Anne Charpentier

Ana S L Rodrigues

Claire Houmard

Alexandre Lefebvre

Krista Mcgrath

Camilla Speller

Laura Van Der Sluis

Antoine Zazzo

Jean-Marc Pétillon

What's in a whale bone? Combining new analytical methods, ecology and history to shed light on ancient human-whale interactions

Keywords: Ancient DNA, archaeology, ethnography, exploitation, stable isotopes, whale, ZooMS

des établissements d'enseignement et de recherche français ou étrangers, des laboratoires publics ou privés.

Introduction

Whales have been important resources to coastal communities for millennia (e.g. [START_REF] Álvarez-Fernández | Occurrence of whale barnacles in Nerja Cave (Málaga, southern Spain): Indirect evidence of whale consumption by humans in the Upper Magdalenian[END_REF][START_REF] Clark | Whales as an Economic Factor in Prehistoric Europe[END_REF][START_REF] Grier | The organization of production in prehistoric Thule whaling societies of the central Canadian Arctic[END_REF][START_REF] Huelsbeck | Whaling in the precontact economy of the central Northwest coast[END_REF][START_REF] Jacob | Arctic whaling: proceedings of the International Symposium Arctic Whaling[END_REF], each individual providing a bounty of food and other valuable products [START_REF] D'auteroche | Voyage en Sibérie fait par ordre du roy en 1761 : contenant les moeurs, les usages des Russes, et l'etat actuel de cette puissance[END_REF][START_REF] Vazquez De Espinosa | Compendium and description of the West Indies[END_REF]. Whale stranding can happen along all coastlines, and their opportunistic exploitation is undoubtedly the earlier form of close contact between humans and whales [START_REF] Savelle | The development of indigenous whaling: prehistoric and historic contexts[END_REF]. However, stranding events are unpredictable, and so whales observed alive in the sea must have been tantalizing to coastal dwellers. In a wide diversity of regions across the world, humans developed the cooperative methods and technology that enabled them to drive or bring to shore some of those whales [START_REF] Reeves | The origins and character of 'aboriginal subsistence' whaling: a global review[END_REF][START_REF] Reeves | A taxonomy of world whaling[END_REF]. Approaching such huge animals was fraught with danger, but came with substantial rewards, not only in terms of resources but also personal prestige [START_REF] Arima | The Whaling People of West Coast of Vancouver Island and Cape Flattery[END_REF][START_REF] Drucker | The Northern and Central Nootkan tribes[END_REF][START_REF] Losey | Opportunistic whale hunting on the southern Northwest Coast: ancient DNA, artifact, and ethnographic evidence[END_REF]. Judging from ethnographical records, whaling was associated with complex cultural practices, involving many individuals [START_REF] Bodenhorn | I'm Not the Great Hunter, My Wife Is": Iñupiat and anthropological models of gender[END_REF], not only for the whaling act itself but also in its preparation as well as subsequent butchering and processing of the captured whales [START_REF] Grier | The organization of production in prehistoric Thule whaling societies of the central Canadian Arctic[END_REF].

Ancient relationships between humans and whales are however difficult to study, because whales can be quite invisible in the archaeological record [START_REF] Smith | The invisible whale[END_REF]. Indeed, given their enormous size, they were necessarily processed on the shore (Figure 1), their skeleton then broken down and dispersed by the action of the waves, leaving few or no archaeological traces. Even though whale bones can provide a valuable material, they are so large that they are seldom brought inland intact, being instead found as highly fragmented or transformed remains that are impossible to identify through traditional anatomical approaches. Other valuable whale resources -such as meat, blubber, and baleen -leave few or no archaeological records. This invisibility likely results in an underestimate of the value of whales to coastal populations, leaving many gaps in our understanding of ancient interactions between humans and the world's largest animals. Yet, a revolution in analytical methods is now making ancient whales more visible than ever, sometimes even in surprising contexts. For example, analyses of environmental DNA in Greenlandic midden deposits suggest that intensive use of whale products started 4,000 years ago, far earlier than presumed from zooarchaeological remains [START_REF] Seersholm | DNA evidence of bowhead whale exploitation by Greenlandic Paleo-Inuit 4,000 years ago[END_REF]. Chemical analysis of cemented organic residues and charcoal samples in Late Iron Age slab-lined pits in Norway revealed their purpose for rendering oil from the blubber of marine mammals, rather than as graves or turf houses as initially presumed [START_REF] Heron | Application of lipid biomarker analysis to evaluate the function of 'slab-lined pits' in Arctic Norway[END_REF].

We present here a multidisciplinary review of the literature, focused on supporting archaeologists confronted with whale bones. By bringing together sources from archaeology, ethnography, history, ecology, and molecular and isotopic techniques our goal is to help these researchers better contextualize and thus interpret their findings, by presenting the most recent methodological advances and discussing the remaining challenges.

We structured this review into three sections. The first section is devoted to the whale bones themselves, and the information that can potentially be extracted directly from them: species identity, bone age, evidence of use, and information on the whale population. The second section reviews the type of evidence that can be obtained from other types of archaeological records besides bones: whale barnacles, lipid and protein residues, environmental DNA, baleen, and the bones of potential human consumers. In the third section, we discuss how the whale may have been obtained (i.e., whether it was scavenged or captured), reviewing traditional methods of whale exploitation, the evidence that these methods can leave, and emphasizing the importance of reliable identification of whale species, given that pre-industrial whaling focused on only a fraction of them.

Given the breadth of the disciplines studied, we have not attempted to provide a systematic literature review in each individual field, but we have focused on the most recent developments (especially with regard to methodological advances) and on the most illustrative examples we have found.

Information obtained directly from whale bones

Whale bone specimens have often been overlooked or misidentified, so we begin this section with a brief overview of their main osteological characteristics.

We then review the main types of information that can potentially be extracted directly from the bones: species identity (using collagen and ancient DNA); bone age (based on radiocarbon dating); evidence of use; and ecological information about whale populations themselves (using stable isotope and ancient DNA).

Characteristics of cetacean bones

Most whale bone found in terrestrial archaeological contexts consists of relatively small and indistinct fragments [START_REF] Buckley | Species identification of archaeological marine mammals using collagen fingerprinting[END_REF]. When recovered in contexts where the use of whale bone is not documented by historic or ethnographic sources, such artefacts can easily be overlooked for lack of trained specialists. For example, [START_REF] Pétillon | Circulation of whale-bone artifacts in the northern Pyrenees during the late Upper Paleolithic[END_REF] and [START_REF] Lefebvre | Interconnected Magdalenian societies as revealed by the circulation of whale bone artefacts in the Pyreneo-Cantabrian region[END_REF] re-discovered over 160 artefacts of whale bone in 23 Magdalenian sites across the Pyreneo-Cantabrian region, mainly large projectile heads previously archived as antler artefacts.

In reality, whale bone -and more generally cetacean bone -is a biomaterial with singular characteristics.

Long bones lack a medullary cavity and the skeleton is highly porous, reflecting cetaceans' skeletal adaptations for improved hydrostatic locomotion (buoyancy control), and deep-sea diving (de Ricqlès and [START_REF] De Ricqlès | Bone histology, heterochronies and the return of tetrapods to life in water: Where are we?[END_REF][START_REF] Gray | Sink or swim? Bone density as a mechanism for buoyancy control in early cetaceans[END_REF] (Figure 2). Histology shows that whale bone presents no clear distinction between compact and cancellous tissues.

The bone tissue is porous with an evenly-spaced network of alveoli [START_REF] Campbell-Malone | Biomechanics of North Atlantic right whale bone : mandibular fracture as a fatal endpoint for blunt vessel-whale collision modeling[END_REF][START_REF] Felts | Structural orientation and density in cetacean humeri[END_REF][START_REF] Margaris | Reconsidering Raw Material Selection[END_REF][START_REF] Pétillon | Circulation of whale-bone artifacts in the northern Pyrenees during the late Upper Paleolithic[END_REF][START_REF] Reiche | Synchrotron radiation and laboratory micro X-ray computed tomography-useful tools for the material identification of prehistoric objects made of ivory, bone or antler[END_REF] (Figure 2). These characteristics help differentiate whale bone from other osseous raw materials, and often make it easy to identify the bone (or the bone object) as coming from a large cetacean. However, on highly fragmented and/or poorly preserved specimens, these characteristics are not always obvious.

Identifying the species

Even for whale bones found in coastal archaeological contexts and correctly identified as marine mammal remains, morphological observation alone seldom allowed their identification to the species level, given the frequent and extensive fragmentation of cetacean bone, the osteological similarities between species, and the scarcity of complete reference collections [START_REF] Speller | Barcoding the largest animals on Earth: ongoing challenges and molecular solutions in the taxonomic identification of ancient cetaceans[END_REF]. In recent years, the development of molecular methods has allowed for the accurate taxonomic identifications of even small fragments. Collagen peptide mass fingerprinting (or ZooMS) has recently been developed and tested for archaeological cetacean identification [START_REF] Buckley | Species identification of archaeological marine mammals using collagen fingerprinting[END_REF][START_REF] Kirby | Identification of collagen-based materials in cultural heritage[END_REF][START_REF] Speller | Barcoding the largest animals on Earth: ongoing challenges and molecular solutions in the taxonomic identification of ancient cetaceans[END_REF]. ZooMS can differentiate baleen whales to the genus level, and in some cases to the species level. Closely related species -such as bowhead whales (Balaena mysticetus) and the three species of right whales (Eubalaena glacialis, E. japonica, E. australis), for example-cannot be differentiated; likewise, odontocetes can usually only be identified to broad taxonomic units [START_REF] Buckley | Species identification of archaeological marine mammals using collagen fingerprinting[END_REF]. ZooMS, however, is ideally suited as a taxonomic screening approach. Rapid and cost-effective, ZooMS can be used to screen a large number of samples to reconstruct species distributions within archaeological sites, or to search for particular taxa of interest, and can easily identify any non-cetacean fragments in the assemblage (e.g., [START_REF] Evans | Using combined biomolecular methods to explore whale exploitation and social aggregation in hunter-gatherer-fisher society in Tierra del Fuego[END_REF]. For example, ZooMS screening has been used to identify the most northerly [START_REF] Hufthammer | Grey whale (Eschrichtius robustus) in Norwegian waters 2000 years ago[END_REF]) and southerly distribution [START_REF] Rodrigues | Forgotten Mediterranean calving grounds of grey and North Atlantic right whales: evidence from Roman archaeological records[END_REF] of the Atlantic gray whale -a population that was extirpated from the North Atlantic before formal analysis could be made of its ecology, migration routes and breeding grounds. ZooMS has also been effective in documenting the diversity of cetacean species exploited in Medieval Europe [START_REF] Van Den Hurk | Cetacean exploitation in Roman and medieval London: Reconstructing whaling activities by applying zooarchaeological, historical, and biomolecular analysis[END_REF][START_REF] Van Den Hurk | Medieval Whalers in the Netherlands and Flanders: Zooarchaeological Analysis of Medieval Cetacean Remains[END_REF]. Minimally destructive compared to other molecular analyses, ZooMS is particularly useful for highly fragmented and/or modified remains, such as artefacts and tools manufactured from whale bone. For example, [START_REF] Hennius | Whalebone Gaming Pieces: Aspects of Marine Mammal Exploitation in Vendel and Viking Age Scandinavia[END_REF] were able to identify the use of North Atlantic right whale (Eubalaena glacialis) bone in the manufacture of Viking age Scandinavian gaming pieces. Moreover, there are several non-destructive ZooMS methods that can be employed including the analysis of collagen adhering to the plastic bags in which artefacts were stored, or collected through gentle rubbing with a PVC eraser (e.g., [START_REF] Mcgrath | Identifying Archaeological Bone via Non-Destructive ZooMS and the Materiality of Symbolic Expression: Examples from Iroquoian Bone Points[END_REF]. These minimally-invasive methods however, have yet to be systematically tested on whale bone remains. Genetic analyses can provide higher resolution taxonomic information than ZooMS, providing identifications to species or population level (Foote et al., 2012), and potentially even identifying specific ecotypes [START_REF] Foote | Ancient DNA reveals that bowhead whale lineages survived Late Pleistocene climate change and habitat shifts[END_REF]. DNA identifications are particularly effective in the case of odontocetes, where ZooMS lacks the taxonomic resolution to differentiate many beaked whale, dolphin and porpoise species [START_REF] Biard | Genomic and proteomic identification of Late Holocene remains: Setting baselines for Black Sea odontocetes[END_REF].

Identification to the species level is crucial for understanding the past composition of marine mammal assemblages, as well as the spectrum of interactions that humans may have had with whales, from an opportunistic utilization of stranded or drift whales (possible for all species) to active whaling (plausible for only a few species in pre-industrial times; [START_REF] Rodrigues | Ancient whale exploitation in the Mediterranean: species matters[END_REF]). This aspect is discussed in the section 4.3 species matters.

Bone age

Radiocarbon dating is crucial to properly contextualize the information obtained from whale bones. Dating requires destructive sampling (in quantities even larger than those required by genetic and collagen based analyses), which is not very appealing to museum conservators, particularly when dealing with worked artefacts. Fortuitously, continuous developments in sample preparation and radiocarbon dating techniques have considerably reduced the required sample size, making radiocarbon dating possible using tens of milligrams of bone [START_REF] Cersoy | Radiocarbon dating minute amounts of bone (3-60 mg) with ECHoMICADAS[END_REF][START_REF] Cucchi | Tracking the Near Eastern origins and European dispersal of the western house mouse[END_REF][START_REF] Rofes | Detecting stratigraphical issues using direct radiocarbon dating from small-mammal remains[END_REF], depending on the state of bone preservation.

Accurate dating of marine animals is difficult because of the marine reservoir effect [START_REF] Ascough | Methodological approaches to determining the marine radiocarbon reservoir effect[END_REF][START_REF] Olsson | Content of 14C in Marine Mammals from Northern Europe[END_REF]. As CO2 from the atmosphere enters the ocean by exchange at the water surface, it takes time for it to distribute evenly across the entire water body. In the deep ocean, radiocarbon will start to decay, leading to lower 14 C concentrations, while new radiocarbon is constantly added via the surface. This will inevitably result in a mixture of both young and old water (i.e., waters with varying 14 C concentrations) being present in the oceans. The global average reservoir age is 400 years, although there are spatial and temporal variations. Where the animals feed (e.g., open water, at depth close to the sea floor, or in coastal areas) and what they feed on (their food sources will also display typical feeding behavior), will all have an effect on the single radiocarbon measurement produced by a bone from a marine animal. As such, knowing the whale species of a particular bone can add useful information on their feeding habitat and choice of prey. Whale species feeding in fresh or brackish water can be even more challenging to radiocarbon date, as their food sources can also include terrestrial carbon originating from specific types of bedrock, such as limestone, which can bring very old (dead) radiocarbon to the equation [START_REF] Philippsen | The freshwater reservoir effect in radiocarbon dating[END_REF]. In addition, whales can make long journeys or seasonally migrate, during which they may supplementary feed in locations with different 14 C signatures [START_REF] Eisenmann | Radiocarbon as a Novel Tracer of Extra-Antarctic Feeding in Southern Hemisphere Humpback Whales[END_REF].

Additionally, when archaeological whale bone is found in close context with terrestrial material, it can be extremely useful for paleoceanographers to investigate ocean reservoir offsets in the past [START_REF] Ascough | Methodological approaches to determining the marine radiocarbon reservoir effect[END_REF][START_REF] Reimer | Marine radiocarbon reservoir corrections for the midto late Holocene in the eastern subpolar North Atlantic[END_REF], as it enables the comparison of a fully marine with a fully terrestrial sample that should be of the same age (i.e. the samples died at roughly the same time). Thus, the difference in age can be assigned to the reservoir effect, which, for example, has been done using closely associated marine mollusks shells and terrestrial material in northern Iberia (e.g. [START_REF] García-Escárzaga | Bayesian estimates of marine radiocarbon reservoir effect in northern Iberia during the Early and Middle Holocene[END_REF][START_REF] Soares | Marine Radiocarbon Reservoir Effect in Late Pleistocene and Early Holocene Coastal Waters off Northern Iberia[END_REF].

Evidence of utilization

Modification and marks in whale bones (together with other contextual archaeological information) can shed light on their utilization and thus value to past communities. Some bones were used whole as structural elements, such as ribs and skulls for components of housing or fences, vertebrae as stools or tables, scapulae as shovels. For example, huts made of whale ribs, and whale vertebrae probably used as stools, were excavated at a pre-19 th century seasonal pastoralist Khoisan camp on the coast of Namibia [START_REF] Smith | The invisible whale[END_REF]. In Nuu-chahnulth sites in Vancouver Island (British Columbia, Canada), whale bones architectural elements served both for retaining walls and diverting water away from the houses (Huelsbeck, 1994, 289;McMillan et al., 2012, 83-86). Eighteenth Century Kamchatka Indigenous peoples used the lower jaws to make sliders for their sledges and vertebrae as mortars (D'Auteroche, 1768). The use of unmodified bonesmostly vertebrae-of large whale species as passive tools such as anvils, mortars and cutting boards is documented in the Mediterranean Basin at least from the Bronze Age to the Antiquity [START_REF] Benito | Ballenas y delfines en la prehistoria peninsular. los restos de cetáceo de la Vital (Gandia)[END_REF]; for example, four vertebra of sperm whale (Physeter macrocephalus), excavated at the 6 th -5 th century BC Phoenician site of Motya in Sicily, were used as platforms for breaking purple-dye shells with stone tools [START_REF] Reese | Whale Bones and Shell Purple-Dye at Motya (western Sicily, Italy)[END_REF].

Other whale bones were instead processed to produce a large diversity of tools and utensils (e.g. [START_REF] Betts | The Mackenzie Inuit whale bone industry: raw material, tool manufacture, scheduling, and trade[END_REF][START_REF] Christensen | L' industrie osseuse des chasseurscueilleurs: le cas des nomades marins de Patagonie et Terre de Feu[END_REF][START_REF] Clark | Whales as an Economic Factor in Prehistoric Europe[END_REF][START_REF] Cunliffe | Whales and whale bone technology in New Zealand prehistory[END_REF][START_REF] Margaris | Reconsidering Raw Material Selection[END_REF], often related to weaponry (projectile points and harpoon heads). Sometimes their specific use can be readily identified in terms of functionality based on design, as in the case of the previously mentioned Magdalenian projectile points [START_REF] Lefebvre | Interconnected Magdalenian societies as revealed by the circulation of whale bone artefacts in the Pyreneo-Cantabrian region[END_REF][START_REF] Pétillon | Circulation of whale-bone artifacts in the northern Pyrenees during the late Upper Paleolithic[END_REF] and other weaponry, common tools such as scrapers or wedges, or even gaming pieces [START_REF] Hennius | Whalebone Gaming Pieces: Aspects of Marine Mammal Exploitation in Vendel and Viking Age Scandinavia[END_REF] and other context-specific tools such as stone pressure flakers [START_REF] Christensen | L' industrie osseuse des chasseurscueilleurs: le cas des nomades marins de Patagonie et Terre de Feu[END_REF], wool weaving combs and peat spades [START_REF] Mulville | The role of cetacea in prehistoric and historic Atlantic Scotland[END_REF].

However, some larger artefacts can be harder to interpret, as their cut marks and reductions can be suggestive of either butchery or intentional modification, leading to more tenuous functional suggestions [START_REF] Bernal-Casasola | Ancient whale exploitation in the Mediterranean: the archaeological record[END_REF][START_REF] Mulville | The role of cetacea in prehistoric and historic Atlantic Scotland[END_REF]. This ambiguity can potentially mean that such elements remain largely ignored in archaeological studies.

In addition, some worked whale bones are the byproducts of processing, rather than worked artefacts. For example, a large quantity of whale bone cutting fragments excavated from a 15 th century Māori coastal fishing camp in New Zealand, reduced longitudinally using a chipping technique, likely reflects a first step in a bone processing industry, whereby large bones were reduced into portions that could be easily transported inland, where they could then be further manufactured into artefacts [START_REF] Cunliffe | Prehistoric Whale Bone Technology in Southern New Zealand[END_REF]. Similarly, at the McKinley Inuit sites, the large majority of the whale bone sample recovered is made of waste material from the different stages of the tool-making process [START_REF] Betts | The Mackenzie Inuit whale bone industry: raw material, tool manufacture, scheduling, and trade[END_REF].

A less known but potentially significant past use of whale bones is as fuel. Whales are unique among vertebrates because of the enormous amount of oil their bones contain. Lipid content varies depending on the species and type of bone, but can reach more than 50% of the bone composition [START_REF] Higgs | Bones as biofuel: a review of whale bone composition with implications for deep-sea biology and palaeoanthropology[END_REF]. Bones could have been broken to facilitate the release of oil they contain (by dripping) or to be used directly as fuel (e.g. [START_REF] Hambrecht | Whale bone as fuel at an inland farm in early modern iceland[END_REF][START_REF] Huelsbeck | The utilization of Whales at Ozette[END_REF]cited by McMillan, 2015;[START_REF] Monks | An oil utility index for whale bones[END_REF][START_REF] Monks | The Cultural Taphonomy of Nuu-chahnulth Whale Bone Assemblages[END_REF]).

In the case of odontocetes, teeth can also be found in the archaeological record, either whole (e.g. carved sperm and pilot whale teeth dated to the Magdalenian from Spain; Corchón-Rodríguez and [START_REF] Corchón-Rodríguez | Nuevas evidencias de restos de mamíferos marinos en el Magdaleniense: los datos de La Cueva de Las Caldas[END_REF] or carved into objects (e.g., ivory buttons from Chalcolithic Portugal, [START_REF] Schuhmacher | The use of sperm whale ivory in Chalcolithic Portugal[END_REF]. Scrimshaw -engraved handicrafts by mariners-frequently employed sperm whale (Physeter macrocephalus) teeth and bone [START_REF] Pichler | Nondestructive DNA extraction from sperm whale teeth and scrimshaw[END_REF].

Information regarding the whale populations

Analyses of whale bone can also reveal information about the ancient populations of the respective whale species. Carbon and nitrogen stable isotope analyses can shed light on the animals' diet and feeding habits, with δ 15 N values reflecting the trophic level and δ 13 C values differing between pelagic and benthic/coastal feeding areas [START_REF] Newsome | Using stable isotope biogeochemistry to study marine mammal ecology[END_REF][START_REF] Riccialdelli | Isotopic assessment of prey and habitat preferences of a cetacean community in the southwestern South Atlantic Ocean[END_REF][START_REF] Sharp | Principles of Stable Isotope Geochemistry[END_REF]. Erreur ! Signet non défini.Carbon and nitrogen stable isotope analysis of whale teeth have also been used to investigate past nursing strategies [START_REF] Walker | Dietary Studies of Marine Mammals Using Stable Carbon and Nitrogen Isotopic Ratios of Teeth[END_REF]. Oxygen stable isotope analysis on bone can in some cases shed light on population structures, and on the use of feeding grounds and migration (e.g. [START_REF] Vighi | Stable isotope analysis and fin whale subpopulation structure in the eastern North Atlantic[END_REF], although the latter is likely to be more informative combined with hydrogen stable isotope analysis of baleen plates (e.g. deHart and Picco, 2015).

Genetic analyses can be used to distinguish different whale populations, as well as the sex of individuals (e.g. [START_REF] Morin | Interfamilial characterization of a region of the ZFX and ZFY genes facilitates sex determination in cetaceans and other mammals[END_REF][START_REF] Pichler | Nondestructive DNA extraction from sperm whale teeth and scrimshaw[END_REF][START_REF] Szpak | Sexual differences in the foraging ecology of 19th century beluga whales (Delphinapterus leucas) from the Canadian High Arctic[END_REF]. Analysis of ancient DNA (particularly when contrasted with modern DNA) can be used to reconstruct the former genetic diversity, population size, and demographic history, including anthropogenic and/or natural bottlenecks and thus be used to understand the impacts of whaling [START_REF] Alter | Prewhaling genetic diversity and population ecology in eastern Pacific gray whales: insights from ancient DNA and stable isotopes[END_REF][START_REF] Béland | Using pre-and postexploitation samples to assess the impact of commercial whaling on the genetic characteristics of eastern North Pacific gray and humpback whales and to compare methods used to infer historic demography[END_REF]. For example, in their analysis of Northeast Pacific humpback and gray whale populations, [START_REF] Béland | Using pre-and postexploitation samples to assess the impact of commercial whaling on the genetic characteristics of eastern North Pacific gray and humpback whales and to compare methods used to infer historic demography[END_REF] were able to demonstrate that commercial whaling had relatively little effect on the overall mitochondrial or nuclear genetic diversity indices of these two species. Nevertheless, patterns of genetic characteristics were affected, with coalescent-based modelling identifying a genetic bottleneck arising from commercial hunting of these species. These studies can inform ongoing conservation efforts, by providing baselines to inform recovery targets. In addition, quantification of distinct mtDNA haplotypes or microsatellite profiles can provide insight into the minimum number of individuals found in a given area (e.g. [START_REF] Arndt | Ancient DNA Analysis of Northeast Pacific Humpback Whale (Megaptera Novaeangliae)[END_REF][START_REF] Béland | Using pre-and postexploitation samples to assess the impact of commercial whaling on the genetic characteristics of eastern North Pacific gray and humpback whales and to compare methods used to infer historic demography[END_REF][START_REF] Evans | Using combined biomolecular methods to explore whale exploitation and social aggregation in hunter-gatherer-fisher society in Tierra del Fuego[END_REF], which is often otherwise exceedingly difficult to estimate in highly fragmentary assemblages.

Evidence from beyond the bones

Until recently bones, and teeth for odontocetes, were the only whale remains that could dependably be found in the archaeological record. However, advances in analytical methods are now bringing to light other direct and indirect evidence of ancient whale utilization. In this section, we start by reviewing historical and ethnological records of ancient uses, processing, storage and transportation of whale products, as context for archaeological records other than whale bones. We then review the main types of such records and the information that can be derived from them: whale barnacles; lipids and protein residues; environmental DNA; baleen; and isotopic analyses of the bones of human consumers.

The many uses of whales

Ethnological records reveal the wide diversity of whale uses. For example, Jean Chappe d'Auteroche (1768) described how the Indigenous peoples in Kamchatka took advantage of every part of the gray whales they hunted (D'Auteroche, 1768). Meat was dried and eaten, with the tongue and fins particularly appreciated. Blubber was also smoked and eaten, as well as rendered into oil for illumination and for heating. The whale skin was dried and beaten, and then transformed into shoe soles and belts. The baleen was used to sew canoes and to make nets for fishing and hunting. Intestines were used as containers, bladders as floaters, as well as to make waterproof clothing. Nerves and veins were converted into ropes. Careful reading of ancient texts can also bring these whale parts to life. Two passages could be examples of such textual evidence in the Mediterranean: Petronius (Satyricon, chapter 21, transl. Arrowsmith and Arrowsmith 1959) mentions a "whalebone-cane" plausibly made of baleen, and Aelian (On the Characteristics of Animals, book 17, section 6, transl. [START_REF] Scholfield | Aelian -on the Characteristics of Animals[END_REF] explains that sinews of large sea monsters are useful for the stringing of harps and other instruments, and even for engines of war.

A single large whale provides tons or tens of tons of meat and blubber, and a main challenge is to quickly preserve these products for future consumption before they spoil. In the Arctic, cellars dug in the permafrost could have been used to freeze them year round, as is still done today by the Iñupiat whalers in Alaska [START_REF] Krupnik | The Material Culture of Inupiat Whaling: An Ethnographic and Ethnohistorical Perspective[END_REF]. Elsewhere, available options for preserving whale products included smoke drying (e.g. in Kamchatka, 18 th century; D'Auteroche 1768), salting (e.g., in Medieval Europe; [START_REF] Szabo | Monstrous Fishes and the Mead-Dark Sea: Whaling in the Medieval North Atlantic[END_REF], and drying by burying in the sand (e.g., 17 th -19 th century south-western Africa; [START_REF] Smith | The invisible whale[END_REF].

Blubber could also have been rendered into oil, which keeps for a long time if stored properly, and has for millennia been a major output of whale exploitation [START_REF] Monks | An oil utility index for whale bones[END_REF]. Simple methods to render oil included letting it drip from the blubber (e.g., 18 th century Olioutores, Kamtchatka; D'Auteroche 1768) and heating in the sun (e.g., 18 th century Chango Indians from Northern Chile; Vazquez de Espinosa 1862). More sophisticated methods involved using hot stones (e.g., Norway Iron Age, [START_REF] Heron | Application of lipid biomarker analysis to evaluate the function of 'slab-lined pits' in Arctic Norway[END_REF], boiling (e.g. 17 th century Khoi, South Africa; [START_REF] Smith | The invisible whale[END_REF], and furnaces (e.g. 19 th century Basque whalers; Duhamel du Monceau 1782, figure 1). Oil was used directly as food (e.g., Changos, Vazquez de Espinosa 1862; Khoisan, [START_REF] Smith | The invisible whale[END_REF], as fuel for illumination and heating (e.g., Kamchatka Indians; D'Auteroche 1768), in the impermeabilization of ropes, leather or boats, as well as in the manufacture of products like soap and paint (e.g. 17 th century Europe, [START_REF] Viret | Les us, et coutumes de la mer[END_REF]. Spermaceti, a liquid wax found in the head cavity of sperm whales, was particularly prized in Medieval Europe as a lighting fuel because it burns without producing smoke and odour.

Dried blubber or meat could have been transported without particular containers, including as packs directly attached to the body (e.g., hunter-gatherers of southern South America; Bove 1883 cited by [START_REF] Lefèvre | Chasseurs terrestres, chasseurs marins ? L'exploitation des ressources animales dans le locus 1 / Cazadores terrestres, cazadores marítimos ? Explotación de los recursos animales en el locus 1[END_REF] Baleen plates are the filtering system used by baleen whales (Mysticeti) to remove prey from the water. Consisting of long strains of keratin, they vary in size and physical properties depending on the whale species. Right whales, and even more so bowhead whales, have particularly long baleen plates (up to 4 m in the latter) that have long been appreciated as a strong and flexible material. Traditional uses of baleen by Arctic whalers included weaved baskets and mats, nets, weirs, fishing lines, and snares [START_REF] Lauffenburger | Baleen in museum collections: its sources, uses, and identification[END_REF] but also some kinds of knives, end-and side blades, and lashing strips for tools and weapons, as well as for drying racks [START_REF] Mathiassen | Archaeology of the Central Eskimos[END_REF]. In historic times, baleen was a prized commodity from commercial whaling. Using heat and pressure, baleen could be fashioned into complex shapes, and was incorporated into objects like collars and corsets, buggy whips, brushes and bed springs.

The degradability of meat, blubber, oil and baleen means that evidence of their use is largely absent from the archaeological record [START_REF] Smith | The invisible whale[END_REF]. Nonetheless, direct and indirect evidence may sometimes remain, as reviewed below.

Whale barnacles

The oldest archaeological evidence for the transport and use of whale blubber is unexpectedly provided by whale barnacles, i.e., cirripede crustaceans that live deeply embedded in the skin of some whale species. Because barnacles can only be removed by cutting the skin, their presence in archaeological contexts associated with past human settlements is evidence of the transport of skin and its underlying blubber, and potentially also meat, for human use [START_REF] Kandel | Scavenging and processing of whale meat and blubber by Later Stone Age people of the Geelbek Dunes, Western Cape Province, South Africa[END_REF]. Furthermore, as certain barnacle species are quite specific to particular whales [START_REF] Hayashi | A checklist of turtle and whale barnacles (Cirripedia: Thoracica: Coronuloidea)[END_REF][START_REF] Mcmillan | Whales and Whalers in Nuu-chahnulth Archaeology[END_REF], they can be used to understand which whale species was likely used. For example, Tubicinella major and Cetopirus complanatus are barnacle species found today almost exclusively in association with right whales [START_REF] Hayashi | A checklist of turtle and whale barnacles (Cirripedia: Thoracica: Coronuloidea)[END_REF], so their finding in a Magdalenian site in Spain is strong evidence that right whales were exploited by contemporary maritime-oriented forager human groups [START_REF] Álvarez-Fernández | Occurrence of whale barnacles in Nerja Cave (Málaga, southern Spain): Indirect evidence of whale consumption by humans in the Upper Magdalenian[END_REF]. Similarly for the finding of C. complanatus in the late Middle Pleistocene human settlement of Pinnacle Point in South Africa [START_REF] Collareta | Cetopirus complanatus (Cirripedia: Coronulidae) from the late Middle Pleistocene human settlement of Pinnacle Point 13B (Mossel Bay, South Africa)[END_REF]. Findings of Coronula diadema in Later Stone Age sites in southern Africa point instead to the likely use of humpback whales (Megaptera novaeangliae; [START_REF] Kandel | Scavenging and processing of whale meat and blubber by Later Stone Age people of the Geelbek Dunes, Western Cape Province, South Africa[END_REF]. Coronula reginae, associated to humpback whales, were discovered in the Nuu-chahnulth sites (northern coast of Vancouver Island, Canada), together with Cryptolepas rachianecto, specific to gray whales [START_REF] Wessen | An account of the Ozette shellfish taxa[END_REF]cited by McMillan, 2015).

Lipid and protein residues

Surfaces that came into contact with whale meat, blubber or oil may still hold residues that reveal their previous uses. For example, characterization of organic residues in fragments of large ceramic containers, taken from a 16 th -17 th century warehouse in Lekeitio (Basque Country, northern Spain), indicate that these were whale oil containers [START_REF] Blanco-Zubiaguirre | GC-MS and HPLC-ESI-QToF characterization of organic lipid residues from ceramic vessels used by Basque whalers from 16th to 17th centuries[END_REF].

In another study, chemical analyses of fatty acids in cemented organic residues and charcoal samples of Late Iron Age slab-lined pits in Arctic Norway revealed that they were used for rendering oil from the blubber of marine mammals [START_REF] Heron | Application of lipid biomarker analysis to evaluate the function of 'slab-lined pits' in Arctic Norway[END_REF]). These pits -elliptical or rectangular depressions (2-4 m in length, and 1-2 m in width, up to 1 m in depth), often lined with stone slabs -were initially interpreted as graves or as turf houses. Signs of thermal influence in the slabs, as well as abundant finds of charcoal and fire-cracked rocks are consistent with an oil rendering method involving alternate layers of blubber and heated stones. More than 700 pits are known, usually in groups up to 5, indicative of a widespread use of marine mammal oil, yet the only known faunal remain is a single vertebra of a small whale [START_REF] Heron | Application of lipid biomarker analysis to evaluate the function of 'slab-lined pits' in Arctic Norway[END_REF].

Besides containers, other promising artefacts to study are those associated with illumination. For example, analyses of Mesolithic shallow oval bowls in the Baltic region (a combination of lipid biomarker and bulk and single-compound carbon isotope analyses) demonstrated their past used as oil lamps fueled with marine animal fat [START_REF] Heron | Illuminating the Late Mesolithic: residue analysis of "blubber" lamps from Northern Europe[END_REF][START_REF] Nilsen | Marine Mammal Train Oil Production Methods: Experimental Reconstructions of Norwegian Iron Age Slab-Lined Pits[END_REF]). However, the degradation of fats and oils within archaeological pottery vessels, particularly if burnt, makes it difficult to specify the species or group of species concerned. For example, the very few chemical analyses of Antique oil lamps in the Mediterranean region have sometimes revealed the presence of animal fat but without being able to distinguish between bovine, ovine, fish or marine mammal origins [START_REF] Copley | Gas chromatographic, mass spectrometric and stable carbon isotopic investigations of organic residues of plant oils and animal fats employed as illuminants in archaeological lamps from Egypt[END_REF][START_REF] Kimpe | Analysis of oil used in late Roman oil lamps with different mass spectrometric techniques revealed the presence of predominantly olive oil together with traces of animal fat[END_REF]. Accordingly, the chemical analysis of burnt fat and other organic residues remains particularly challenging and requires more developed methods to reveal their precise origin [START_REF] Heron | Application of lipid biomarker analysis to evaluate the function of 'slab-lined pits' in Arctic Norway[END_REF][START_REF] Lucquin | Untangling complex organic mixture in prehistoric hearths[END_REF]. Analysis of entrapped proteins on ceramic vessels may potentially provide greater taxonomic insights as proteins are often more species-/genus-/family-specific while lipids and isotope composition are dependent on dietary and environmental factors. For example, proteomic analysis of Iñupiat potsherd fragments from Alaska revealed seal myoglobin (muscle) and hemoglobin (blood) proteins [START_REF] Solazzo | Identification of protein remains in archaeological potsherds by proteomics[END_REF]. As specific proteins are frequently expressed only in particular tissues (e.g., blood, skin, muscle), this approach may provide greater insights into particular parts of whales processed or stored in ceramic vessels.

Environmental DNA

High-value portions of whales like meat and blubber tend to degrade rapidly, but may leave biomolecular traces within archaeological sediments. For example, [START_REF] Seersholm | DNA evidence of bowhead whale exploitation by Greenlandic Paleo-Inuit 4,000 years ago[END_REF] analyzed sedimentary DNA (sedaDNA) within midden deposits from Greenland, detecting the exploitation of bowhead whales dating back 4,000 years. Indeed, they found that in some Paleo-Indian contexts, marine mammals, and particularly bowhead whale, dominated the recovered DNA profiles, suggesting that the importance of whale products was far greater than would be presumed based on zooarchaeological remains alone.

Baleen

The degradability of baleen means that archaeological specimens are rare, being mostly found in association with Arctic whaling cultures (e.g. [START_REF] Sinding | Minimally destructive DNA extraction from archaeological artefacts made from whale baleen[END_REF][START_REF] Solazzo | Molecular markers in keratins from Mysticeti whales for species identification of baleen in museum and archaeological collections[END_REF], although there are exceptions (e.g. 12 th century fragments found in waterlogged organic midden layers in Scotland, [START_REF] Moffat | The use of Baleen for Arms, Armour and Heraldic Crests in Medieval Britain[END_REF]. More recent baleen artefacts can be found in museum collections (e.g., 18th-19th century corset busks; an early 20 th century Kinguktuk basket, [START_REF] Lauffenburger | Baleen in museum collections: its sources, uses, and identification[END_REF].

The analysis of baleen can uncover valuable information regarding the identity of the species as well as of contemporary environmental conditions. For example, [START_REF] Sinding | Minimally destructive DNA extraction from archaeological artefacts made from whale baleen[END_REF] successfully extracted and amplified DNA from small fragments of baleen dated back more than 4,000 years ago, highlighting its potential value as a source of ancient DNA for studies on whale population genetics. More recently, application of a keratin peptide mass fingerprinting technique to 29 fragments of unidentified and partially degraded samples of baleen excavated from prehistoric archaeological sites in Labrador (Canada), demonstrated a dominance of bowhead whale in the archaeological assemblage [START_REF] Solazzo | Molecular markers in keratins from Mysticeti whales for species identification of baleen in museum and archaeological collections[END_REF]. Baleen can also provide a valuable archive of the individual's lifetime, given that it forms continuously over the years and becomes metabolically inert after synthesis. Indeed, stable isotope analyses (of carbon, nitrogen, oxygen and hydrogen) can reveal dietary changes across time, shedding light on environmental changes experienced by the individuals [START_REF] Bentaleb | Foraging ecology of Mediterranean fin whales in a changing environment elucidated by satellite tracking and baleen plate stable isotopes[END_REF][START_REF] Schell | Declining carrying capacity in the Bering Sea: Isotopic evidence from whale baleen[END_REF]. For example, sequential carbon and nitrogen stable isotope analysis on baleen revealed there was limited exchange in foraging grounds between two subpopulations of fin whale in the Atlantic and Mediterranean [START_REF] Giménez | Isotopic evidence of limited exchange between Mediterranean and eastern North Atlantic fin whales[END_REF], while oxygen and hydrogen stable isotope analyses of baleen plates have been used to investigate the migratory routes of bowhead whales, also highlighting the value of baleen as a record of historical sea ice concentration and Arctic climate [START_REF] Dehart | Stable oxygen and hydrogen isotope analyses of bowhead whale baleen as biochemical recorders of migration and arctic environmental change[END_REF]. Depending on the state of preservation, even certain hormones can be retrieved from whale baleen, which can show insights into the individuals' stress levels [START_REF] Rolland | Chronic stress from fishing gear entanglement is recorded in baleen from a bowhead whale ( Balaena mysticetus )[END_REF]. Baleen preserved in museum collections can thus provide a rich isotopic and genetic archive for whales living within the last few centuries.

Isotope analyses of human bones

If whales were an important part of human diet, this will have left a signature in human bones. Indeed, the nitrogen stable isotope values of human bone collagen reflects the trophic level of diets [START_REF] Kelly | Stable isotopes of carbon and nitrogen in the study of avian and mammalian trophic ecology[END_REF], which combined with carbon stable isotope values can be used to distinguish marine from terrestrial consumption [START_REF] Chisholm | Stable-Carbon Isotope Ratios as a Measure of Marine Versus Terrestrial Protein in Ancient Diets[END_REF]. For example, stable isotope analyses of Eastern Arctic foragers from the Thule period indicates that they relied on bowhead whales for circa 12% of their dietary intake [START_REF] Coltrain | Sealing, whaling and caribou: the skeletal isotope chemistry of Eastern Arctic foragers[END_REF].

How the whale might have been obtained

Given a whale bone in an archaeological context, it is inevitable to wonder: was it obtained from a whale found naturally stranded on a beach? Or was it the product of ancient whaling? This section provides context for helping researchers to answer this question. Firstly, we draw on ethnographic and historical sources to overview the diversity of methods humans have developed for exploiting whales. Secondly, we discuss the different types of evidence that whaling can leave, including written sources, graphical representations, archaeological records of whaling implements, and evidence of transformation and use of whale products. Finally, we highlight the need to identify whale bones to the species level to discuss a possible active capture of whales.

Naturally stranded or drift whales can be found along all coastlines, although they are more frequent in some regions ('stranding hotspots'; [START_REF] Brabyn | Oceanography and coastal topography of herd-stranding sites for whales in New Zealand[END_REF] and in particular seasons [START_REF] Evans | Periodic variability in cetacean strandings: links to large-scale climate events[END_REF]. Most stranding events correspond to single individuals, but mass stranding of multiple individuals (of the same or different species) can also occur. These events are unpredictable, but they are sufficiently frequent for coastal communities to have developed techniques to take advantage of the abundant resources they bring (e.g., well-developed regulations in Medieval Norse societies regarding rights over beached whales; Lindquist 1997). Freshly deceased whales provide vast amounts of edible meat, blubber as well as other useful materials (veins, tendons, intestines…). Dead whales decompose very rapidly, and so stop being edible [START_REF] Huelsbeck | Whaling in the precontact economy of the central Northwest coast[END_REF], but bones can remained available for exploitation for years or decades (although they progressively lose their oil, and hence their value as fuel).

The active killing of whales requires preparedness and cooperation, but not necessarily sophisticated technologies. Maritime cultures around the world have developed a wide variety of whaling methods, in regions as diverse as the Arctic, both coasts of the North Pacific, Atlantic, South Pacific, Indonesia, the Lesser Antilles, and Equatorial Guinea [START_REF] Reeves | The origins and character of 'aboriginal subsistence' whaling: a global review[END_REF][START_REF] Reeves | A taxonomy of world whaling[END_REF].

The first step in whaling is detecting the whale. Before the 16 th century (when the navigation technology necessary for offshore whaling developed), only those whales that naturally came very close to the shore were accessible to early whalers.

Once a whale is detected, a much more substantial challenge is to kill it and to retrieve it, particularly for larger whales. One approach involves forcing the whale to strand, which some whalers did by surrounding it with small boats and using nets and/or making much noise to direct it to shore (e.g., gray whale hunting in early 20 th Century, Korea Andrews 1916; and in 18 th Century Kamchatka, D'Auteroche 1768; ancient and modern pilot whale hunting in the Faeroes, [START_REF] Lindquist | Peasant fisherman whaling in the Northeast Atlantic area, CA 900-1900 AD[END_REF]. Other options involve the use of traps, natural or artificial. For example, medieval Norse whalers took advantage of natural traps like closed bays or rivers: of a whale entered it, they would then prevent it from leaving [START_REF] Lindquist | Peasant fisherman whaling in the Northeast Atlantic area, CA 900-1900 AD[END_REF]. In estuary of the Saint Laurence River, 18 th -20 th century whalers installed wooden weirs that allowed whales in during the high tide, while becoming barriers in the low tide [START_REF] Reeves | Whaling in the St. Lawrence[END_REF], a similar mechanism to that of Medieval whaling stone dams in Norway [START_REF] Lindquist | Peasant fisherman whaling in the Northeast Atlantic area, CA 900-1900 AD[END_REF]. The whaling scenes in the magnificent Bangudae petroglyphs in southeastern Korea (date undetermined) seem to represent the use of nets and wooden weirs to capture large baleen whales [START_REF] Lee | Chasseurs de baleines : La frise de Bangudae[END_REF]. In the Arctic, natural traps also occur as gaps in the sea ice, with whales that surface to breathe being vulnerable to attack (with harpoons) from above.

More active forms of killing whales necessarily involved the use of piercing instruments. Lances or arrows hurled with sufficient energy to pierce deeply across the blubber would cause the whale to die in a few days [START_REF] Lindquist | Peasant fisherman whaling in the Northeast Atlantic area, CA 900-1900 AD[END_REF], a process sometimes accelerated through the use of poison (e.g., among Medieval Norse and the North Pacific whalers; [START_REF] Reeves | A taxonomy of world whaling[END_REF]). However, this method does not necessarily guarantee recovery of the whale, as the dead animal could sink in the high seas, or strand far away. To increase the likelihood of recovery, medieval Norse [START_REF] Lindquist | Peasant fisherman whaling in the Northeast Atlantic area, CA 900-1900 AD[END_REF] and Alaskan Inuit [START_REF] Reynolds | Ownership Marks and Social Affinity in Northwest Alaska during Late Prehistoric Times[END_REF] whalers marked their lances individually as a means of claiming rights over whales subsequently found by others.

Harpoons -i.e., barbed projectiles with an attached rope -are a technological solution to improve the odds of recovering the whale. In the case of the Makah in the northeastern Pacific [START_REF] Scammon | The Marine Mammals of the North-Western Coast of North America, Described and Illustrated; Together with an Account of the American Whale-Fishery[END_REF] as well as in the Arctic [START_REF] Reeves | A taxonomy of world whaling[END_REF], whalers approached whales in small boats, and handthrew harpoons attached to inflated sealskin floats (Figure 3). The floats created drag on the harpooned whale's escape and signaled its presence as it resurfaced, allowing whalers to approach it again, throw additional harpoon-line-float arrays, until the whale was sufficiently weakened to be killed with hand lances (Figure 3). The first reliably dated whaling scene, carved in a walrus ivory piece dated to 3,000 BP from the Bering region [START_REF] Pringle | Signs of the First Whale Hunters[END_REF], may illustrate this method, showing the use of sealskin floats in the capture of a (likely) gray whale. It represents the boat fastened to the whale by a rope, which may indicate that it too was being used to create drag, apparently supported by a drogue (a device trailing behind to provide additional resistance). Dragging behind a large whale would however have been a very dangerous endeavor (as a diving whale can easily sink a small boat), and so this scene may have represented the final stages of whaling (the boat fastened to the whale only after the animal became tired). Medieval Basque whalers developed a method that explicitly required them to remain fastened to the whale. They used lines so long that the harpooned whale could sprint away or even dive without endangering the boat, while allowing the whalers not to lose track of it. By following the line, the whalers could then re-approach to attach more harpoons and lines. Once the whale was very tired, the whalers approached it and killed it with long lances (Duhamel du Monceau, 1782). This very efficient method became the foundation of European (and subsequently American) industrial whaling [START_REF] Reeves | A taxonomy of world whaling[END_REF]. Another highly valuable line of evidence comes from graphical representations of whaling, particularly when they realistically illustrate the quintessential scene of a whaling boat linked to a harpooned whale by a line. This is the case with the above-mentioned carved walrus ivory pieve from the Bering region [START_REF] Pringle | Signs of the First Whale Hunters[END_REF], whaling scenes represented in the Pegtymel petroglyphs in Chukotka, Russia [START_REF] Dikov | Mysteries in the Rocks of Ancient Chukotka (Petroglyphs of Pegtymel)[END_REF], and in the El Medaño rock art from Northern Chile [START_REF] Ballester | Rock art, marine hunting and harpoon devices from the Atacama Desert coast, northern Chile[END_REF]. Less typical (and thus more ambiguous) are the realistic representations of whales superposed by nets and wood weirs in the Bangudae petroglyphs of South Korea, which have also been considered as evidence of past whaling [START_REF] Lee | Chasseurs de baleines : La frise de Bangudae[END_REF]. Graphical representations do not necessarily exist, or they may have existed but not have survived, or they may not appear as obvious representations of whaling to a modern observer.

Large harpoons can be strong evidence of past whaling, and have been found in several archaeological contexts associated with whaling cultures (e.g., [START_REF] Monks | Nuuchah-nulth whaling: Archaeological insights into antiquity, species preferences, and cultural importance[END_REF][START_REF] Ballester | Rock art, marine hunting and harpoon devices from the Atacama Desert coast, northern Chile[END_REF]). However, harpoons were also used for the capture of other large marine animals (e.g., seals, turtles, sharks, manta rays) and so their presence is not by itself irrefutable evidence of whaling [START_REF] Losey | Opportunistic whale hunting on the southern Northwest Coast: ancient DNA, artifact, and ethnographic evidence[END_REF]. Conversely, harpoons may also be rare even in deposits from areas where whaling is definitely confirmed; as prized possessions, they were not often discarded, but reused [START_REF] Monks | Nuuchah-nulth whaling: Archaeological insights into antiquity, species preferences, and cultural importance[END_REF]. Furthermore, as discussed in the previous section, harpoons were not used in all whaling methods. Their absence from the archaeological record is thus not a proof that whaling did not take place. More broadly, many of the above-mentioned methods for killing whales involve implements (such as small boats, nets, ropes, stone or wood weirs) that are not necessarily specific to whaling and could have been employed in the context of other fisheries (Bekker-Nielsen and Bernal-Casasola, 2010). With ancient whaling being a coastal activity, the boats involved were not necessarily seaworthy, and may appear unremarkable in the archaeological record, or not at all. For example, the Changos in Northern Chile used inflatable rafts made of sea-lion skins (Vazquez de Espinosa 1862; Ballester 2018).

Given the huge size of whales, it is rare to find direct marks on the whale bones that can indisputably prove that humans captured the live animal. Rare examples include findings of bone points embedded in whale bones, such as in a phalange of a humpback whale found in Oregon [START_REF] Losey | Opportunistic whale hunting on the southern Northwest Coast: ancient DNA, artifact, and ethnographic evidence[END_REF], or in two bones from the Neolithic site Hwangseong-dong in South Korea [START_REF] Yoo | A study on excavated whale form in Ulsan and the capturing[END_REF], providing strong evidence that the animals were actively attacked. More commonly, though, the harpoons merely pierce the blubber without touching any bone. Other methods of whaling (e.g. using traps) would be even less likely to leave such marks on the bones.

Whaling produces very large quantities of whale bone, so in theory it should result in conspicuous archaeological evidence. In practice, though, shore erosion or sea level rise mean that much of it is currently underwater [START_REF] Mcleod | Bowhead whales, and not right whales, were the primary target of 16th-to 17thcentury Basque whalers in the western North Atlantic[END_REF]. Conversely, given that a substantial amount of bone can be obtained from a single stranded whale, a relatively high abundance or frequency of bone artefacts (e.g., evidence of a Magdalenian whale bone industry; [START_REF] Lefebvre | Interconnected Magdalenian societies as revealed by the circulation of whale bone artefacts in the Pyreneo-Cantabrian region[END_REF][START_REF] Pétillon | Circulation of whale-bone artifacts in the northern Pyrenees during the late Upper Paleolithic[END_REF] does not by itself demonstrate past whaling. Nonetheless, an increase in the use of whale bone may indicate an increase in its availability. For example, archaeological records show that nonwhaler Pre-Inuit groups manufactured some utensils from whale bones, such as shovels from scapulae in Ipiutak settlements [START_REF] Larsen | Ipiutak and the Arctic whale hunting culture[END_REF], architectural elements in a Late Dorset semisubterranean structure [START_REF] Ryan | A Late Dorset semi-subterranean structure from the Bell Site (NiNg-2), Ekalluk River, Victoria Island[END_REF], or sled shoes from ribs in Dorset campsites [START_REF] Houmard | Caractérisation chrono-culturelle et évolution du paléoesquimau dans le golfe de Foxe (Canada) : étude typologique et technologique des industries en matières dures d'origine animale[END_REF]. However, the quantity and diversity of whale bone implements increased markedly after the 7 th century A.D. (in the Old Bering Sea and the subsequent Birnirk and Thule culture) to include the manufacture of tools, weapons, and amulets [START_REF] Mason | Out of the Cold: Archaeology on the Arctic Rim of North America[END_REF][START_REF] Mason | Walrusing, whaling and the origins of the Old Bering Sea culture[END_REF][START_REF] Mathiassen | Archaeology of the Central Eskimos[END_REF], indicating that whale hunting became more intensive then.

Whaling would also have produced large amounts of meat and blubber. These products are even less likely than bones to have been preserved in the archaeological record (but see the above-mentioned analysis of environmental DNA by [START_REF] Seersholm | DNA evidence of bowhead whale exploitation by Greenlandic Paleo-Inuit 4,000 years ago[END_REF], but other types of archaeological evidence can come from the structures used to process these products. Such structures are unlikely to have existed if whale use was opportunistic (and thus unpredictable in space and time), but may have been built by communities able to bring whales regularly and predictably to designated coastal points. This is the case of tryworks, brick furnaces built by 16 th -19 th century European and American whalers to render blubber into oil [START_REF] Paterson | The Sleaford Bay Tryworks: industrial archaeology of shore based whaling stations, Flinders University maritime archaeology monograph series[END_REF]. It may also be the case of the hundreds of Iron Age slab-lined pits found in Arctic Norway, for which residue analysis indicates a prior use for rendering oil of marine mammals. However, their presence alone is not evidence of whaling, as they may also have been used for processing other marine animals such as seals. Conversely, the hundreds of large tanks (cetariae) found in coastal Roman archaeological contexts and known from historical records to have been used for processing tuna [START_REF] Trakadas | The archaeological evidence for fish processing in the Western Mediterranean[END_REF] might have also been used to process whales [START_REF] Rodrigues | Forgotten Mediterranean calving grounds of grey and North Atlantic right whales: evidence from Roman archaeological records[END_REF]. Look-out towers for detecting the whales are another type of physical structure associated with whaling (e.g. Basque atalayas; Aguilar 1986) but again they could have served other purposes too (e.g., detecting enemies).

Overall, in the absence of reliable ethnographic records, realistic graphical representations or very large harpoons (ideally all three), it is very difficult to demonstrate that active whaling took place. Accordingly, the tendency among archaeologists is to fall back on the default assumption that whale remains correspond to scavenged animals [START_REF] Losey | Opportunistic whale hunting on the southern Northwest Coast: ancient DNA, artifact, and ethnographic evidence[END_REF]. However, a lack of whaling evidence should not be mistaken for evidence that whaling did not occur.

Species matters

Until quite recently, only those whales that naturally came close to shore (or, in the Arctic regions, close to the edge of the sea-ice) were available to whalers. For this reason, pre-industrial whaling focused mostly on a narrow range of whale species [START_REF] Reeves | The origins and character of 'aboriginal subsistence' whaling: a global review[END_REF]. These include smaller odontocetes, such as beluga (Delphinapterus leucas), narwhal (Monodon monoceros), pilot whales (Globicephala sp.), and killer whales, that are either coastal or approach the coast at predictable seasons. Among the larger whales, the focus was on those that are coastal year round (gray whale) or that come close to shore for calving and during migration (right, bowhead, and humpback whales), with the calves being particularly targeted [START_REF] Aguilar | A review of old Basque whaling and its effect on the right whales (Eubalaena glacialis) of the North Atlantic[END_REF][START_REF] Krupnik | Prehistoric Eskimo whaling in the Arctic: slaughter of calves or fortuitous ecology?[END_REF]Savelle and McCartney, 1991). In a few coastal areas adjacent to deep seas, high-seas species that came very close to shore were taken too (e.g. sperm and Bryde's whales off Lamalera, Indonesia; [START_REF] Barnes | Sea Hunters of Indonesia: Fishers and Weavers of Lamalera[END_REF], but otherwise the technologies for killing and retrieving non-coastal whale species were only developed recently, particularly for Balaenoptera species that are quite fast and sink once killed (which only started being exploited systematically in the late 19 th century; [START_REF] Reeves | A taxonomy of world whaling[END_REF].

For this reason, knowing the identity of whale species (and thus its ecology) is key to contextualizing whale bones found in archaeological contexts, and specifically for talking the question of how the bones might have been obtained. Indeed, whereas all whale species could have been found stranded, only a subset could have plausibly been actively taken with pre-industrial technology [START_REF] Rodrigues | Ancient whale exploitation in the Mediterranean: species matters[END_REF]. Accordingly, archaeological species profiles with an abundance of large, off-shore, fast-swimming species (e.g., fin or blue whale) point towards an opportunistic exploitation of beached animals, whereas a species profile dominated by smaller odontocetes, or slower moving, near-shore baleen whales (e.g., right, gray or humpback whales) provides more support for active acquisition strategies [START_REF] Rodrigues | Ancient whale exploitation in the Mediterranean: species matters[END_REF]. For example, ZooMS and genetic analyses of bones found in Roman and pre-Roman archaeological sites revealed the past presence of two coastal whales species (North Atlantic right whales E. glacialis; gray whales Eschrichtius robustus) in the Gibraltar region, opening the possibility of their exploitation during the Roman period [START_REF] Rodrigues | Forgotten Mediterranean calving grounds of grey and North Atlantic right whales: evidence from Roman archaeological records[END_REF]. Similarly, genetic analyses of bone fragments from archaeological and paleontological sites in New Zealand revealed a diversity of whales species used by precontact New Zealand Māori, including three small toothed whale species (killer whale Orcinus orca; Cuvier's beaked whale Ziphius cavirostris, unspecified Delphinidae) that could have been plausibly actively whaled [START_REF] Seersholm | Subsistence practices, past biodiversity, and anthropogenic impacts revealed by New Zealand-wide ancient DNA survey[END_REF]). On the Northwest Coast of North America, genetic analyses of pre-contact archeological cetacean bones, associated with the traditional whaling culture, the Nuu-chah-nulth, have revealed a persistent focus on gray and humpback whales dating back 4,000 years [START_REF] Arndt | Ancient DNA Analysis of Northeast Pacific Humpback Whale (Megaptera Novaeangliae)[END_REF][START_REF] Béland | Using pre-and postexploitation samples to assess the impact of commercial whaling on the genetic characteristics of eastern North Pacific gray and humpback whales and to compare methods used to infer historic demography[END_REF] -the same species identified in oral histories and documented in ethnographic whaling accounts [START_REF] Drucker | The Northern and Central Nootkan tribes[END_REF][START_REF] Monks | Nuuchah-nulth whaling: Archaeological insights into antiquity, species preferences, and cultural importance[END_REF].

Biomolecular analyses may also confirm or refute hypotheses regarding the dominant target species within historic time frames. For example, right whales were believed to be the dominant species taken by Basque whalers in the Western Cantabrian Sea in the 13 th -18 th centuries, and this has been confirmed by analyses of cetacean remains from whaling harbours [START_REF] Rey-Iglesia | The genetic history of whaling in the Cantabrian Sea during the 13th-18th centuries: Were North Atlantic right whales (Eubalaena glacialis) the main target species?[END_REF]. In contrast, whereas right whales were also believed to have been the focus of 16 th -17 th century Basque whaling, in the Strait of Belle Isle and Gulf of St. Lawrence, biomolecular analyses of bones revealed instead a strong focus on bowhead whales [START_REF] Mcleod | Bowhead whales, and not right whales, were the primary target of 16th-to 17thcentury Basque whalers in the western North Atlantic[END_REF][START_REF] Rastogi | Genetic analysis of 16th-century whale bones prompts a revision of the impact of Basque whaling on right and bowhead whales in the western North Atlantic[END_REF].

Conclusions

The general invisibility of whales in the archaeological record makes the finding of isolated whale bone artefacts difficult to interpret: are they the outcome of an opportunistic use of beached animals, or the last vestiges of an ancient close connection between humans and whales? Here we argue that while the latter is difficult to prove, it is a possibility that archaeologists should keep in mind. Indeed, the high value of whale products has prompted a wide diversity of societies across the world to develop approaches for actively accessing them, many of which involving relatively simple technologies [START_REF] Reeves | The origins and character of 'aboriginal subsistence' whaling: a global review[END_REF][START_REF] Reeves | A taxonomy of world whaling[END_REF]. It seems likely that a good number of such societies disappeared without leaving undisputable evidence of their whaling activities, such as credible ethnographic records or realistic rock art. Coastal dynamics and sea level rise may have also eliminated much of the possible archaeological evidence [START_REF] Szabo | Monstrous Fishes and the Mead-Dark Sea: Whaling in the Medieval North Atlantic[END_REF]. Furthermore, in some cases the coastal whales themselves disappeared [START_REF] Rodrigues | Forgotten Mediterranean calving grounds of grey and North Atlantic right whales: evidence from Roman archaeological records[END_REF], putting the possibility of whaling even further beyond the radar of archaeologists.

We thus recommend keeping an open mind and taking a broad perspective when analyzing evidence of ancient whale utilization. This involves placing archaeological records in an ecological context, keeping in mind the diversity of uses as well as whaling approaches known from ethnographic records, and taking advantage of modern molecular analytical techniques that are now revealing the presence of whales at the bottom of a container, in an oil lamp, as the tip of a weapon, or even in a dusty archive. The nature of pre-industrial human-whale interactions is still very poorly understood and its prevalence is certainly underestimated. Molecular and isotopic techniques, be it ancient DNA, ZooMS, radiocarbon dating and stable isotope analysis, residue analyses (or better yet the combination of several of these), are gradually helping us to draw the outline of these interactions, rendering visible previously unnoticed giants.

Figure 1 :

 1 Figure 1 : Whale processing by Basques in Spitzbergen, showing a whale being towed to shore (A), blubber extraction (B and C), blubber chopping (D), oil rendering in ovens (E) and storage in wood casks (G) (Duhamel du Monceau, 1782).

Figure 2 .

 2 Figure 2. Examples of Magdalenian objects made in whale bone from Ermittia -level III (Guipúzcoa, Spain), 1: projectile point, 2: indeterminate object made on rod blank, a-d: close-up of the material. The porosity of whale compact bone tissue is perceptible by the presence of alveoli evenly distributed both inside (a, c) and on the surface (b, d) of the objects (modified from Lefebvre et al., 2021).

Figure

  Figure 3: Whaling scenes: (a) English whalers in the North Atlantic, 19th century (Cheever, 1853) and (b) Makah Indians, Washington State, USA (Elliott, 1886)

  . Oil required containers, but simple options mentioned in historical texts included the intestines of whales (Olioutures; D'Auteroche 1768) or of seals (Changos; Vazquez de Espinosa 1862), dried kelp (Khoisan; Smith & Kinahan 1984), and skins (19 th century Makah from the northwestern United States; Scammon 1874). Basque whalers used wooden casks for the transport and trade of oil (Duhamel du Monceau, 1782).

Acknowledgements

This study is an output of the PaleoCet project, funded by the Agence Nationale de la Recherche (Projet-ANR-18-CE27-0018).

Author contributions

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.