for hyena mobility and consumptive activity, with the interesting possibility of at least partial specialization in the predictable expoitation of Neanderthal hunting leftovers. Neanderthal sites like Les Pradelles and Jonzac document the recurrent use of certain locales for reindeer processing by hominins and include evidence for noncomprehensive carcass treatment during the Quina Mousterian (Costamagno et al. 2006;Niven et al. 2012). The mixing of Neanderthal and hyena signatures at these places would have thus created a powerful background for distributed, multispecies place-making, where hominins and hyenas were not only drawn together but co-created the defining sense, experience and signification of place through mutually reinforcing actions. It is this special association of 'Neanderthal-ness' and 'hyena-ness' that arguably crystalizes in the incised hyena femur from layer 2a.
In Western Europe, especially in France (e.g., Brugal and Jaubert 1991;Airvaux et al. 2012;Discamps et al. 2012), Germany (e.g., Müller-Beck 1968;Hülle 1977;Böttcher et al. 2000;Uthmeier et al. 2018) and on the Iberian peninsula (e.g., Rodríguez-Hidalgo 2010; Brugal et al. 2012;Picin et al. 2020), there is also growing evidence for Neanderthal ventures into 'domestic' hyena spaces as indicated by 'type three' sites with a reduced hominin fingerprint, often associated with hyena maternity dens with plenty of coprolites and youngster-dominated mortality profiles (cf., Table 2, Table 3). For instance, localities interpreted as important hyena dens or hideouts in Spain such as Cueva del Camino ), Cueva de la Buena Pinta (Huguet et al. 2010) or Portalón del Tejadilla (Sala et al. 2020) have yielded sparse evidence for hominin presence, often a few isolated lithic artefacts only. A cutmarked lynx humerus with superimposed carnivore toothmark from the hyena den of Cueva de la Zarzamora may also result from Neanderthal excursions into hyena living grounds, even though other interpretations are also possible . In Iberia, hyenas generally became hyperubiquitous carnivores at the end of the Middle Palaeolithic (Sauqué et al. 2017) and the tightened intersection of hominins and hyenas documented there archaeologically is thus perhaps also a consequence of this emerging ecological salience Picin et al. 2020).
Practices of visiting hyena spaces and their varying material correlates can be viewed in connection to the special importance of hyenas as landscape agents and proximate neighbors. Visiting alien spaces 'stained in hyena' may then, generally speaking, express situated strategies of relating to hyena others, affirming and reproducing lived ecocultural adjacency and propinquityand this holds independently of whether or not hominin presence was concomittant with hyena den occupation or not since hyena-stained places can be understood as part of the extended phenotype and personhood of hyenas in a wider, shared landscape. In a similar vein as Politis (1996) has argued that human foragers 'move to produce', we consequently suggest that Neanderthals may in part have 'moved to relate', and some nodes in their mobility networks can thus probably be re-examined as interspecies 'contact zones'. This mode of relating to hyena others arguably points to a broader ethos of sharing that goes beyond mere resource competition and antagonism, indicating that hyenas participated in the larger social field of hominins and were recognized as significant others, perhaps even persona (Fig. 3). Rather than contradicting with, this explicitly encompasses practical uses of hyena dens as part of the 'giving environment' (pace Bird-David 1990), serving as valuable faunal caches (cf., Discamps et al. 2012) and thus facilitating the acquisition of specific, thought-after animal bones and body parts. In modern-day India, for instance, scavengers facilitate the collection of high-quality cattle bones since they efficiently clean the animals' carcasses and make them readily available, thereby providing important 'cultural ecosystem services' (Markandya et al. 2008). Bradley's (2000) arguments on the importance of landscape biography and 'natural' places in the making and evolution of human landscapes are also worth recalling here. The frequently encountered occupational sequence of alternating habitations of hyenas and Middle Paleolithic hominins, often beginning with hyena denning activities followed by more-or-less structured hominin domestic use of space (e.g., Spagnolo et al. 2020) may then be positively turned-as reasserting Table 3 Selected archaeological sites from Western, Central and Southern Europe dated to between MIS 5 and 3 with evidence for different modes of co-habitation among hominins and hyenas or bears. Sites are classified according to the criteria listed in        hominin-hyena complementarity, home sharing and the lived significance of hyena-shaped locales. Former hyena occupations were inscribed into hominin site biographies and as such became phenomenologically foundational to hominin place-making. Hominin site selection in former hyena country, overlaying former hyena homes, may then in itself represent a strategy of relating and thus a means of promoting pluripresence with hyena neighbors. At sites like Teixoneres in Northeastern Spain, where detailed intraspatial analysis of Unit III has shown that Neanderthals respected the habitation area of hyenashominins occupying the areas close to the cave entrance and hyenas denning in the inner sheltered zonesand recurrently used the same sectors to house their short-term occupations (Zilio et al. 2021), it is certainly difficult to dismiss the layered multispecies history of the site as a place-defining feature. The structured palimpsest of these sites can be viewed as framing the phenomenology of place, rather than reducing palimpsest to a barrier of insight (cf., Bailey and Galanidou 2009), and hence testifies that Neanderthal landscapes also incorporated 'the powerful fact that life must be lived amidst that which was made before' (Meinig 1979: 44), both human and non-human. Additional evidence for potentially meaningful ecocultural and material negotiations between hominins and hyenas comes from the LRJ ('Lincombian-Ranisian-Jerzmanovician'; sensu Flas 2011) and related Late Middle Paleolithic/Initial Upper Paleolithic (IUP) technocomplexes in Northwestern and Central Europe (Bolus 2004a;Uthmeier 2004;Jacobi et al. 2007). At the classic type-locality of Ilsenhöhle near Ranis in Thuringia, Germany (Hülle 1977), MIS 3 hominins left behind a small assemblage of finished Jerzmanovice blade points and symmetrical bifacial leaf-points in a cave bear and hyena-rich depositional context in close proximity to possibly penecontemporary hyena dens. The bifacial leaf points are particularly well-crafted and are made of high-quality Baltic flint, imported to the site from a distance of at least 30 km. These artefacts exhibit only limited traces of prior use or evidence for recycling and edge-maintenance as is otherwise characteristic for similar bifacial tools (cf., Iovita 2014; Weiss 2020). Fifteen of the 24 blade points and 13 of the 19 bifacial leaf-points were discarded in a complete, near-mint state (Hülle 1977: 106). The lack of any conclusive evidence for on-site toolkit maintenance opens up the interesting possibility that these certainly valued objects were deposited or at least abandoned in hyena country. This tentalizing reading provides an important challenge to standard subsistence-economic or simplicityforegrounding interpretations of Late Middle Paleolithic/IUP hominin behavior rendering them as mere discard products, and, in line with the broader re-conceptualization of hominin-nature relations attempted here, thus re-configures the burden of proof as well as received assumptions on the hierarchy of evidence.
Other 'type two' and 'type three' assemblages from the Central European Blattspitzengruppen are similarly dominated by finished tools, especially partial and fully bifacial points, bifacial foliates and various bifacial knife-types (Richter 2009), and the operational sequences documented at these sites are frequently fragmented, with little or no evidence for initial stages of tool confection and primary blank production (Bolus 2004b; cf., Table 3). Furthermore, two notable examples for the recovery of a small number of finished tools in hyena contexts are provided by a single but typical Jerzmanovice point alongside a few plain flakes from Zwergloch cave (Müller-Beck 1968) and the exceptional Late Middle Paleolithic leaf points from Haldenstein (Riek 1938) in Southern Germany, both found in hyena-rich deposits. Another example is Kirchberghöhle near Schmähingen in Southeastern Germany, dated to between ca. 40 and 48 kya, where only four lithic artefacts, three classic Jerzmanovice blade points and a basal blade point fragment, plus a nondiagnostic fragment were recovered alongside a large faunal assemblage bearing a diagnostic hyena fingerprint (Uthmeier et al. 2018). Such potential practices of signifying hyena-invested places with non-local tools embodying hominin individual and group biographies may not only be understood as a symptom of hominin landscape socialization and enculturation, but coevally as a historically situated attempt to negotiate the place of Neanderthals in hyenasaturated, carnivore-rich worlds.
A similar pattern of hominin-hyena coassociation has been noted for the LRJ in Great Britain (Jacobi et al. 2007;White and Pettitt 2011). Sites such as Kent's Cavern, Hyena Den, Pin Hole and Robin Hood's Cave all represent hyena-dominated archaeological deposits containing only a handful of finished lithic tools in mint or close-to-mint condition, again especially Jerzmanovice points and cognate pointed blade tools, as well as a few bifaces and bifacial points of likely slightly older Middle Palaeolithic origin (cf., Wragg Sykes 2017; Table 3), thus possibly similarly referencing the significance of these nonhuman places and the related deposition or placement of selected items of material culture as a form of relating, interspecies negotiation and meaning-making (cf., Fig. 3). At Coygan cave in South Wales, hyenas were the primary agents of accumulation while three Late Middle Palaeolithic handaxes and two flakes attest the short-term presence of Neanderthals at the site (Pettitt and White 2012: 336-337). The handaxes exhibit extended object biographies and are manufactured from non-local raw materials imported from sources ca. 20-40 km away. The curated artefacts were found in a peculiar position within the cave interior, turned against the north wall, and their deliberate caching has been considered (Pettitt and White 2012: 337) but remains difficult to securely establish given the poor documentation of the original excavations. Another special find illustrating possible depositional hominin behaviors in hyena territory comes from Bench Quarry 'tunnel' where a single leaf point on a blade was placed directly underneath a hyena jaw and taphonomic or postdepositional processes can be excluded as formative agents of this lithichyena association (Jacobi et al. 2007: esp. Figs. 54a-b).
At Grange Farm in Central England, radiocarbon dated to between 40 and 44 kya, a small assemblage of LRJ lithic artefacts including a complete and a broken leaf/blade point and a small number of trimming flakes related to on-site blade point maintenance were recovered next to an open-air hyena den and a few proposed meat storage pits for hyena cubs (Cooper et al. 2011). The articulation and intermixture of hominin and hyena signatures at Grange Farm is interpreted as evidence for MIS 3 hominins venturing into hyena territory (pace Pettitt and White 2012: 324), possibly to scavenge food caches in or close to hyena maternity dens (Cooper et al. 2011: 91). The archaeological data from Grange Farm (a 'type three' site) is thereby broadly consistent with a short-term 'raiding party' scenario involving only a few hominins, and generally supports the emerging complexity of hominin-hyena relations in MIS 3, including the interpretation of hyena spaces as an integral part of the broader 'giving environment' of Late Pleistocene hominins (sensu Bird-David 1990; see above). Drawing on Bird-David's (1990: 190-191) anthropological insights, it might indeed be sensible to re-contextualize the broader observations from the Late Middle Palaeolithic and the LRJ in terms of an integrated system of giving and takingand thus in light of generalized transspecies exchange theory (Nadasdy 2007) rooted in the social anthropology of the Maussian 'gift' (Mauss 1967;Peterson 1993).
As Nadasdy (2007) has pointed out, hunting-gathering lifewaysespecially in northern latitudes and arguably independently of their specific hominin contextrevolve around human-animal intimacy, codependency, mutuality and continuous exchange. Because of the existential significance of hunting and cognate practices in many small-scale Northern foraging communities, even seemingly inculpable, subsistence-oriented activities are fundamentally social and articulate the need to negotiate, maintain and reproduce human-world relations in which animal others regularly encountered or engaged are focal nodes. 'Hunter-gatherer', then, is not just an economic label, an adaptive signification or a 'mode of production' but instead denotes a general mode of being-in, relating to, and experiencing the world (Ingold 2000), with often varied but decisively non-Western ontological ramifications. Not only is there widespread recognition of the hunted animal as a 'gift' incurring future obligations and potential 'counter-gifts' on behalf of the hunter, animals are met as sociopolitical interlocutors, powerful as well as potentially dangerous 'trading partners' and thus demand respect and reverence, often safeguarded by sophisticated cultural techniques (Nadasdy 2003(Nadasdy : 83-108, 2007. Ingold (2000: 61-76) argued that the relationship between the hunter and their prey is based on 'trust', in turn resting on bilateral autonomy and co-dependency. Indigenous people's talk of animal others as 'offerings' or as 'giving themselves' to the hunting communityfor example well-described in the context of circumpolar whale (Demuth 2019) or elk hunting (Willerslev 2007: 102) -is therefore not just 'metaphorical' or 'symbolic' as Nadasdy (2007) aptly reminds us, but instead expresses the lived social fact that animals are an integral part of society and as such participate in cycles of exchange and affirmation, just like other humans do, for example when they share meat or circulate objects to ascertain relatedness or belonging (i.e., as seen in the 'Kula ring' of Melanesia or potlach practices across North America; cf., Malinowski 1922;Nadasdy 2007: 28). Fuglestvedt (2012: 18-21, 2018.8) and others (cf., Corry 2011: 53f.) have framed the notion of the 'hunting cycle' to capture this reciprocal dynamic of giving and taking permeating the human-animal interface in many foraging societies. Importantly, however, and as already highlighted by Mauss (1967: 23), gifts are not always freely given and often require particular strategies to be facilitated. The implied reciprocity is also not always positive: reciprocal gift exchange may thus easily involve potentially adversarial relationships. What is more, the exchange relations themselves can be asymmetrical insofar as different 'trading partners' may give and receive gifts in different ways and by different means, and this makes social exchange particularly challenging to detect and examine in interspecies contexts.
Hominin depositional practices broadly construedi.e., without necessarily involving structured or formal depositionincluding assemblage mixing, object transformation and discard behaviors may then be reconsidered in the light of a broader logic of material and social exchange at the hominin-hyena interface. Taking from hyena places and hideouts, for example by scavenging and foraging bones and meat caches, may have implied the obligation to give something 'hominin' in return, and some of the finished and curated tools and the isolated pieces of human-modified bone encountered in hyena places may consequently be interpreted as evidence for situated and multi-stranded processes of interspecies negotiation and exchange. Material transformation and fragmentation may be regarded as a particularly potent locus of such generalized exchange relations since hyenas are the osteophagic raw material transformators and fragmentators par excellence and mimicking this behavior thus not only maps hominins onto hyenas, and in this way produces analogy, similarity and proximity, but perhaps also signifies the conviction to submit hominin materiality to hyena domains. Mixing itself may then be viewed as a community-creating practice. From this perspective, the image of the 'raiding party' invoked above is minimally misleading and reinforces resource-competition tropes rather than being attentive to the larger ecologies of exchange in which such practices are often embedded.
This integrated ecocultural framework also recognizes that abandoning hunting implements or stone tools more generally can hardly be considered a neutral act, as foraging people variously regard such objects as part of their 'extended personhood' (González-Ruibal et al. 2011;Walls 2019;Snow 2020) and/or maintain quasi-interpersonal relationships with them (Weedman Arthur 2018), again undermining received interpretive wisdom in Paleolithic archaeology based on long-standing nature-culture divisions. Given that proximate animal others are often regarded as 'extended kin' in such small-scale societies (cf., Norton-Smith 2010;Willerslev and Ulturgasheva 2012;Bird-David 2017a;Wisher and Kay 2020: 131), material exchange and mutual interference at the hominin-hyena interface have the capacity to promulgate lived multispecies communities transcending hominin-exclusive domains and behavioral registers and therefore potentially indicate non-analogue practices of transspecies 'kin-making'.
Returning to the LRJ, another observation on MIS 3 hominin-hyena entanglements is interesting. As noted by Discamps (2009), there is an interesting increase in hyena-bearing sites towards the end of MIS 3, both paleontological and archaeological, and different complementary lines of evidence point to a marked demographic expansion of Western European hyena populations in this timeframea 'Hyena event'. Even though hyenas arrived in Britain already during the Eemian (Varela et al. 2010) and earlier (Turner 2009), the remarkable cluster of British LRJ sites, many of which yield 'type three' signatures and represent short-term occupations in or close to hyena dens with only a handful of lithic artefacts, mostly hunting implements and points (see Table 3), may be regarded as a reflection of LRJ hominins tracking hyena landscapes, and thus as indicative of the increasing abundance of hyenas in this area during the second part of MIS 3. Put more provocatively, irrespectively of whether Neanderthals or other Late Pleistocene hominins including potentially contemporaneous IUP H. sapiens groups were the bearers of this techno-complex (cf., Hajdinjak et al. 2021;Prüfer et al. 2021), the LRJ then possibly delineates the crystallization of longterm, high-mobility adaptations to hyena environments already prefigured in some Late Middle Paleolithic contexts. Re-formulated in multispecies terms, the LRJ may give testimony to a derived archaeological signature of being-with and becoming-with hyena others in the open and vast landscapes of Northern Europe. The LRJ may then circumscribe a mutually constructed nicheor a multispecies 'meshwork' to take up Ingold's (2013) formulationarticulating large-scale sociomaterial consequences of Late Pleistocene hominin-hyena coordination and nice construction (see esp. Hussain and Will 2021 for the possibility of multispecies niche construction). At the very least, it is certainly worthwhile to reconsider the broad geographic correspondence between the larger LRJ phenomenon and the North European Plain which has likely acted as a major landscape attractor and mobility catalyst for Late Pleistocene hyenas. As a final consideration, it is worth returning to the earlier mentioned circumstance that some Middle Paleolithic sites where both hominins and hyenas acted as formative agents (esp. 'type two' and 'type three' sites), especially in France, Italy and Spain (Diedrich 2014: Table 1), have also yielded isolated hominin teeth or other skeletal parts. Traditionally, it has been assumed that these hominin remains are intermingled with carnivore-derived deposits due to taphonomic processes, hominins falling prey to predators, for example as a result of 'confrontational scavenging' (Bunn and Kroll 1986;Baynes-Rock 2015b) or succumbing otherwise and then being transported to nonhuman occupation sites by various carnivores (cf., Camarós et al. 2016), and/or, finally, that the palimpsest character of the archaeological sediments makes it unlikely that the encountered hominin remains are contemporary with the activities of hyenas and other carnivores in the first place (e.g., Villa and Soressi 2000). Some hominin bones further exhibit carnivore modification marks and some even yield evidence for digestion by hyenas as seen at Les Pradelles (cf., Maureille et al. 2017), which is taken to lend support to the cited scenarios. All of these points are valid considerations but given the scarcity of formal, well-defined burial contexts in the European Middle Paleolithic (Pettitt 2011) and following the previous arguments for socioecological immediacy and a broader reciprocal 'give-and-take' logic underpinning Neanderthal-nature relationsabove framed under the theoretical umbrella of generalized multispecies exchange relations (pace Nadasdy 2007) -other scenarios emanating from the situated sensitivities and animal-directed concerns of multispecies life should also be considered.
It is for instance possible that Neanderthals and other hominins engaged in what may sensibly be termed 'ecological' funerary practices, in which the deceased were offered to specific nonhuman co-inhabitants to be scavenged and ultimately recycledbroadly analogous to Tibetian sky burials and related cultural practices already described by anthropologists (Martin 1996). It should be noted that Neanderthal 'funerary caching' (Mussi 1988: 99;Pettitt 2011: 92) is not inconsistent with this scenario (cf., Pettitt 2018), especially if we accept hyenas as important agents of hominin landscape enculturation, and given that the animals' ecosystem role is tied to recycling key nutrients in bones (Abraham et al. 2021) and, generally speaking, transforming bodiesi.e., their agency is phenomenologically linked to the transposition of corpses and takes part in the wider ecological chaîne opératoire of bodily forms, thus fundamentally regulating and curating Middle Paleolithic 'bodyscapes'. Isolated hominin remains at carnivore sites may accordingly attest to the deliberate fragmentation of hominin bodies linked to the deposition of some body parts, as discussed in the context of Les Pradelles and similar to the meaningful placement of stone artefacts considered above, at liminal places in the landscape where the relationship between human and nonhuman was at stake and negotiative action was required. And, lastly, some of these scattered remains may simply give testimony to the hazardous nature of venturing into the territories and homes of carnivores, yet not merely as deprived evidence for a 'nasty, brutish and short' life (pace Hobbes) but as part of ongoing efforts to relate to and address significant nonhuman co-inhabitants.

Bears: Slumbering companions walking between worlds
The so far developed perspectives on the significance of carnivore others, especially hyenas, can be complemented by a brief foray into Neanderthal relationships with bears. Cave bear remains are frequently encountered at Neanderthal sites across Eurasiaeven though many instances are interpreted as alternating, non-related occupations (Patou 1988;Stiner 1999) -and they co-occur with hyena assemblages in many important 'type two' and 'type three' sites (Hoffecker and Cleghorn 2000;Viranta and Grandal d'Anglade 2012;Skinner 2012). Interestingly, there is a marked east-southwest gradient, with cave bears being more dominant and consistently entangled with Middle Paleolithic sites in Central, East-Central and Southeastern Europe (cf., Daujeard et al. 2016;Romandini et al. 2018: Fig. 1), while hyenas seem to have been the primary spelaeoid interlocutors for hominins in Western and probably Southwestern Europe (cf., Jimenez et al. 2021;Fig. 4; see Table 3). This geographic division is probably linked to the characteristic heterogeneous relief of East-Central Europe and the circumstance that many of these areas retained a noteworthy forest or forest-steppe component and even acted as cryptic refugia for various tree species during glacial conditions, especially below 45 • N and for example in the Balkans (Birks and Willis, 2008;Tzedakis et al. 2013;Ravnsbaek Holm and Svenning 2014). The boreal refugia and mountainous environments of Southern Europe, notably Italy and Austria, probably played a similarly conducive role for hominin-bear encounters and interactions.
This division between Western and Central/Eastern Europe coincides with bear palaeogeography: in Western Europe, Neanderthals mainly had to deal with the common cave bear, U. spelaeus, whereas Central European and Eastern Neanderthals shared their environment with the largest and most powerful Late Pleistocene bears, U. ingressus and U. arctos 'priscus' (cf., Baca et al. 2014;Marciszak et al. 2019;Marciszak and Lipecki 2020: Fig. 2). This not only suggests divergent conditions of interaction and modes of lifeworld intersection but also raises the interesting possibility that geographically distinct bear assemblages coevolved with different Neanderthal subpopulations reflected in genetic  Table 3 (site numbers correspond to site IDs in  Table 3; Zaskalnaya VI and Matuzka are located outside of the shown map section). Colors indicate inferred site category and shape of symbols represent period/ techno-complex contexts (MP = Middle Paleolithic; LRJ = Lincombian-Ranisian-Jerzmanovician). Animal outlines from http://phylopic.org/ via the Public Domain. (Dalén et al. 2012; but see Hajdinjak et al. 2018) and lithic technological data (Ruebens 2013). Such structured cogeographies have been demonstrated for contemporary First Nation groups and grizzly bears in Western Canada (Henson et al. 2021), indicating that foragers and bears often have aligned needs in terms of space, food and other resources and are thus drawn to the same habitats, so that bears easily emerge as cardinal nodes in integrated human socio-ecological systems.
In contrast to hyenas who tend to be gregarious, inquisitive and resource-competitive, cave bears are unlikely to have assumed a role as tangible and physically contiguous, 'apresent' neighbors (sensu Schütz 1993) and encounters between bears and hominins were likely framed by specific situations and bound to particular locales. As shown by occasional evidence of Middle Paleolihic encounter hunting (e.g., Münzel and Conard 2004) cave bears and Late Pleistocene hominins were drawn together in karstic environments (Daujeard et al. 2016) and potentially interfered with one another while seeking shelter and, in the case of spelaeoid bears, places for hibernation. Based on archaeozoological evidence from Balve cave, Kindler (2012) has suggested that Neanderthals must have developed a habit for scouting and perhaps even clearing caves before establishing their own occupations. But Neanderthals also visited cave bear retreats independently of occupational concerns, sometimes in remote places such as high-up in the Alps (Richter 2018), and bequeathed isolated hominin cut-marks on bear bones (e.g., Martínez-Sánchez et al. 2012). In addition, earlier interglacial Neanderthals recurrently, and sometimes systematically, 'harvested' cave bears close to hibernation sites, in wetland settings and/or at prominent river confluences, and took their fur and perhaps meat in addition to high-value consumables such as the tongue (Wragg Sykes 2020: 158). This targeted hunting pattern is for example documented at Taubach in Germany (Bratlund 1999), Biache-Saint-Vaast in France (Auguste 1992) and Krapina in Croatia (Miracle 2007) and aligns with ungulate predation tactics discussed before.
In Taubach, bear skulls are particularly numerous and Neanderthals also took special interest in the paws of the animals (Bratlund 1999). In contrast to what might be expected, U. arctos bones yielded most cutmarks at Biache and Taubach (Auguste 1992;Bratlund 1999;cf., Romandini et al. 2018: 87) and this suggests that Neanderthals to a lesser extent took advantage of the marked 'homing behavior' and den fidelity of spelaeoid bears but instead preferred to intercept large carnivorous brown bears in opportunistic foraging situations or when attempting to hibernate close to cave entrances (and not deep inside cave bodies; cf., Marciszak and Lipecki 2020: 888). This is consistent with Auguste's (1995) observations regarding the Biache material, where cubs are generally missing and the age and sex distribution of killed bears supports an interception scenario of solitary individuals, rather than hominin stalking of hibernating ursids in underground cavities. The clear processing focus on cranial bones and especially mandibles, jaws and paws derived from prime-aged bears (cf., Bratlund 1999: 118) seems hardly coincidental then, andsupports the view that Neanderthals were also being concerned with 'harnessing' specific behavioral and foraging qualities liked to these body parts, which clearly define brown bears as foragers. This possibly indicates a general concept of character transposition and trait fluidity at the Neanderthal-animal interface, which is well-documented among contemporary and historic hunter-gatherer societies and frequently provides a fundamental ontological rationale for the selection, transformation and utilization of specific zoomaterials (Hill 2012(Hill , 2019Qu 2017).
This pattern of targeted bear exploitation with an emphasis on MIS 5 contrasts with Neanderthal-ursid engagements in later Middle Paleolithic contexts, especially in the Italian Alps of MIS 3, where hominins seem to have conducted foraging trips to mountainous and sometimes high-altitude bear denning locales and indiscriminately processed hibernating spelaeoid bears of all age classes during winter and spring (Romandini et al. 2018;Wragg Sykes 2020: 158). These alpine contexts, comprising sites like Fumane, Rio Secco, Generosa Cave and Cio Ciara, document a more integrated foraging strategy where rugged terrain prey such as ibex and deer complement opportunistic but nonetheless intense bear exploitation of entire carcasses including the fatty chest, limb parts and also bone marrow extraction, with occasional evidence for the transport of high-value meat-bearing parts between sites (Wragg Sykes 2020: 157-158). The responsible Neanderthals thus clearly took advantage of the seasonal schedule and rhythm of characteristic cave bear 'homing behavior' (Marciszak and Lipecki 2020: 888), rendering ursids a seasonally predictable resource.
Yet bear hideouts and hibernation dens in or at the edge of mountainous landscapes were not only foraging grounds in the Late Middle Paleolithic, they also framed multispecies 'contact zones' with their own distinct material affordances and possibilities. They offered rich, natural bone caches with a raw material potential of their own, and hominins sometimes manipulated and dislocated skeletal material from these bear-derived bone beds (Romandini et al. 2018). The recently discovered cave bear bone retouchers from Scladina (Abrams et al. 2014), Rio Secco (Romandini et al. 2018) and Fumane (Jéquier et al. 2012) may be regarded as 'emergent materialities' of this larger interspecies constellation, in which hominin-bear relationships were negotiated at specific localities and situations in a wider 'hunter's landscape' (pace Ogden 2011; Fig. 3). The sequentially incised cave bear bone recovered from the hyena and cave bear-dominated sediments of Pešturina cave in modern-day Serbia can similarly be interpreted as such situational material engagement linked to special bear-endowed places in the wider landscape (Majkić et al. 2018). In the Northern Caucasus, the role of cave bear-invested places may even be reflected in the intermediate or 'transitory' position of sites such as Matuzka in larger Neanderthal landuse systems, occupying a key nodal position between lower and higher elevations (Hoffecker and Cleghorn 2000). Matuzka has yielded only a small number of Middle Paleolithic stone artefacts within otherwise cave bear-dominated sediments and its archaeology is most parsimoniously interpreted as a reflection of multiple recurrent but short-term hominin stays (Hoffecker and Cleghorn 2000: 372, Fig. 2). Some bear localities at the periphery of Neanderthal core territories thus likely developed into ephemeral but persistant places (pace Maher 2021) or integral waypoints in seasonally differentiated cultural landscapes of the terminal Middle Paleolithic. After all, landscape is always, as Crumley (1994) has pointed out, 'an array of lived experiences made manifest' (Maher 2021: 249) where hominin interference with space and its inhabitants contributes to the making of 'place'.
The importance of ursid places in Neanderthal landscapes may also be gleaned from Middle Paleolithic burial practices. As outlined by Camarós et al. (2017), the Neanderthal burial of La Chapelle-aux-Saint in Southwestern France may provide tentative evidence for the utilization of refurbished cave bear hibernation pits. The heated controversy on the intentional character of Neanderthal burials (Rendu et al. 2014(Rendu et al. , 2016Dibble et al. 2015) may then, in some instances at least, beg the actual question at hand and only detract attention from the 'hybrid' quality of the burials as partly anthropogenic and partly zoogenic. Following Hill (2000), such interments can be regarded as vibrant 'microcosms' of lived human-animal relations, and they powerfully counter received Western bifurcations of 'nature' and 'culture' (or the supposed cognitive and cultural implications of formal vs. informal burials). As Pettitt (2015) has already noted, Neanderthal-type interments are hardly intelligible as proper 'ritual burials', but instead appear to signal a set of cultural practices for which the fabrication and affirmation of proximity to the surrounding land was imperative. Such 'ecological' funerary behavior embedded in multispecies geographies and tied to specific more-than-human places must be examined as born out of the conditions and dynamics of multispecies encounter and interaction rather than merely signposting their aftereffects. Material fragmentation and the disarticulation of bodies was an integral part of these Neanderthal practices of relating and they may be understood, as proposed above, as part of broader efforts to negotiate familiarity and intimacy via mimicry and re-encactmentcommunicative modalities that often play a key role in how contemporary hunter-gatherers address a range of key actors in their environments including other animal species (e.g., Lewis 2014). Although the co-optation of bear-dug structures by Middle Paleolithic hominins remains somewhat speculative until more supportive evidence is recovered, bear places themselves also frame some Neanderthal interments. At Regourdou, for example, a Neanderthal burial is associated with a natural, albeit catastrophic denning profile of brown bears and three cut-marked bear bones indicate at best sporadic hominin visits to an otherwise bear-vested locale (Cavanhié 2011), thus encouraging the interpretation that the placement of the deceased into bear country was not fortuitous.

A new narrative?
Multispecies theory sets the stage for new narratives and alternative understandings of early human evolution and the making of our species. Patterns and trajectories of hominin becoming are often the product of mutual and ongoing co-shapings of humans and nonhumans, and animal others have played a key but understudied role in this process. We have tried to argue here that much of the archaeological evidence traditionally deployed to reconstruct hominin subsistence behavior or the symbolic and cognitive capacities of hominins in fact presents a rich corpus for rethinking interspecies relationships in deep prehistory. The resulting redressing of hominin-animal interactions, through the prism of integrative multispecies theory and interdisciplinary anthrozoology, draws attention to the reciprocal interweaving of hominin and animal behaviors and the productivity of the emerging, often co-adaptive interfaces in modulating behavioral constraints and opportunities, broader land-use and mobility patterns, and signatures of material culture. Other-than-H. sapiens material culture which references animals, such as deliberately modified and decorated animal bones or extracted and then co-opted animal body parts such as claws, feathers and limb bones, thereby actively takes part in the construction and continuous negotiation of interspecies relations. These instances of material culture, rather than indexing abstract symbolic or cognitive deus-ex-machina qualities, are consequently imbricated in the very hominin-animal interactions they shape, affirm and reproducethey do not stand 'outside' of or transcend these relationships and their study is thus essential.
Making and handling material culture is a mode of relating in itself, and the archaeological challenge is to reinsert different material expressions of the archaeological past back into their multispecies 'meshworks' (sensu Ingold 2013). Deliberately incised bones of corvids, hyenas and giant deer (Leder et al. 2021) or the suspended or otherwise worn eagle and owl talons of Eurasian Neanderthals then first and foremost attest to the socioecological and phenomenological significance of concrete lifeworld intersections between the involved hominins and the addressed animal others. There is no need to account for this material culture in terms of a priori biocognitive determinisms, prioritizing inbuilt capacities, extended phenotypes or fossil minds. To the contrary, such animal-imbued material culture can be understood and contextualized by taking stock of the tangling of behavior and lived experience at prehistorically situated hominin-animal interfaces, precarious or not, and by recognizing that different types of interspecies neighborhoods and modes of landscape sharing can modulate particular 'horizons of concern' and sociomaterial practices on both sides. These social practices of course have cognitive and aesthetic extensions, but not the other way around.
This reframing of hominin-animal relationships not only suggests that animals such as raptors, hyenas and bears actively shaped, and thus co-defined, the lifeways and material cultures of Neanderthals and other Late Pleistocene hominins, it also, and perhaps more importantly, deconstructs received dualist evolutionary narratives, which too often divide the human story into an earlier, ancestral stage 'poor' in culture and mind and a later, derived stage 'rich' in both of them. The herepresented multispecies exploration, conversely, suggests that the Neanderthal world was deeply, and probably from the beginning, impregnated with culture, expansive sociality and mind, and interspecies relations were an important locus for all of them. Nevertheless, Neanderthals probably lived in differently constituted 'naturecultures' when compared to later Upper Paleolithic H. sapiens, and hominin modes of engaging, living and becoming-with other animals were generally subjected to substantial evolutionary and contingent historical change. The various issues and questions raised by at least a decade of heated paleoanthropological research and debate on behavioral modernity and complexity and their many extensions and sidediscussionseven though having certainly passed their heyday nowhave thus unfortunately resulted in a misplaced concern with 'origins', rather than fostering the development of a comparative and deeply contextual approach to the shifting variables of hominin culture and society, including its anchoring interspecies nexus. What it means to be a Neanderthal in different times and places can only be answered by integrated ecocultural approaches confronting hominin and nonhuman realities, productions and concerns, and such analysis needs to make space for non-analogue variation in cultural behavior and interspecies dynamics to avoid the pitfalls of sapienscentrism. Responding to Slimak's (2019) timely call for a 'cultural anthropology' and serious comparative 'ethology' of Neanderthals therefore requires to develop new contextual approaches that belabor the multispecies conditions of Neanderthal life, culture and materiality.
This lurking synthesis of 'ethnographic' and 'ethological' aspirations in Neanderthal archaeologyin sync with Lestel's (2006) call to arms in animal studiesrequires taking on the broad picture, and to also rechart hominin-herbivore interfaces in more-than-economic terms. A privileged starting point would be Neanderthal-elephant relations since elephants were, except for a few cases, probably opportunistically taken and scavenged (Uthmeier 2006;Smith 2015), yet their skulls were nonetheless transported to Neanderthal sites, and in Spy perhaps used to structure hominin space (Wragg Sykes 2020: 146-147), while elephant ribs were sometimes transformed into hammock-like pointed tools or points (Gaudzinski et al. 2005). The role of reindeer in the Quina Mousterian, briefly mentioned above, is another obvious locus of multispecies investigation in the Middle Paleolithic and might shed new light on the extent and nature of Neanderthal reindeer adaptations. More generally, it is striking how particular bones of selected animal taxa were used and transformed in particular ways (Wragg Sykes 2020) and this implicit normativity of raw material selection and transformationfor example the strong entanglement of bovid bones with lissoirs  or the link between cervid femura and organic retouchers (Jéquier et al. 2015;Castel et al. 2017;Costamagno et al. 2018b) -is certainly indicative of lived concerns and interspecies relations embedded with meaning. These cases illustrate that not only carnivores but other animals as well co-structured the behavior and material culture of European Neanderthals, and that these hominins, in turn, maintained intimate, phenomenologically-informed and culturally consequential relationships with a broad range of nonhuman others. This not only demonstrates that Neanderthals lived in a vibrant ecology of relations but warrants further anthropologically-inspired investigations into the motivations and ontological ramifications of coopting particular animal parts as tools and ornaments, and not others (see Hussain and Breyer 2017;Hill 2019; see above).
The Late Pleistocene is arguably a privileged arena for investigating the material consequences of multispecies relationships because ecological and faunal communities experienced severe continual reorganizations and the archaeological record documents major changes in hominin lifeways and culture from Western Eurasia to Southeast Asia and into Sahul (Roebroeks and Soressi 2016), most notably the emergence of representational imagery, which in Europe and Oceania centers on animal motifs (Bourrillon and White 2015;Brumm et al. 2021). An explicit multispecies framing of hominin-animal interstices in the Late Pleistocene helps to throw these consequential developments into novel perspective. Faunal and taphonomic evidence from the site of Les Cottés in Western France covering the Middle-to-Upper Paleolithic transition has been interpreted as indicating changing expositions between hominins and carnivores through timefrom 'shared access' to sites and bones in the Middle Paleolithic to the increasing 'exclusion' of other carnivores from hominin occupations and their ultimate demise during the Early Upper Paleolithic . Similar trajectories of hominin-carnivore dynamics have been reported from other key stratigraphies in Central and Western Europe (e.g., Castaños 2005;Davis et al. 2007;Arsuaga et al. 2011;Enloe 2012;Conard et al. 2012;Kitagawa et al. 2012;Kitagawa 2014;Bertacchi et al. 2021). The periods following the Middle-to-Upper Paleolithic transition in Central and Western Europe are also associated with an increasing grasp of hominins on their surrounding landscape, resulting in more extensive settlements and a growing anthropogenic fingerprint (Baumann et al. 2020;Bocherens 2018), indicating changing modes and logics of hominin landscaping, place-making and ecosystem impact (cf., .
The Middle-to-Upper Paleolithic transition additionally marks a tipping point with regard to the systematic exploitation of animal carcasses anchored in the transformation or co-optation of nearly all animal body parts and animal-furnished raw materials, including previously less utilized materials such as teeth, antlers and ivorya development described by Richter (2018: 202) as preempting a momentous 'bone technique revolution' in later MIS 3. This development may imply, or lay the foundations of, a fundamental rupture with obligatory and facultative scavengers co-inhabiting the landscape with Paleolithic hominins, and thus might have changed the role and significance of hominins as carcass accumulators for these other animals, perhaps most notably for bone-cracking hyaenids. Later hominins, most likely H. sapiens populations, laid claim to the totality of hunting yields and in this way substantially reduced the sense of co-ownership and sharing with other carnivores, possibly even promoting 'amensalism' (negative association). Given the strong emphasis on Neanderthal-carnivore competition and antagonism in the expert literature, it is somewhat ironic that it might have been this essentially post-Middle Paleolithic development largely tied to H. sapiens practices that possibly tipped the scales towards acute resource antagonism, and hence subverting the fragile but sympatric hominin-carnivore intersections that have evolved over thousands of years. If incipient H. sapiens-constructed cultural niches comprised domestic dogs, as recently proposed by Shipman (2015aShipman ( , 2015b, this would have introduced yet another distance-promoting and 'horizon of concern'-shifting element with potentially cascading effects for hominin-carnivore ecosystem expositions. Another long-term trajectory in the evolution of hominin lifestyles and nature relations is associated with the Late Pleistocene drawing to a close: while archaeological evidence for 'delayed-return' foraging economies (sensu Woodburn 1980Woodburn , 1982 remains largely absent from the Middle Paleolithic record, some Upper Paleolithic societies document the development of food storage practices (Soffer 1985;Soffer et al. 1996;Soulier and Costamagno 2018), anticipatory lithic blank caching or even 'hoarding' (Angevin and Langlais 2009), the expansive labor-intensive investment into delayed-return technologies including organic tools with extended life history (Liolios 2006;Bon 2015) and perhaps socially-controlled access to space, resources, knowledge and skill (Pelegrin 2013;Julien and Karlin 2014;Langlais 2020). Although these landmark developments seem to be staggered in time and do not emerge as a single coherent package, they nonetheless signal a timetransgressive, albeit not necessarily irreversible, departure from strict 'immediate-return' systems of ecocultural organization (pace Woodburn 1980; cf. Testart 2012) which have charcaterized much of the European Middle Paleolithic. This shift may be epochal as Bird-David (2006) has developed a range of convincing arguments suggesting that especially delayed-return foragers, for example the many contemporary Indigenous societies of the circumpolar North (Morphy 1999;Plumet 2006), are prone to develop and invest into elaborate animal-depicting visual culture, while their present-day immediate-return counterparts tend to exhibit reduced interest in such representational imagery.
This relationship does not reference some mysterious anthropological universal, however, but rather appears to be a result of the specific interplay of economy and sociality in these communities and the proximity-dissolving consequences of delayed-return foraging systems, in which immediacy and co-presence are not regulative ideas or world-making principles anymore. In elaborate art traditions which are in many cases also characterized by the sophisticated decorative practicesrepresentational and non-representationalwith regard to everyday objects, the visualization of 'others' including animals serves in part as a means to engage, evoke or address these others and to produce immediacy, adjacency and social proximity in the first place (Bird-David 2006). A key function of material culture bearing figurative images, then, is often to re-enact the agency of the depicted beings in order to foster pluripresence and communication. For immediate-return foragers, by contrast, there is often no acute need to invest in such social technologies or cultural techniques that establish copresence with their communities of significant nonhuman others because pluripresence is safeguarded and continuously reproduced by their very way of living. Against this borader background of human-nature relations and their cultural consequences, we propose that some carnivoresespecially hyenas, raptors and bearsneed to be considered socially significant others for Late Pleistocene hominins, and Neanderthals in particular seem to have invested material and cultural resources into forging reciprocal ties with these animals. We suggest that this is not just indicative of the ecological status of Neanderthals in the larger carnivore guilds of their time, but points to the phenomenological and societypermeating significance of animals with whom they shared, and regularly intersected in, the landscape. Being a carnivore and relating to carnivore others was likely a cultural concern for Neanderthals, and the biocultural evolution of these hominins must thus probably be understood in relation to broader patterns of evolving carnivore ecologies. The ecocultural entanglement of Neanderthals, hyenas and raptors across the vast expanses of the Eurasian mammoth steppes can therefore arguably be recognizedechoing Stewart's (2005Stewart's ( , 2007 seminal ecological arguments cited in the beginning of this paperas a coadapted community of on-par carnivore neighbors and interlocutors, stabilized by particular material and sociocultural practices of engagement, attunement, recognition and affirmation. In this view, the demise of Late Pleistocene scavenger-rich ecological communities in which Neanderthals actively participated is perhaps not only a byproduct of global dynamics of Late Quaternary megafaunal extinctions, but was possibly also fueled and accelerated by the emergence of cultural behaviors violating the balance of sharing and disrupting a sensitive matrix of co-adapted and carefully coordinated behaviors at the dawn of the Upper Paleolithicundermining a broader Middle Paleolithic carnivore 'commons' (pace Baynes-Rock 2013).
Interestinglyand even though the archaeological and palaeogenetic evidence on the likely hominin bearers of the Northwestern and Central European LRJ is likely to tip in favor of early H. sapiens populations rather than late Neanderthals (or show that both demes contributed to its formation) -the LRJ and other 'transitional' or 'Initial' Upper Paleolithic complexes such as the Bachokirian (Hublin et al. 2020), the Bohunician (Svoboda 2003;Valoch 2013b) and possibly the Western European Châtelperronian (Rios-Garaizar et al. 2012;Discamps et al. 2019) seem to showcase a typical Middle Paleolithic signature when carnivore faunal fingerprints, site formation histories and relationships are concerned. This in turn may suggest that the significance of carnivores for Paleolithic societies was not primarily a question of hominin taxonomy and hominin-carnivore correlations, and that the latter were possibly continuous across the Middle Palaolithic-IUP transition. Yet, the reorganization of hominin societies precipitated by the developments between ca. 60 and 40 kya and eventually leading to the fully developed Upper Paleolithic still marks a turning point in the deep prehistory of Western Eurasia. Not only in terms of long-term material culture trajectories but equally so in the evolution of hominin-carnivore interactionsand thus likely heralding shifting regime dynamics of ecosystem assembly and hominin as well as carnivore roles therein, linked to both changing ecoclimatic conditions and hominin technological and sociocultural practices.
The discovery of 'recent Neanderthal ancestry' in incipient H. sapiens populations associated with the IUP from Bacho Kiro (Hajdinjak et al. 2021) certainly adds to this emerging picture of a complex mosaic of continuity and rupture at the Middle-to-Upper Paleolithic juncture, and, instead of reassuring untenable Neanderthal-H. sapiens polarities, points to overlapping lifeworlds and cultural logics supporting similar carnivore relationshipsa contestation perhaps indirectly supported by the emerging aDNA picture, according to which Europe between 45 and 40 kya was populated by more-or-less Neanderthal-like populations rather than genetically distinct Neanderthal and H. sapiens subpopulations (Fu et al. 2014(Fu et al. , 2015Prüfer et al. 2021). The pierced bear teeth recovered from the IUP layers of Bacho Kiro cave (Hublin et al. 2020) at least seems to continue a trajectory of intensification, and perhaps diversification, in hominin-bear relationships already prefigured in the later Middle Paleolithic.

Conclusions
This paper has taken some initial steps to unravel the 'humanimal' dynamics and conditions of later hominin evolution (pace Haraway 2007) beyond the confines of H. sapiens prehistory. We have shown that the deep-time evidence harnessed by Paleolithic archaeologists is by no means mute about the involvement of other animals in the making of hominin cultures and livelihoods, and suggests that perhaps especially carnivores have played a salient role in long-term human-nature negotiations among Neanderthals and other Late Pleistocene hominins with elevated carnivory. We have drawn on macroarchaeological evidence on the patterned associations and intersections of MIS 5-3 hominins and carnivore others in Western Eurasia to argue that engaging and addressing nonhuman carnivore others, especially hyenas but also large raptors and some bears, was an important dimension of cultural niche construction and world-making among pre-Upper Paleolithic societies, underscoring the lived significance of finding a place in the larger carnivore community. Carnivores thus actively participated in Neanderthal societies and relating to them was a fundamental sociological, and thus more-than-economic concern. The redressing of hominincarnivore interaction attempted here therefore suggests that sympatry was likely a foundational pillar of Neanderthal-carnivore interaction.
Contrary to common views and previous interpretations in (Middle) Paleolithic archaeology, we have proposed that the competitive exclusion of other carnivores and their increasing detachment from hominin's immediate 'horizons of concern' was bound to a fundamental regime shift in hominin-carnivore relations at the passage from the terminal Middle Paleolithic to the developed Upper Paleolithic. We raise the possibility that a new set of hominin cultural behaviors, in particular the aspiration to claim total prey carcasses for supporting a diverse portfolio of organic tools and visual cultures, eventually undermined co-evolved socially and materially productive ties with other carnivores and thereby foreshadowed the ultimate extinction of some species, notably obligatory scavengers adapted to hominin carrion landscapes, in the course of the Pleistocene. Our multispecies exploration further suggests that the development of animal-themed visual culture and figurative art in the Upper Paleolithic may have less to do with so-called 'modern human behavior' than with the conditions and situated logics of humananimal relationships and the (changing) role and contribution of nonhuman animals to hominin life, material culture and society.
Taken together, this paper cautions against the image of the ecologically 'authentic' carnivore, as other carnivores have lived in close neighborhood to hominins for over 3 to 4 million years and thus had plenty of opportunity to adapt to various hominin-shaped and inhabited landscapes. Hominins alike are too easily dismissed (or overlooked) as past ecosystem agents, even though their cultural practices and behavioral proclivities can profoundly modulate ecosystem dynamics. Hominins and carnivores have shaped each other for thousands of years and the resulting behaviors, adaptations and dispositions must be considered a shared yet contested evolutionary legacy. Reconsidering Neanderthals and other hominins in the light of multispecies theory may unlock and clarify some of these legacies and opens up new windows into sociocultural coordinates of Pleistocene life.

Declaration of Competing Interest
The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper.

Acknowledgements
This paper has a history of co-becoming of its own. We wish to express our gratitude to our students, who, through their enthusiasm and inspiration, have contributed to the completion of this paper. We thank our home institutions and affiliate research groups for providing intellectually stimulating and safeguarded environments. This paper is dedicated to the recognition of our multispecies past and its many legacies as well as the contemporary conservation of hyenas, raptors and bears.

Funding information
STH acknowledges funding through the CLIOARCH project supported by the European Research Council under the Horizon 2020 programme (grant agreement No. 817564). TKN received funding from the Independent Research Fund Denmark (case number 9062-00027B).