Human-beaver cohabitation in the Early and Mid-Holocene of Northern Europe: Re-visiting Mesolithic material culture and ecology through a multispecies lens

Macro-patterns in zooarchaeological beaver assemblages

In contrast to the Terminal Pleistocene of higher latitude Europe where beaver remains are rarely encountered in archaeological contexts (cf. Veil et al., 2012; Weber et al., 2011: 661), the Eurasian beaver becomes an important and recurrently encountered component of faunal assemblages from the Early and Mid-Holocene of Northern Europe (Figure 1 and Supplementary Information M, available online). We summarize the available zooarchaeological data in relation to the three delineated macro-regions, paying particular attention to changing beaver abundances and faunal co-associations.

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Figure 1.

Overview of key sites from the Northern Mesolithic and associated beaver remains as well as beaver-related material culture: (a) Early Mesolithic; (b) Middle Mesolithic; (c) Late Mesolithic/Early Neolithic. Data provided in Supplementary Information 1, available online.

Northeastern Europe

The Early and Middle Mesolithic in the Baltic and the Russian West are represented by the Early Kunda, Veretje and earlier Butovo complexes (Damlien, 2016; Hartz et al., 2010; Manninen et al., 2021; Zhilin, 1996), while the Late Mesolithic/Early Mesolithic comprises Late Kunda, later Butovo and Janislawice as well as the regional variants of the emerging Early Comb Ware complex (Oras et al., 2017; Piezonka, 2021; Zvelebil, 1994). Site types and conditions of archaeological recovery vary greatly between and within regions. In the earlier Mesolithic, sites are mainly located above river and lake terraces or in extended wetlands and are often represented by small flint scatters or mixed deposits, while in the later part of the Mesolithic settlement systems become more structured and a duality between habitation and extraction sites as well as between coastal and inland occupations begins to take shape (Piezonka, 2021). For the Russian West, well-preserved faunal assemblages are mainly associated with the many bog and wetland sites of the Volga-Oka interfluve (Zhilin, 1996, 2004, 2007, 2014b), and the record may thus be biased towards specific localities and forager activities in the landscape.

A directly dated beaver-gnawed piece of wood from the important Ivanovskoye peat bog dated to around 10,000 years ago (Zhilin, 2019) provides evidence for prolonged histories of human-beaver co-residence in the Russian Northwest. In the Early and Middle Mesolithic, beaver makes up between 5% and 47% of total mammalian NISP in the Baltic and between 19% and 36% in Western Russia, while the Late Mesolithic/Early Neolithic is characterized by beaver NISP frequencies between 3% and 61% in the Baltic and between 9% and 34% in Western Russia (Figure 2). In the Early Mesolithic of Western Russia and the Baltic, beaver is often the second-most frequent mammal after the elk, pointing to the beaver’s economic relevance and, possibly, elevated abundance in the ecosystem. Muskrats (Ondatra zibethicus) and martens (Martes sp.) are also an important faunal component in this spatiotemporal context.

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Figure 2.

Number of identified mammal specimens (NISP%) from the Baltic countries and Western Russia from the Early to the beginning of the Mid-Holocene: (a) Early Mesolithic; (b) Middle Mesolithic; (c) Late Mesolithic/Early Neolithic). Horizontal bars represent median values with standard deviations. Animal silhouettes have been redrawn from https://www.phylopic.org/. Raw data are provided in Supplementary Information M Tables 1 and 2, available online.

From the Middle Mesolithic onwards, the importance of elk diminishes in the wider area, while wild boar and red deer (Cervus elaphus) become increasingly important in the Baltic and the water vole (Arvicola amphibius) in Western Russia. Interestingly, the increasing importance of water vole in the Northeastern inland Mesolithic, reaching up to 61% of recorded NISP at individual archaeological sites, corresponds to a general trend of decreasing faunal representation of the beaver during the Middle and Late Mesolithic of the region, and this may be related to long-term beaver-modulated changes in riverine-lakeland habitats, strongly promoting water vole populations on a local scale.

In the Baltic, beaver frequencies are overall less stable across archaeological sites and the pattern is more punctuated than in Western Russia: some sites harbour fairly low beaver abundances (c. 3–7% of NISP), while others exhibit increased beaver yields (c. >20–60%), pointing to functional differences in site formation and perhaps more diverse human-beaver interactions. The Late Mesolithic/Early Neolithic site of Dąbki 9 in Northern Poland, for example, has been argued to represent a seasonal special purpose locality mainly geared towards fur-extraction and possibly exchange (Schmölcke and Nikulina, 2015). It is notable that among the other species from Dąbki 9, the otter makes up c. 6% of the documented mammalian NISP, thus representing the third-most frequent animal in the whole faunal assemblage. Although otter remains are not generally frequent at beaver-bearing sites, they are often a common yet low-abundance feature. This association is conspicuous since otters and beavers are known for their sympatric relationships, as otters benefit from beaver-engineered and disturbed habitats, especially in riverine higher latitude woodland environments (LeBlanc et al., 2007; Reid, 1984; Tumlison et al., 1982).

The possible presence of muskrat (NISP = 28%) alongside beavers (NISP = 30%) in Early Mesolithic Stanovoye 4 at the Volga-Oka interfluve in Northwestern Russia (Zhilin, 2004) is notable and may similarly indicate that human foragers took advantage of the facilitative effects of beavers on their animal co-inhabitants (cf. Crego et al., 2016).

Northwestern Europe

The Early and Middle Mesolithic of Southern Scandinavia is characterized by the Maglemose complex (9500–6400 BC) and the earlier part of Kongemose (6800–5400 BC), known in particular from the many well-preserved bog sites across Denmark and Northern Germany. Two types of sites are known from the Early Mesolithic: so-called deposition sites and habitation sites (Sørensen et al., 2018). Well-preserved faunal material only derives from deposition sites in Southern Scandinavia, while in Northern Germany habitation sites also occur (Sørensen et al., 2018) – likely a consequence of site preservation. The respective lithic and faunal assemblages are often very small and fragmented, indicating short-term visits. The Middle Mesolithic sites from Southern Scandinavia as a whole are well-preserved, derive from wetland deposits, and have mostly been described as habitation sites (‘base camps’: Gramsch, 2000; Groß, 2017; Schuldt, 1961: 185), but we may generally miss the more ephemeral sites from the period. Early and Late Maglemose sites from Jutland and Eastern Denmark show patterned differences in preservation and depositional context (Blankholm, 1996; Nielsen, 2006). In the Netherlands, only a couple of archaeological sites with faunal remains are known from the Early and Middle Mesolithic. The zooarchaeological assemblages are small and beaver remains number only a handful, making the comparison of Northeastern Europe and Southern Scandinavia difficult for the Early Holocene.

In Early Mesolithic Southern Scandinavia, beaver remains make up an average of 8% of total MNI, ranging between 5% and 14% (Figure 3). Most assemblages from this period are very small (<50 total MNI), making it difficult to draw firm conclusions from species compositions. At Sværdborg, where several excavations have yielded larger faunal assemblages, beavers are clearly present (MNI between 5% and 9%), as are other fur-bearing mammals such as otter (5–8%) and badger (Meles meles) (2–5%). In general, otters are relatively common in the Early Mesolithic of Southern Scandinavia (2–33% of MNI), supporting the observations made for Northeastern Europe. Overall, however, there is a greater emphasis on large fauna, such as wild boar and roe deer (Capreolus capreolus). Despite the noted differences in faunal preservation for habitation and deposition sites between Southern Scandinavia and Northern Germany, there is no apparent divergence in beaver MNI representation.

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Figure 3.

Number of identified mammal specimens (NISP%) or minimal number of individuals (MNI%) from Southern Scandinavia and the Dutch wetlands from the Early to the first part of the Mid-Holocene: (a) Early Mesolithic; (b) Middle Mesolithic; (c) Late Mesolithic/Early Neolithic. Note that data for the latter region is only available for the final phase and that MNI is used in (a) and (b). Horizontal bars represent median values with standard deviations. Animal silhouettes have been redrawn from https://www.phylopic.org/. Raw data are provided in Supplementary Information M Tables 3 and 4, available online.

In the Middle Mesolithic faunal assemblages from Southern Scandinavia, beaver remains continue to be present, but in consistently small numbers. They average c. 9% of total mammalian MNI, ranging between 2% and 18% respectively. Larger mammals are again found in greater numbers, but, as noted above, most of the respective assemblages are too small and fragmented to draw broader inter-site conclusions.

The Maglemose and Kongemose complexes are succeeded by the Ertebølle Culture (EBK; 5400–3950 BC), identified first and foremost by its famous coastal shell-midden sites. The EBK represents the end of the Mesolithic in Southern Scandinavia and many sites reflect transitionary phases to the Funnel Beaker Culture (TRB) marked by the appearance of domesticated animals and cereals (cf. e.g. Gron and Sørensen, 2018). EBK zooarchaeological assemblages derive from a variety of site contexts, such as bogs, submerged/waterlogged sites and shell middens, introducing systematic taphonomic and recovery biases (Gron and Robson, 2016). Beavers decrease to an average of only 0.4% of total NISP in EBK, with values ranging between 0% and 2% at individual sites (cf. Figure 3). Other fur-bearing animals, in particular marten and wild cat (Felis silvestris), make up a more substantial part of the faunal assemblages than beavers, while large wild fauna continue to be important. In EBK, marine mammal shares increase significantly in abundance and together with marine fish become a key part of the human diet (Rowley-Conwy, 1999).

In the Netherlands, the Late Mesolithic/Early Neolithic has produced a wealth of faunal data from Swifterbant Culture (ca. 5500–3400 BC) sites in the Dutch wetlands. The Swifterbant Culture has long been considered a transitionary archaeological complex at the forager-farmer interface, but recent evidence demonstrates that from at least 4250 BC on these communities had established agricultural practices while also relying on diverse natural resources (Brusgaard et al., forthcoming; Huisman and Raemaekers, 2014; Raemaekers et al., 2021). Swifterbant Culture sites are all habitation sites, some occupied seasonally and some on a year-around basis. In the Early Neolithic, Linearbandkeramik (LBK) farming communities also inhabited the southernmost zone of what is now the Netherlands, but faunal remains exist from only one site and are very few (eight total identified mammal NISP), so we have to restrict the discussion to the Swifterbant Culture in this region.

Beaver remains are altogether abundant at Swifterbant Culture sites, averaging c. 20% of total NISP, but the numbers are highly variable between sites, ranging between 0.4% and 49%. Beaver remains are predominant at the oldest sites, Hardinxveld-Giessendam Polderweg and De Bruin (5500–4250 BC) (34% and 49% of total NISP, respectively) and Schokland P14 (4900–3300 BC) (37%), where they rival or even outnumber larger ungulates such as wild boar and red deer. At the Hardinxveld-Giessendam sites, the total percentage of beaver NISP masks the notable increase in beavers over time, reaching frequencies of 83% at Polderweg and 51% at De Bruin in the final occupation phases, while the number of large fauna specimens decreases (Oversteegen et al., 2001; van Wijngaarden-Bakker et al., 2001). This is possibly a result of increasingly wet conditions due to sea level rise in the area – conditions in which beaver would have flourished and foragers may have become increasingly reliant on aquatic resources (Brusgaard et al., 2022a).

At the Swifterbant type sites S2, S3 and S4 (4300–4000 BC), where domesticated livestock are also present (Brusgaard et al., in press; Zeiler, 1997), beaver remains make up between 12% and 23% of total NISP. Beaver-gnawed willow branches were found at S3 (Casparie et al., 1977) and Prummel (2017) has recently highlighted that the Swifterbant environment would have been ideal for human exploitation due to beaver engineering. In contrast, at the archaeological sites Tiel, Hoge Vaart and Nieuwegein, beaver represents less than 5% of the total NISP, and there is more emphasis on large wild fauna. Otter remains fluctuate almost on par with beaver remains at each site, ranging between 29% at Hardinxveld-Giessendam Polderweg (where beavers are abundant) and <1% at Tiel (where beavers are few). The only exception to this pattern is Schokland where the NISP of beaver is high but only few otter remains were recovered.

From an environmental perspective, the prevalence of beaver remains at many of the Swifterbant Culture sites is not surprising considering their location in freshwater riparian landscapes. The Dutch data discussed here is generally biased towards such locations because faunal remains from archaeological sites on the sandy (drier) soils have not been preserved and no coastal sites are known, probably due to erosion (Vos, 2015). It is therefore presently unknown which wild fauna was exploited in these other regions and ecotones. While the presence of beavers at Mesolithic and Early Neolithic wetland sites is thus not unanticipated, the relative abundance of beaver remains points to some form of concentrated exploitation of these animals (and of otters) by Swifterbant communities. The main synchronic and diachronic trends in beaver remains appear to be negatively structured by the frequency of large prey animals such as cervids and wild boar, suggesting a strategic trade-off between either beaver and otter or the targeting of such larger species. There is, however, no evidence for a switch to other fur-bearing animals such as martens, as often observed in more recent periods, and which for example appears to be reflected in the faunal data from EBK hunting locales in Southern Scandinavia (see above).

Macro-patterns in beaver-related material culture

Beaver-related material culture has been recovered from across the Northern European wetland and boreal zones in the Early and Mid-Holocene (cf. Figure 1; Supplementary Information 1, available online). This material culture can be grouped into four primary groups: (i) incisor tools, (ii) mandible tools, (iii) tooth pendants, and (iv) other modified and unmodified but selected beaver bones not directly tied to food-getting or tooling endeavours, such as ankles. Incisor tools are made from the front teeth of beavers without their associated bone sockets, either by manipulating the teeth themselves or by simply using and/or subsequently re-sharping them. Mandible tools consist of completely or partially removed beaver mandibles including the front teeth, often modified and roughly shaped to facilitate effective instrumentalization. Tooth pendants are defined as beaver teeth that are either grooved or perforated for suspension.

Incisor and mandible tools

Incisor tools are found across the entire region, from the Early Mesolithic to the Late Mesolithic/Early Neolithic, whereas mandible tools are mainly known from the Baltic and the Russian West, where they are abundantly found at Mesolithic sites of all ages (Lozovskaya and Lozovski, 2015; Lozovskaya et al., 2017; Zhilin, 2001, 2020). For Russian contexts alone, Zhilin (2020) reports more than 1400 beaver mandible and incisor tools dated to the Mesolithic, mostly from the key wetland sites. Most of these objects represent prepared and/or used mandibles (n = 1388) and only a handful conform to modified frontal teeth (n = 34). The number of such objects varies greatly among archaeological sites, ranging from sites with only a small amount of such tools to archaeological sites such as Ozerki 5/IV and Veretje 1 bearing more than 100 objects. Mandible tools are least frequent in the Early Mesolithic and increase from the Middle to the Late Mesolithic/Early Neolithic (Zhilin, 2020). Incisor tools are currently unknown from the Early Mesolithic and most of these artefacts derive from Middle Mesolithic contexts within the region.

In Northwestern Europe, beaver-sourced tools from the Mesolithic are much less abundant but they have been reported in the literature (Figure 4 and Table 2). A single worked mandible has for example been described from the Early Mesolithic occupations of Star Carr in Britain (Knight et al., 2018), attributed to the Maglemose complex. Even though Star Carr lies outside of the study area, it showcases the early origin of beaver-related material culture also in northwest. In Southern Scandinavia and Northern Germany, mandible tools are presently only known from Maglemose contexts, including some of the classic Maglemose localities from Zealand in Eastern Denmark (Broholm, 1924; Hatting, 1970; Lautsen Lomborg, 2021). They mainly date to the Boreal/earliest Atlantic (‘Middle Mesolithic’ in the here-adopted terminology) and thus likely belong to the later part of the Early Holocene, even though this material should be radiocarbon dated to confirm this tentative placement. Beaver incisor tools, even though mostly represented by isolated pieces, were found at the Middle Mesolithic sites of Holmegård, Ørgård and Sværdborg in Denmark (Hatting, 1970; Lautsen Lomborg, 2021) and at Hohen Viecheln I (Schmölcke et al., 2017; Schuldt, 1961) and Rothenklempenow 17 (Schacht and Bogen, 2001) in Northern Germany. Beaver-sourced tools, both mandibles and incisors, were also found in a likely EBK context at the site of Heidemoor in the German Northeast (Ewersen, 2011), but this attribution similarly awaits future corroboration and is currently an isolated case.

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Figure 4.

Selected beaver-sourced tools from the Mesolithic of the Northwestern European wetland zone. 1–4: beaver incisor tools; 6–7: beaver mandible tools. 1: Hohen Viecheln (Germany), Middle Mesolithic (Schmölcke et al., 2017: Figure 6; photograph: H. Lübke, ZSBA Schleswig); 2: Hardinxveld-Giessendam Polderweg (The Netherlands), Late Mesolithic (Coles and Kooijmans, 2001: Figure 2); 3: Holmegård (Denmark), Middle Mesolithic (Hatting, 1970: Figure 10); 4: Øgårde (Denmark), Middle Mesolithic (Hatting, 1970: Figure 9b); 5: Spjellerup (Denmark), Middle Mesolithic? (Hatting, 1970: Figure 4); 6: Lynby (Denmark), Middle Mesolithic (Hatting, 1970: Figure 8b).

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Table 2.

Overview of published beaver-related material culture from the Mesolithic and earliest Neolithic of continental Northwestern Europe.

Site	Country	Dating/chronology	Period	Description	References
Star Carr a	England	Preboreal, Early Maglemose, c. 9500–8500 BC	Early Mesolithic	A single worked beaver mandible half associated with a few other beaver bone remains in the earliest Mesolithic occupation	Knight et al. (2018), Milner et al. (2018a)
Hardinxveld-Giessendam Polderweg	The Netherlands	Atlantic, c. 5500–4650 BC	Late Mesolithic/Early Neolithic	Six modified beaver incisors, probably for use as chisels. The site also has produced >1000 NISP of beaver remains	Coles and Kooijmans (2001), Louwe Kooijmans et al. (2001), Dreshaj et al. (2023)
Hardinxveld-Giessendam De Bruin	The Netherlands	Atlantic, c. 5450–4250 BC	Late Mesolithic/Early Neolithic	Eight modified beaver incisors, probably for use as chisels. The site also yielded >1000 NISP of beaver remains	Coles and Kooijmans (2001), Louwe Kooijmans et al. (2001), Dreshaj et al. (2023)
Lundby	Denmark, Zealand	Boreal/early Atlantic, Late Maglemose	Middle Mesolithic	A single beaver mandible with traces of anthropogenic removal and potential use, associated with a small number of beaver bones (quantity unknown)	Hatting (1970)
Spjellerup	Denmark, Zealand	Boreal/Atlantic	Middle Mesolithic?	Isolated beaver mandible with traces of anthropogenic use recovered from a bog	Hatting (1970)
Ravnsbjerggård	Denmark, Zealand	Boreal/Atlantic	?	Two beaver mandibles with traces of anthropogenic use	Hatting (1970)
Holmegård	Denmark, Zealand	Late Boreal/early Atlantic, Late Maglemose, c. 6500 BC	Middle Mesolithic	A single split and worked beaver incisor within an assemblage of 70 beaver remains including skull fragments, mandibles and isolated teeth	Hatting (1970), Lautsen Lomborg (2021)
Øgårde	Denmark, Zealand	Boreal/early Atlantic, Late Maglemose	Middle Mesolithic	A single incisor with polish and reworking traces as well as a few worked beaver mandibles associated with >200 beaver remains including some beaver mandibles and a few isolated teeth	Hatting (1970), Lautsen Lomborg (2021)
Sværdborg	Denmark, Zealand	Boreal/early Atlantic, Late Maglemose	Middle Mesolithic	Two removed and used beaver incisor tools within a larger assemblage of beaver remains including mandibles and teeth (n = 219)	Hatting (1970), Lautsen Lomborg (2021)
Hohen Viecheln I	Germany	Boreal, Maglemose	Middle Mesolithic	A few used pairs of frontal beaver incisors (n = 3) and isolated incisors with use marks (n = 3) within a small assemblage of beaver remains. The frontal part of one such incisor tool is recorded to have been attached to a wooden stick	Schuldt (1961), Schmölcke et al. (2017)
Rothenklempenow 17	Germany	Boreal, Maglemose	Middle Mesolithic	A single pair of frontal incisors glued together with birch tar, found in association with a few beaver remains (number unknown), otter bones and a notable quantity of fish, especially carps, pike, perch and zander	Schacht and Bogen (2001); Schmölcke et al. (2017)
Heidemoor	Germany	Later Atlantic, Ertebølle?	Late Mesolithic/Early Neolithic	Six modified (polished) beaver mandibles and characteristic chipping of used incisors on mandibles; assemblage contains a large assemblage of mandibles and mandible fragments and 53 isolated lower jaw beaver incisors	Ewersen (2011)

a

Star Carr is only mentioned for comparative reasons as the earliest evidence for beaver-related material culture outside of the study area.

No mandible tools have been found in the Netherlands. Only incisor tools are known and they derive from the two Late Mesolithic Hardinxveld-Giessendam sites (Coles and Kooijmans, 2001; Louwe Kooijmans et al., 2001, 2001). These tools bear a remarkable similarity to the incisor tools recovered from the Danish Middle Mesolithic sites. Esser et al., in preparation) further draw attention to the circumstance that at Tiel-Medel, while beaver mandibles have been recovered, only a very small number of beaver teeth occur in the assemblage, which is taphonomically unlikely, and may thus point to anthropogenic selection and filtering. It is for example possible that the missing beaver teeth have been removed for use or ornamentation elsewhere or were simply exported from the site but this hypothesis requires future empirical substantiation. That said, an analogous case is possibly framed by the Early Mesolithic site of Huseby Klev on the west coast of Sweden, where only beaver teeth and a single nearly complete mandible were found among the faunal remains and strontium isotope data points to a non-local origin of the material, contrasting with the rest of the zooarchaeological assemblage (Boethius et al., 2022).

The evidence from Northwestern Europe is extremely sparse, despite researchers being keenly aware of such finds and thus on the lookout for them (e.g. Esser et al., in preparation; Enghoff, 2011), suggesting this is most likely not due to researcher bias. The Northwestern earlier Mesolithic record thus appears to be structurally different from what is observed in the Northeast. Most relevant beaver-related material culture from the region either dates to the Middle Mesolithic or is associated with the earlier part of the Mid-Holocene, even though the precise dating of some of the objects remains problematic. It is worth noting that for the EBK specifically, tools made from animal bones are rare overall, so this difference may have less to do with the role of beavers in particular and more with the status of animal-related material culture in general. In Swifterbant Culture contexts, by contrast, a wide diversity of species appears to have been involved in tool production – from wild boar to swans to caprines (Aal and van Gent in preparation; Kranenburg and Prummel, 2020; Louwe Kooijmans et al., 2001) – suggesting little species-level discrimination. More detailed analysis, for example with regard to body part selectivity (see e.g. Hill, 2019 and Hussain et al., 2022 for the key importance of the latter), may be warranted, however.

The function of beaver-sourced tools has been discussed extensively in the literature (e.g. Coles, 2006; Lozovskaya et al., 2017; Schmölcke et al., 2017; Zhilin, 1997, 2020). At Middle Mesolithic Veretje 1, the first ever securely identified beaver mandible tool was reportedly still bound with a strip of bark, revealing its tool-character and the way it was instrumentalized (Oshibkina, 1983; Zhilin, 1997). Similar mandible tools attached to wooden handles are for example known from Indigenous people from Alaska who used them as scrapers, the molars functioning like a rasp (Osgood, 1940; Schmölcke et al., 2017). Following Zhilin (2014a, 2014b, 2020), Mesolithic instances of such beaver-sourced tools were mainly used as scrapers, knives, and chisels or as pressure flaking devices. Woodworking was an important task of many of these tools but some scraper-like mandible tools were apparently also used for bone-working (Zhilin, 2020). The utilization of these objects therefore largely mirrors the capacity of the respective body-parts in a living beaver, which has led Schmölcke et al. (2017: 8) to suggest that ‘perhaps by observing these animals prehistoric people got the notion that they have built-in woodworking tools’. We return to this point below and take it up again in the discussion.

The in-depth techno-functional analyses of these objects performed by Zhilin (2020) have further shown that beaver-sourced tools were not deployed ad hoc, as might perhaps hastily be inferred from a pragmatic copy-paste logic vis-à-vis beaver woodcutting practices. Instead, the chaîne opératoire of these objects is often surprisingly complex and demonstrates distinct stages of reworking (Zhilin, 2020), suggesting not only that many of these tools were probably in use for quite some time, but also that they were actively curated and thus generally cared for. We must assume that Mesolithic people generally had the capacity and means to acquire beaver bodies to replenish tool stocks if they had wished to, and the extended life-histories of beaver-sourced tools thus strongly suggest that the objects mattered to people, and keeping the same items in human systems for prolonged periods of time was a conscious concern. Furthermore, using and handling these tools would have brought beaver bodies to the centre of human ‘horizons of concern’ (sensu Bird-David, 2017), promoting understanding of and sympathy for the beaver, and thus bringing human and beaver perspectives closer together, with human and beaver bodies as well as perceptive and cognitive horizons literally merging, if only temporally.

Ichtyofaunal patterns

Fish remains from the Northern Mesolithic provide important information on human landscape use and exploited aquatic ecologies, and may further disclose hitherto overlooked evidence for human-beaver cohabitation and encouragement in wetland and freshwater environments. Similar to the data on mammalian faunal compositions, the available evidence on human-procured fish during the Mesolithic indicates a broad distinction between ichtyofaunal assemblages in Northwestern Europe on the one hand and the Baltic and the Russian West on the other (Figures 5 and 6; Supplementary Information F, available online).

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Figure 5.

Number of identified fish specimens (NISP%) from the Baltic countries and Western Russia from the Early to the first part of the Mid-Holocene: (a) Early Mesolithic; (b) Middle Mesolithic; (c) Late Mesolithic/Early Neolithic. Horizontal bars represent median values with standard deviations. Animal silhouettes have been redrawn from https://www.phylopic.org/. Raw data are provided in Supplementary Information F Tables 1 and 2, available online.

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Figure 6.

Number of identified fish specimens (NISP%) from Southern Scandinavia and the Dutch wetlands from the Early to the first part of the Mid-Holocene: (a) Early Mesolithic; (b) Middle Mesolithic; (c) Late Mesolithic/Early Neolithic. Note that data for the Dutch wetlands is only available for the final phase. Horizontal bars represent median values with standard deviations. Animal silhouettes have been redrawn from https://www.phylopic.org/. Raw data are provided in Supplementary Information F Tables 3 and 4, available online.

Northeastern Europe

In Western Russia, northern pike (Esox lucius) dominates the ichtyofauna from the Early to the Late Mesolithic (mean NISP = c. 50–70%) but the relative importance of pike decreases slightly over time, while other large freshwater predators such as the European perch (Perca fluviatalis) become more frequent throughout the Middle and Late Mesolithic (Figure 5). Early Mesolithic assemblages tend to be more monospecific, focusing either on the pike or, in one case, on the predatory zander/pike-perch (Sander lucioperca: NISP = 95%). From the Middle Mesolithic onwards, the ichtyofauna not only becomes more diverse, it is also notably enriched in carps and ground-feeders in general. Wels catfish (Silurus glanis), occasionally found already in the Early Mesolithic, emerges as a regular although low-frequency component of Late Mesolithic/Early Neolithic assemblages. Importantly, the vast majority of represented species forms part of lentic communities (Figure 7) and some of the notable later Early Holocene and Mid-Holocene species suggest that eutrophic conditions must have been well-established in the landscape by then. The eel (Anguilla sp.) remains exceptionally rare even in the Late Mesolithic, when the Littorina Sea was already well developed (Kostecki, 2014) and eel has been shown to be generally present in Baltic waters (Enghoff and Ediger, 2016).

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Figure 7.

Marine versus freshwater fish ecologies within ichtyofaunal assemblages as represented by NISP shares in different regions and periods. Species classification and summary data table are provided in Supplementary Information 2, available online.

In the Baltic, the evidence is sparser and some patterns are repeated. In the Early and Middle Mesolithic, the ichtyofauna is dominated by both northern pike and zander/pike-perch, which together make up at least >60% and often >90% of recorded fish NISP. Yet in contrast to Western Russia, northern pike becomes more frequent in the Late Mesolithic/Early Neolithic, often at the expense of zander, and the Middle and Late Mesolithic are again characterized by a notable catfish input (up to NISP = 28% in the Late Mesolithic). Interestingly, Wels catfish seems to supplant carps in the course of the Mesolithic, but the dataset is not robust enough to securely establish this trend. Kõpu I in Estonia presents a rare example of notable marine fish input during the Late Mesolithic/Early Neolithic, but the fish assemblage is too small (NISP = 6) to extrapolate from this so far isolated context. Given that the site’s mammalian fauna is dominated by seal (Lõugas et al., 2017), the marine fish component is likely the result of specific foraging activities linked to targeted phocid hunting at the emerging Baltic Sea, and is thus not necessarily reflective of a systematic practice and/or well-established marine fishing economies (see esp. Boethius et al., 2017 for similarly cautionary arguments regarding another Baltic Early Holocene context from the Swedish island of Gotland).

Mammal hunting affordances

A ‘beaver lens’ reveals interesting patterns in species co-associations and possible long-term dynamics in multispecies assemblages related to beaver agency. In Northeastern Europe, for example, the predominance of elk alongside beaver is noteworthy as beaver-elk ecosystem associations are reflective of incipient colonization scenarios where beaver-powered wetlands create a mosaic of standing water bodies and promote the growth of riparian or edge vegetation such as willow (Gibson and Olden, 2014), in turn attracting elk (Baker et al., 2005; Nummi et al., 2019; Ripple and Beschta, 2004). Beavers and elk also display a high degree of dietary overlap (Hoy et al., 2019). As previously argued by Petersen (2009: 46), beaver-elk interactions are often beneficial for human foragers and the beaver may have accordingly come into view as a ‘faunal regenerator’ because of this notable ecological interrelationship. Similar ecosystem dynamics are harder to trace in Northwestern Europe, in part because elk is much less frequent in faunal assemblages there. In Southern Scandinavia, for instance, elk altogether disappears from the zooarchaeological record in the course of the Mesolithic, even though, puzzlingly, the species remains culturally important (Bridault, 1992) – but these processes may in fact be interrelated. In the Dutch wetlands, elk is consistently present only in small numbers, despite wetlands being among the preferred habitats of the species (cf. Janík et al., 2021), opening up the possibility that Swifterbant Culture communities culturally opted not to hunt elk systematically. This scenario is not in principle inconceivable for the later Mesolithic of Southern Scandinavia as well.

A recent ecological study conducted in Denmark has shown that red deer and roe deer avoid water in areas with high beaver activity, and that these cervids tend to occur in higher numbers in areas with fewer beavers (Svanholm Pejstrup et al., 2023). This is noteworthy because the Dutch Mesolithic and Early Neolithic record contains few Cervidae and at the same time features high numbers of beavers – precisely the predicated pattern. In Northeastern Europe, the pattern is overall the same, and the few sites with abundant red deer (Zvidze and Zemaitiskes) yielded few beaver remains. In Southern Scandinavia, by contrast, red deer and roe deer are very common and, as noted earlier, beaver much less so than in the other regions. Svanholm Pejstrup et al. (2023) caution that human impact likely plays a formative role in these present-day observations on interspecies dynamics, so it remains an open question as to what extent such dynamics can be extrapolated back in time, but they are in any case worth considering and should further be explored in and tested against the archaeological record. It should at least be considered that beaver geopraxis may have been less prominent and consequential in Southern Scandinavia when compared to the Dutch wetlands and Northeastern Europe as a whole.

The faunal record of Northwestern Europe hosts a diversity of megafauna, including Cervidae, horse, wild boar and aurochs. Cervidae and horse indicate a more open wetland zone interspersed with deciduous and mixed woodland already very early in the Holocene. Over the course of the Mesolithic, these animals would have thus played a role in further opening up the dense alluvial forests through extensive grazing (e.g. Noe-Nygaard et al., 2005). In the Baltic, the documented increase of wild boar, red deer, and eventually aurochs may thereby have been encouraged by well-established openings at the edge of mature beaver habitats and the role of beaver hotspots as landscape attractors for these animals (Fedyń et al., 2022) – probably fuelled by early low-level human woodland impact and perhaps management (cf. Poska et al., 2004). This may point to an easily overlooked interplay – with notable feedback potential – between beaver woodland engineering and clearance on the one hand and ungulate suppression of woodland regeneration on the other hand – a dynamic possibly consequential for Northwestern Europe and its specific Late Mesolithic/Early Neolithic human histories.

Unlike deer, wild boar are strongly attracted to beaver-engineered landscapes and in particular aged beaver ponds provide ideal foraging and wallowing locales for them (Nitsche, 1997; Rosell et al., 2005). Their co-occurrence with beaver remains in the zooarchaeological record in all of the study regions is therefore interesting and probably influenced by such interspecies co-facilitation, in turn providing human foragers with the possibility to intercept and target boar in beaver country. As discussed earlier, wild boar forms an important element of subsistence across Mesolithic Northern Europe, for meat – and for teeth, tools and pendants. It is important to note that that wild boar appears to acquire a new role in human economies in the Late Mesolithic and Early Neolithic at least in some areas, probably due to local anthropogenic encouragement and/or incipient management strategies (Brusgaard et al., forthcoming; Magnell, 2006; Maring and Riede, 2019; Rosvold et al., 2010). This latter process may not be unrelated to the likely facilitation of wild boar in beaver landscapes, as humans foraging in these landscapes would also intersect and increasingly interact with wild boar, perhaps laying the foundation for subsequent management, taming, and domestication.

Some smaller mammals in the dataset may similarly signal beaver-related ecological promotion. In Western Russia, the Early Mesolithic is characterized by the importance of muskrat and marten, animals who have been argued to form an early colonizer assemblage within formative beaver landscapes (Crego et al., 2016; Nummi et al., 2019). The case of the muskrat may be especially instructive as this semi-aquatic rodent is known to parasitize beaver lodges (Mott et al., 2013), so that knowledge on beaver landscapes can promote muskrat foraging and generally supports integrative foraging strategies in Boreal ecologies (cf. Winterhalder, 1981; see below). The relationship between the beaver and the water vole in this region, as noted earlier, is of further interest in this context because of the common ecological association, and succession, of the two species, which is also an important concern of present-day ecological restoration projects (cf. Stringer and Gaywood, 2016). This latter interspecies relationship is perhaps also reflected in geoarchaeological findings from Grabow 15 in Northern Germany, where an early Atlantic beaver burrow system was documented in conjunction with a later, possibly Late-Holocene, water vole-gnawed piece of wood (Tolksdorf et al., 2017), supporting the idea that water vole is an indicator species of developed-matured beaver wetlands.

Otter is consistently present in Mesolithic assemblages in which beavers are also prevalent, probably because of the earlier outlined facilitation of otters through beaver geopraxis (LeBlanc et al., 2007; Reid, 1984; Tumlison et al., 1982). In the Netherlands, the exploitation of both beaver and otter has been shown to be a key feature of Swifterbant Culture sites, as well as Middle Neolithic sites attributed to the so-called Vlaardingen Culture (c. 3500–2600 BC) (Brinkkemper et al., 2011; Lauwerier et al., 2005; Zeiler, 1987). This pattern supports the observations from Northeastern Europe, where these two species tend to converge in the faunal record, and accordingly lends credence to the idea that integrated low-level beaver and otter foraging was often opportune and possibly sustainable. In Southern Scandinavia, marten, mink (Mustela sp.) and otter are also a recurrent but low-abundance faunal component in the Early and Middle Mesolithic, which may similarly be indicative of exploited foraging affordances within beaver habitats. It is not clear, however, whether the decrease in beaver remains documented in the Late Mesolithic of the region denotes a principal change in hunting practices and logics with regard to beavers, or instead signals a switch to other animals occurring in beaver landscapes at the expense of beavers, or both. For example, some sites feature hardly any beaver remains but have yielded abundant other fur-bearing animals like pine marten (Martes martes) and wild cat, who were probably targeted separately within specialized/dedicated fur-getting economies (Price, 1991; Richter, 2005; Richter and Noe-Nygaard, 2003; Trolle-Lassen, 1987). Otter in the EBK was likely targeted in the context of coastal and marine activities (e.g. Price et al., 2001), signalling a reorganization of foraging affordances and perhaps even their partial disintegration, and this may indeed indicate subtle but relevant changes in the role of beaver affordances in regulating larger foraging systems.

Taken together, these successional and interspecies faunal dynamics may point to the crucial significance of integrated foraging systems in the Mesolithic of Northern Europe, centred on animal resources directly accessible at hotspots of beaver activity or at sites of former beaver occupation (legacy sites). In Southern Scandinavia, this constellation may have fundamentally been reconfigured towards the end of the Mesolithic, while demonstrating substantive continuity in the other North European study regions. This human-beaver entanglement would have granted beaver landscapes special significance for human foragers and it is tempting to conceptualize such ‘beaver country’ in analogy to notions of Country perpetuated by Indigenous groups across Australia, emphasizing reciprocity and nourishment through human care (e.g. Urwin et al., 2022). Either way, these dynamics of human-beaver cohabitation would have drawn the beaver closer to the centre of forager lifeworlds and ‘horizons of concern’ (sensu Bird-David, 2017), fostering people’s attentiveness to the animals and their resource provisioning work as well as the attendant multispecies rhythms. Beavers, then, hold a double status as ecological and phenomenological cornerstones of Northern Mesolithic more-than-human landscape ‘dwelling’ (sensu Ingold, 2022). In this optic, the beaver, furthermore, quickly acquires the status of a collaborator – a feral partner – in the pursuit of human sustenance. The ecological keystone status of beavers in Northern Europe – their capacity to anchor and assemble animal ecologies and ecosystem processes – therefore arguably laid the foundations for the ecological facilitation of Mesolithic human foragers living in the same landscapes, who in turn integrated a large suite of beaver landscape affordances into their behavioural repertoire. As Kikvidze and Callaway (2009) point out, facilitation can be a powerful but often overlooked evolutionary factor, structuring multispecies communities and shaping long-term dynamics of history.

Fish-getting affordances

The possibility of beaver facilitation is equally supported by the Mesolithic fish record. The striking feature of the ichtyofaunal record from the earlier part of Holocene Northern Europe is its strong emphasis on larger predatory freshwater fish such as pike, zander and perch in conjunction with a growing importance of lentic bottom-dwellers over time, such as carps and Wels catfish. This accentuation is consistent with some known effects of beaver-fish interactions, while the observed regional variability similarly points to important differences in hydrological systems and aquatic ecologies across Northern Europe – especially the relative importance of larger lakes vis-à-vis riparian corridors – and the changing impacts of the beaver on these systems and flow regimes. Beaver activity generally fosters habitat heterogeneity over larger spatial scales by creating patches of lentic habitat within a corridor of lotic habitat, thus altering and framing new niches for both ‘stream species’ and ‘pond species’ (Collen and Gibson, 2000; Snodgrass and Meffe, 1999: 452). In warm water ecosystems, beaver ponding, eutrophication, and lentic shallow-water and edge-habitat engineering mainly promote potamodromous (i.e. freshwater-only) species such as pike and perch, who can over time outcompete and replace smaller bodied cyprinids (e.g. Gaywood, 2018; Pliūraitė and Kesminas, 2012; Rosell et al., 2005). Pike is found to benefit from beaver ecosystem modification especially when larger ponds or lakes are available (Collen and Gibson, 2000), while perches are attracted by beaver lodge debris (Gibson, 1969). As conditions within ponds become increasingly anaerobic, larger carps and species such as catfish increasingly benefit and typically grow in abundance. Wels catfish, once promoted, can further impact freshwater ecologies due to induced alterations of the trophic chain and physiochemical modifications of the water content (Rodriguez-Labajos et al., 2009), sometimes leading to the displacement and near-disappearance of other species such as the common carp (Cyprinus carpio) (Snodgrass and Meffe, 1998), whilst attendant algae growth may complicate even the spearing of larger fish such as pike.

In all study regions, the fish evidence often reveals a gradual re-configuration of lentic fish communities in the course of the Mesolithic – a successional sequence starting with an emphasis on pike and other large freshwater predators leading to increased attention to cyprinids and eventually catfish. This, then, is unlikely to be reflective only of human foraging preferences but probably also records a latent beaver legacy effect, as these changes are a consequence of consolidating and ageing beaver landscapes including maturing ponds, some of which are ultimately abandoned, offering distinct fishing opportunities. Fish-getting practices and human strategic decision-making during the Mesolithic were thus likely influenced, and dynamically modulated, by the specific fish-getting affordances emerging from long-term beaver activity in the hydroactive wetland and boreal environments of Northern Europe at the edge of former glaciers.

Beavers almost never fully disrupt riparian connectivity (Schlosser, 1995) but they can severely impede the capacity of river migrating species such as salmons and eels to traverse across riparian landscapes (cf. Kemp et al., 2012). Mitchell and Cunjak (2007) found that beaver dams in coastal rivers prevent upstream migration of salmonids and simultaneously, through competitive exclusion, increase fish diversity upstream. This dynamic may apply to, and in part explain, the ichtyofaunal patterns of the Late Mesolithic/Early Neolithic from the Dutch wetlands, where salmonids are conspicuously rare. This pattern indeed continues at Middle Neolithic wetland and coastal sites in the area, where seal and other marine mammals increase in importance, yet the ichtyofaunal emphasis remains on pike, sturgeon, eel and cyprinids (Lauwerier et al., 2005). This could suggest long-lived beaver legacies, hunting affordances, and the resulting enfolded cultural practices. A highly similar pattern emerges from the Middle Mesolithic of the Baltic, where beaver-supported lentic fish including pike, perch, and cyprinids remain the focal target of fish-getting practices, even though human foragers begin to engage in specialized seal hunting (Lõugas et al., 2017). The increasing importance of the sea does therefore, contrary to what might be assumed, not lead to a dramatic shift in ichtyofaunal acquisition patterns, and lotic fish continue to form only small portions of human-foraged fish. Consonant with this broader perspective, Boethius et al. (2017) have previously argued that freshwater fish derived from eutrophic lakes remained a key subsistence staple even in the context of the initial Early Holocene occupation of Gotland in the Baltic basin, while marine mammal hunting, especially the targeted pursuit of younger seals, was secondary and probably primarily oriented towards and motivated by raw material acquisition.

Salmonids and eel inputs to Mesolithic fish assemblages remain generally negligible, perhaps indicating some level of amensalism (cf. Arthur and Mitchell, 1989) between earlier Holocene beaver-powered environments and migratory, lotic fish requiring access to the open sea. This idea may be supported by the faunal evidence from Mesolithic Ireland, where the beaver is not part of the native mammalian fauna, while salmonids and eel are of key importance in the anthropogenic fish assemblages (Kelly, 2005; Warren, 2022), contrasting with the data from Mesolithic mainland Europe and Britain where these species are virtually absent (Robson and Ritchie, 2019; Robson et al., 2018; Zhilin, 2014a). This again suggests that beaver agency co-structured anthropogenic fish assemblages. Alternatively, or perhaps complementarily, lentic fish, especially larger predatory species and fast-growing carps, offered more attractive, more reliable and easier to access food resources due to their predictable association with beaver-fabricated ponds and wetlands.

In this context, a few comments on the evolution of fishing technologies in Northern Europe are useful. Sophisticated and curated fishing installations such as fish weirs, fences and/or proper fisheries only emerge in the course of the Mesolithic and typically date to the later part of the Middle Mesolithic or the Late Mesolithic/Early Neolithic in the study regions (e.g. Amkreutz, 2013; Fischer, 2007; Lozovski and Lozovskaya, 2016). In the Early Holocene, fish was thus probably often acquired via spearing and/or bowing, sometimes but not always in conjunction with angling, and these practices can easily take advantage of fish trapped in ponds, and the shallow water habitats engineered by beavers can greatly facilitate the spotting, spearing and/or bowing of larger freshwater fish such as pike. Eutrophic lakes and ponds have consequently been invoked as potent landscape attractors for Early Holocene foragers in Northern Europe (Boethius et al., 2017) and Welinder (1978) specifically suggested that overgrown lakes formed central elements of Maglemose adaptations. Advanced stages of eutrophication and pond maturation, as discussed above, may conversely complicate spearing and bowing and thus inspire new fish-getting strategies.

Contrasting with the Dutch and Northeastern European data, the Late Mesolithic and Early Neolithic record from Southern Scandinavia shows a clear break in ichtyofaunal species compositions (cf. Figure 6). EBK sites demonstrate an increased emphasis on both marine mammals and marine fish (cf. Figure 7). The emergence of coast-bound and increasingly marine economies in the EBK may have thus considerably weakened the probiotic effect of the beaver on human livelihoods and drawn human attention away from beaver habitats, thus simultaneously defusing, or at least re-configuring, the reliance on beaver-engineered, inland foraging affordances. To over-exaggerate, Early and Middle Mesolithic foragers in this region encountered the beaver as a key society-sustaining agent – as a nonhuman fishing aid or more-than-human fishing technology – whereas Late Mesolithic/Early Neolithic societies increasingly relied on their own fishing infrastructure and self-devised fish-getting technologies at coast-inland interfaces, yet perhaps nonetheless inspired by the transgenerational experience of beaver geopraxis. The nature and significance of a human-beaver ‘contact zone’ as envisioned by Hjørungdal (2019b; 2019a) for the Southern Scandinavian Mesolithic thus likely depended on the critical intersection of lived human and beaver geographies and, perhaps more importantly, the extent to which beaver habitats were routinely visited, and thus integrated into broader forager landscapes. In analogy to other documented, integrated human foraging strategies such as ‘garden hunting’ in the Americas (Guiry et al., 2021; Linares, 1976; Stahl, 2020), we may refer to this strategy as ‘pond hunting’ or ‘wetland foraging’. The development of coastal and open-water economies in the course of the Late Mesolithic in Southern Scandinavia would have contributed, then, to the disruption of these broader foraging schemes centred on the diverse resource opportunities in and close to beaver habitats.

Other multispecies affordances

Other indirect ecosystem effects of beaver geopraxis with benefits for hunter-gatherers include waterfowl encouragement and promotion (Brown et al., 1996; Nummi and Hahtola, 2008; Nummi and Holopainen, 2014) as well as beaver ‘gardening’. Waterfowl encouragement is a notable life service for human co-inhabitants as waterfowl was an important subsistence good for meat and feathers (e.g. Zhilin and Karhu, 2002) and this bird category is prevalent in the Mesolithic archaeozoological record of the study regions (e.g. Çakirlar et al., 2019; Lauwerier et al., 2005; Lõugas et al., 2017; Zhilin, 2014a). Water birds are also occasionally rendered in the visual art of the Mesolithic of Western Russia (e.g. Lozovskaya, 2021), coevally pointing to the prominence and potential abundance of these birds in earlier Holocene environments. Waterfowl presence draws in and sustains larger birds of prey such as ospreys and sea eagles and these species have often received special attention by Mesolithic foragers across Northern Europe (Amkreutz and Corbey, 2008; Hussain, 2023a; Mannermaa, 2013), again indexing the crucial role of beaver habitats in framing human forager life, perception, culture and possibly cosmology in the European north. Another example of the important and consequential role of beavers in modulating larger multispecies communities is provided by the pond turtle (Emys orbicularis), who is known to be strongly promoted by beaver wetland engineering and pond-making (Janiszewski et al., 2014) and is well-represented in the archaeology of the earlier Mesolithic of Northwestern Europe (Groß, 2017: 18). Pond turtles notably also make an appearance in the Dutch Late Mesolithic at the Hardinxveld-Giessendam sites (Oversteegen et al., 2001; van Wijngaarden-Bakker et al., 2001) and equally occur at some Southern Scandinavian Middle Mesolithic sites (Groß, 2017). Interestingly, these reptiles then seem to disappear again from the archaeological record during the Atlantic, and this may be linked to a shift in beaver preponderance, landscape impact and/or proximity to human habitation sites in some northern areas at the end of the study period.

In addition, beaver behaviour effectively conforms to a form of landscape gardening – encapsulated by the notion of a ‘beaver meadow complex’ – which promotes distinct plant community successions (Westbrook, 2021), and can notably encourage aquatic plants that grow and proliferate under eutrophic conditions. Water lilies (Nympahea) and water chesnuts (Trapa) both benefit from beaver-induced damming and pond formation (Benke et al., 1999; Kukuła and Bylak, 2017; Law et al., 2014) and the remains of both plants have been found in Northern Mesolithic sites as well as in human coprolites (e.g. Bakels et al., 2001; Kubiak-Martens and van der Linden, 2022; Price, 1991; Zvelebil, 1994). Beavers are known to strategically feed on white water lilies (Nymphaea alba) while promoting them ecologically (Law et al., 2014), and beaver habitats therefore not only signal the potential availability of high-value plant resources, Mesolithic people may have actually discovered this specific resource potential of wetland-lakeland ecosystems by observing and learning from beaver behaviour. Considering the possibility of beaver-directed mimicry as an important ecocultural practice may be fruitful in this context. Mimicry (sensu Bhabha, 1984; GoGwilt and Holm, 2018) describes the ability to imitate or re-enact a socially relevant other in asymmetric power-relations and may be evidenced by co-opting beaver body parts and using them in the same way as the animals, for example deploying beaver-mandibles for woodworking. Such appropriation of nonhuman capacities through material practices, in this optic, may then not only be an expression of relating to beavers as socially relevant others, but could reflect human attempts to literally assume a beaver gaze, as the beaver comes into view as a nonhuman guide and tutor of the north (see e.g. Stobiecka, 2022 for a general exposition of these latter notions), disclosing unique possibilities of navigating and using the landscape. This perspective brings us close to an understanding of Mesolithic beavers as a nonhuman landscaping technology, even though reducing beavers to purely instrumental roles would obviously undermine the gist of the here proposed argument.

All of this being said, beaver-shaped landscapes clearly invite particular foraging behaviours and offer exceptional possibilities for integrated food-getting strategies, but to exploit these efficiently requires intimate knowledge on beavers and their geopraxes. We thus propose that adapting to earlier Holocene environments in Northern Europe in many cases involved human adaptation to beaver behaviours and landscapes. The beaver, in line with Coles’s (2006) previous arguments for Britain, was thus likely a key agent in the Early Mesolithic (re-)occupation of high-latitude Europe as a whole, and ‘landscape learning’ (sensu Rockman, 2003) was promoted by attunement to and familiarization with beaver activity, including efforts to assume a beaver perspective, drawing the beaver into human affairs, at times merging human and beaver horizons, and thereby fostering human respect and care for beaver others, as ‘thinking with’ and ‘acting with’ the animals emerged as an important touchstone of Mesolithic forager life in different parts of Northern Europe.

Human-beaver cohabitation in the Early and Mid-Holocene

The Mesolithic data reviewed and synthesized here may thus be taken to suggest that exploiting the attractive foraging grounds curated by ongoing beaver ecosystem engineering was a central pillar of the earlier Holocene human occupation of Northern Europe, indicating that forager lifeways were at least in part predicated on beaver agency. Human-beaver cohabitation and its associated behavioural possibilities, in other words, emerged as an important precondition for human sustenance and livelihood within the vast wetland and boreal zones of the European north, only to be disrupted when some Late Mesolithic and Early Neolithic societies turned their attention to the sea (and perhaps to other foci of economic, cultural and cognitive concern). The Mesolithic/Early Neolithic of the Dutch wetlands, interestingly, shows notable similarities in overall beaver-related ecosystem relations with the Baltic and the Russian West, rather than Southern Scandinavia, as may initially be expected based on geographic proximity. This convergence may in part be a consequence of the rich delta landscapes in these regions, acting as biodiversity contraptions with extensive wetlands and catalysing beaver habitation as well as the attendant successional dynamics for human foragers (cf. e.g. Giosan et al., 2014; Richardson et al., 2021). Beavers have been shown to play key roles in the maintenance of these landscapes and the provisioning of associated deltaic wetland resources (Hutchings and Campbell, 2005). The cognitive and less tangible aspects of human-beaver cohabitation in these regions are more difficult to glean and open up a host of new questions. Compellingly, however, the here-adopted beaver perspective suggests that geographically close regions in the European Northwest appear to have embarked on divergent ecocultural trajectories, with notable differences in the place of beavers in multispecies systems.

In contrast to the Northwest, the European Northeast stands out in the richness and diversity of its beaver-related material culture. The productive co-habitation and cross-fertilization of humans and beavers in the Early and Mid-Holocene has thus arguably laid the foundation for an increasingly diverse repertoire of beaver-related material practices, understood here as a consequence of fostered human-beaver intimacies throughout the Mesolithic. Extended life-histories of beaver-sourced tools and the human care put into them (cf. Zhilin, 2020) as well as the association between the teeth and astragali of beavers with human bodies strongly suggests that beavers participated in the making of the human social world and became entangled, albeit differently across regions and time periods, with human bodies. The evidence is therefore consistent with the idea that beaver landscape significances became deeply sedimented into cultural memory systems as beavers and humans curated relatively stable and increasingly meaningful interspecies neighbourhoods.

The observation that beaver-related material culture appears in the archaeological record of Northwestern Europe less frequently than in the Baltic and Western Russia and generally tends to postdate the Early Mesolithic is thereby important, potentially showing that beaver knowledge took more time to crystallize in the respective human societies of the northwest, although the comparison may be hampered by the lack of sites from the Dutch Early and Middle Mesolithic. In all regions, however, there is some evidence for the co-optation of beaver capacities through the use and transformation of selected beaver body parts, which embody and exemplify the environmental agency of the animals.

Animal body-part selectivity is a common feature of forager zoo-materialities, often linked to broader concepts of trait fluidity and bodily transposition rooted in relational epistemologies and situated zooontologies emerging from lived interspecies intimacies (Hill, 2011, 2019; Hussain et al., 2022; McNiven, 2022). Beaver-related materializations linked to human instrumental, ornamental, and burial practices, from this perspective, may have helped to produce human-beaver co-sociality, pointing to the socio-historical efficacy and lived significance of beaver neighbourhoods for Mesolithic people. Beaver-related material culture can then be interpreted to reflect the recognition of beavers as ‘co-workers’ and as ‘community’ (sensu Welden, 2023), and thus as a symptomatic feature of what Bird-David (2017) has termed ‘plurispecies’ societies.

The systems-perspective on human-animal interactions pursued here places particular emphasis on the relational assembly, integration, consolidation and disintegration of humans, beavers, landscapes, and material cultures in the course of the Early and Mid-Holocene. It queries the changing ‘intra-actions’ (Barad, 2007; cf. Kirksey, 2015) within these systems, relationships such as conflict, tension, synergy, cross-pollination and possibly co-constitution. Figure 8 attempts to outline the central place of the beaver in these systems – in human world-making during the Mesolithic of Northern Europe as suggested by our analysis and broader discussion. In total, we argue that the beaver’s role as a potent ecological keystone species in the Early and Mid-Holocene of Northern Europe provided the larger context for a prehistory of human-beaver sympoiesis (sensu Haraway, 2016). The status of the beaver as a socially significant other in the Northern Mesolithic was thereby not given but made, and it emerged out of an extended but inter-regionally diverse history of human-beaver co-habitation in the north. Beaver remains and beaver-related material culture, in this view, trace the millennial-scale transformation of the beaver into a ‘cultural keystone species’ (Garibaldi and Turner, 2004; Jacques-Coper et al., 2019; Platten and Henfrey, 2009). As we have seen, the timing and trajectory of this process differs between Northwestern and Northeastern Europe and within Northwestern Europe, and articulates with other documented patterns in the archaeological record. The cultural keystone status of the beaver appears to be conserved in the Northeast and in the Dutch wetlands in the course of the Mesolithic, while human-beaver relationships in Southern Scandinavia appear to be transformed, and perhaps lose their former significance, as human practices are subject to dramatic changes and other animals such as the wild boar (e.g. Magnell, 2006; Maring and Riede, 2019) and marine mammals including killer whales (Andersen and Crabb, 1996) rise to economic, ecological and cultural prominence during the Mid-Holocene.

OPEN IN VIEWER

Figure 8.

Tanglegram of human and beaver practice in the Northern Mesolithic. Beaver agency and geopraxis structure human behavioural systems and provide key affordances for subsistence (hunting, fishing, fowling and wetland gardening) as well as the production, curation and signification of material culture. Human-beaver co-living provides a key adaptive background for human forager life in the European North during the Early and Mid-Holocene. Human practices and materialities, in this view, cannot be properly understood outside of their attendant multispecies systems.

Outlook: The agriculturalist hypothesis

Based on historical data, Liarsou (2013, 2015) has provocatively suggested that the relationship between humans and beavers is re-tailored as humans introduce and invest into new landscape practices such as pastoralism and farming. There are several reasons for this general tendency and many of them have to do with human encroachment and/or destruction of beaver habitats. Increasing population pressure, intensification of economic activities in aquatic areas, deforestation, and cereal cultivation in sync with growing infrastructural and environmental fingerprints, including expanded riparian transportation, can have detrimental effects on the size and distribution of beaver populations, in addition to curtailing and fragmenting beaver habitat and mobility. Changing human relations to the landscape, especially claims to and early annexation or ‘propertization’ of particular places, can also provoke changes in the perception and conceptualization of beavers, frequently shifting human attitudes, as the attention is readily drawn to interactive tension and possible conflict. The beaver’s landscape-altering capacities are then easily cast as ‘destructive’. This is particularly the case when humans become perennial cultivators themselves since beavers may flood and/or severely damage fields and larger agricultural landscapes (Philip, 2022), thus changing the conditions and context of human-beaver interaction. Unsurprisingly, such dynamics form also a major source of present-day beaver-landowner tensions in reintroduction areas, such as Britain, the Netherlands, and Denmark (Coz and Young, 2020; Jansman et al., 2016; Naturstyrelsen, 2020) .

It is therefore possible that the arrival of agricultural life in Northern Europe marks an important turning point in human-beaver relations, and there is indeed evidence for substantial human population growth in this period (Shennan, 2013; Shennan and Edinborough, 2007), coupled with increasing evidence for aquacultural investment in the form of fishing infrastructure and extractive freshwater economies (Amkreutz, 2013; Beerenhout, 2001b; Price, 1985, 2000, 2015) as well as expanded riverine and oceanic transportation, at least in some areas such as Southern Scandinavia, and new systems of livestock management (Brusgaard et al., 2022b; Gron et al., 2016). In Southern Scandinavia including Northern Germany, the transition to agropastoral systems occurs around 4000 BC (Gron and Sørensen, 2018). A decrease of beaver representation within faunal assemblages can already be observed from the Middle to Late Mesolithic onwards, however, well before this poignant transition. In the Netherlands, sites with the first clear evidence for crop and animal management date to around 4250 BC (Brusgaard et al., in press; Dreshaj et al., 2023) and continue to yield relatively high percentages of beaver remains. While beaver abundances decline later in the Dutch Neolithic, this is the case for wild animals in general, in tandem with the increase of domesticated livestock (Çakirlar et al., 2019; Lauwerier et al., 2005). Wetland landscapes and natural resources remain generally important throughout the Dutch Neolithic and even the Bronze Age, with specialized sites continuing to be used for fur-animal extraction (Dusseldorp and Amkreutz, 2020; Zeiler, 1987).

At first glance, therefore, our analysis reveals no clear indications for a correlation between the transition to agricultural practices and human-beaver relations. Yet full-blown farming systems do not become established in the region before the later part of the Mid-Holocene and perhaps even later, so these changes may be delayed. It is also important to note that the expected developments in human-beaver relations may manifest themselves in different ways in the (zoo)archaeological records. Agricultural conditions may for example greatly favour the classification of beavers, together with other wildlife, as ‘pests’ (Liarsou, 2013: 177) and may thus foster concerns to remove them from human landscapes and lifeworlds, in turn motivating targeted ecological suppression or even overhunting. For example, agro-horticulturalists in Mexico carry out subsistence hunting of species that otherwise pose a threat to crops (Santos-Fita et al., 2012), which ironically occurs at the edges of human-shaped habitat which provides attractive habitat for these species. Other lines of evidence, such as harvesting profiles which can inform on the sustainability of hunting practices, may offer additional insights here in the future. Çakirlar et al. (2019) have for example concluded that there is currently no indication for beaver overhunting at Dutch Late Mesolithic sites as the corresponding harvesting profiles indicate mostly adult-oriented hunting, data which these authors interpret as being consistent with a stable source population. However, these profiles are based on a now outdated method of age-determination for beavers (Iregren and Stenflo, 1982) and thus need to be revisited. It is furthermore important to consider that forager hunting patterns may have been less influenced by utility-oriented decision-making predicated on universal, neoclassical notions of ‘rational choice’ than by situated animal ethologies and lived predator-prey responses, including so-called ‘ecologies of fear’ (Brown et al., 1999; Brusgaard et al., 2022a, 2022b; Holmern et al., 2006; Hussain, 2023c; Zanette and Clinchy, 2019). To account for such dynamics requires to acknowledge the ‘bounded rationality’ (Simon, 1957; Wheeler, 2020) of human behaviour, drawing attention to a possible role of diverse nonhuman agents in steering and stabilizing human foraging systems and their intrinsic rationalities. Beaver agency in Mesolithic hunting practices thus certainly demands further comparative investigation, which may in turn shed novel light on the influence of emerging agricultural systems and other landscape practices on beaver relations, behaviours and populations.

An interesting perspective for future work on later Holocene societies are the grazing affordances potentially emerging in aged beaver landscapes. Beaver landscapes where the animals’ activity has turned formerly flooded areas into meadows (Westbrook, 2021) may so provide localized feeding opportunities for human-accompanying livestock. Meadows can in this way act as ‘alluvial grasslands’ (Hejcman et al., 2013; see esp. Ritchie, 2017 for an illuminating historical account) and so serve as natural pastures (Liarsou, 2013: 175), thus offering land-use affordances which differ from the kind of beaver-supplied affordances for foragers without livestock (cf. Coles, 1992). Eriksson (2020) has thoroughly illustrated how meadows, once created, recruit novel human-animal entanglements and thus become central places or even focal points for people in the landscape, with important consequences for the assembly and dynamics of larger multispecies assemblages.

When contrasted with these possible later structural transformations in human-beaver systems, Mesolithic engagements with beavers can be described as ‘commensal’ (Liarsou, 2013: 178; see O’Connor, 2013 for a general account of commensalism) – with human foragers being commensal to beavers. This divergence suggests a changing status of beavers as ‘companion species’ in the sense of Haraway (2008) in the past, linked to important transformations in human lifeways and behavioural regimes. Contextualizing the available archaeological data for human-beaver interactions from the Early and Mid-Holocene of Northern Europe in this way helps to recognize that beavers may have offered different affordances and life services for different types of human societies, and that beaver landscapes, as a result, were likely unequally perceived, valued, and imagined because of these differences. While beaver agency in the earlier Mesolithic helped to anchor and spatially organize forager lifeways, and thus co-structured and fuelled human ‘taskscapes’ (sensu Ingold, 2022), beavers appear to have been gradually excluded from the centre of Southern Scandinavian lifeworlds in the course of the later Mesolithic, while mutually conducive relationships were seemingly maintained in the Dutch and Northeastern European Late Mesolithic. The latter thereby promoted large-scale diversification and promulgation of beaver-related material culture with continuities at least into the developed Neolithic.

In general, the documented dynamics clearly expose the fragility of northern systems of human-beaver conviviality that have evolved over millennia and that greatly depended on human practices allowing beavers to enter the realm of social significance and to garner human concern and care. These archaeological insights on the millennial-scale dynamics of multispecies systems involving beavers and humans – echoing Liarsou’s (2013, 2020) earlier arguments – are important sources of information for the ecological management and restoration of beavers in the present, as they demonstrate the inseparability of human lifeways and the functioning of human-animal systems that also feature beavers. Coupled archaeological and geological research can make highly significant contributions here as the vast majority of millennial-scale beaver landscape legacies remain undocumented, and this even though the cumulative ecosystem and geomorphological impacts of the hundreds of millions of beavers who once modified rivers and floodplains across the northern hemisphere can hardly be underestimated (cf. Wohl, 2021).