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Nature of the beast? Resolving drivers of prey choice, competition and resilience in wolves

Lead Research Organisation: University of Bristol
Department Name: Geographical Sciences

Abstract

Wolves were well-established members of the Pleistocene (Ice Age) carnivore community in Europe but today, many surviving populations of these charismatic animals are endangered because of human persecution and environmental change. As keystone predators, wolves play a vital role in maintaining biodiversity, particularly in keeping mammalian herbivore and medium-sized carnivore numbers in check, thereby limiting over-browsing on vegetation and over-predation on small vertebrates respectively. In this regard, they are the most influential large predator in the northern Palaearctic. The ripples from their activity can therefore be felt in diverse positive ways throughout the ecosystem but serious concerns exist as to the viability of European wolf populations under different scenarios of environmental and climate change. A key goal is therefore to understand how wolves have adapted to changing circumstances so that current and future conservation policy can be appropriately tailored.

One of the best ways to approach this issue is through the study of diet, since this is closely linked to climate and environment (determining which prey species are available) and to competition for resources from other carnivores. Our previous research into the British fossil wolf record revealed marked changes in the size and shape of the jaws and teeth over the last half a million years, which together with evidence from tooth breakage and wear, indicate that wolves modified their diet (consuming more/less meat versus non-meat foods) in response to changing environmental parameters.

Such morphological change cannot readily be measured in the short time scales (years to decades) of modern ecological studies but the rich Pleistocene fossil record offers a chronologically well-resolved series of wolf specimens spanning tens to hundreds of thousands of years, allowing patterns of change to be fully tested against diverse variables such as changing climates, environments, carnivore competition and prey availability. We successfully tested these palaeodietary assumptions in two NERC-funded studies on fossil wolf remains from three different climatic episodes (glacial and interglacial), using direct measurements of bone chemistry through carbon and nitrogen stable isotope analysis in order to verify changing prey choice through time. We now propose to expand this study in what will be the most comprehensive and state-of-the-art examination of diet in modern and recent fossil (<250,000 years) European wolves by using a series of independent proxies operating on different temporal scales.

Working with conservation biologists, a key aim is the integration of morphological and dietary evidence from modern wolves from Sweden, Poland and Croatia, using a combination of GPS data on radio-collared wolves to identify kill sites, analysis of the contents of wolf scats, and stable isotope evidence from recently culled or dead specimens. As well as revealing seasonal and geographical variation in wolf diet, our research will allow for the first time: (1) direct comparison of modern, Holocene and Pleistocene wolf diet; (2) investigation of the degree to which direct (stable isotope, dental microwear) and indirect (morphometric) measurements of diet are in step with real-time dietary evidence from scat analyses and kill sites; (3) evaluation of the influence of diet on the morphology of modern wolves and (4) the opportunity to "ground truth" the evidence generated by current palaeodietary approaches, by assessing whether it replicates that obtained from analyses of modern wolf diet. By understanding the ecological trajectory of past and current wolf populations, we will generate a new, evidence-based view of the impacts on European large carnivores of climate, prey choice and environment.

As well as academic beneficiaries, we will reach new audiences through public outreach at the Wildwood Trust and art commissions for gallery and online display.

Publications

10 25 50

Related Projects

Project Reference Relationship Related To Start End Award Value
NE/W006103/1 31/08/2022 30/07/2024 £578,582
NE/W006103/2 Transfer NE/W006103/1 31/07/2024 28/05/2026 £212,514
 
Title The Nature of the Beast 
Description This artistic work is a collaboration between graphic artist and animator, Sean Harris, and the Nature of the Beast research team. The work takes the form of a 33 minute long film, which showcases aspects of the research methods (stable isotope geochemistry, dental microwear texture analysis) and commentary by the PI, co-I and postdoctoral researcher, and combines these with Sean's artworks - animations of grey wolves alongside other elements of the Pleistocene mammal fauna. The animation explores the ecological resilience of the wolf in the face of climatic and environmental change, as well as changing prey and competitors, before reflecting on the ecological consequences of the extinction of this species. 
Type Of Art Artwork 
Year Produced 2026 
Impact The piece has only so far been shown to an in person audience of c.130 people before being launched online, but it has inspired discussion on the ecological role of the wolf and its rewilding potential from viewers. It is being presented in March 2026 as part of a talk by Sean Harris at the National Library of Wales in Aberystwyth on 'The Nature of the Beast and the Mind in the Cave: Resurrecting the British Wolf through Art and Science.' 
URL https://vimeo.com/1169548951
 
Title Dental Microwear Texture Analysis (DMTA) of tooth surfaces of fossil wolves from Britain 
Description One Excel workbook reporting dental microwear texture analysis (DMTA) raw data. These are values from scale-sensitive fractal analysis (SSFA), and areal surface texture analysis (based on a suite of parameters, most of which are defined by ISO-25718-2) for dental microwear texture analysis of tooth surfaces of wolves from Britain (MIS 2, MIS 3, MIS 5a). This study examines the dietary adaptability of European grey wolves using dental microwear texture analysis (DMTA). These datasets offer new insights into the palaeoclimatic and palaeoenvironmental conditions under which grey wolves endured climate change in the past, free of anthropogenic influence. Tooth surface casts were produced using high-resolution, which is optimised for imaging purposes. Epoxy tooth replicas were scanned with a Sensofar S neox. MountainsMap Imaging Topography was utilised to quantify dental microwear texture parameters from Sensofar scans. 
Type Of Material Database/Collection of data 
Year Produced 2026 
Provided To Others? Yes  
Impact N/A 
URL https://www2.bgs.ac.uk/nationalgeosciencedatacentre/citeddata/catalogue/3d4a8583-ebc7-4cce-ad06-c4f8...
 
Title Dental Microwear Texture Analysis (DMTA) of tooth surfaces of fossil wolves from Britain and modern wolves from Poland 
Description Dental microwear texture analysis (DMTA) raw data. These are values from scale-sensitive fractal analysis (SSFA), and areal surface texture analysis (based on a suite of parameters, most of which are defined by ISO-25718-2) for dental microwear texture analysis of tooth surfaces of wolves from Britain (MIS 5e & MIS 7 a-c) and modern Poland wolves. This study examines the dietary adaptability of European grey wolves using dental microwear texture analysis (DMTA). We compare modern Polish wolf populations from the present interglacial (Holocene), with fossil specimens from two contrasting Pleistocene interglacials: the warmer Last Interglacial (Marine Oxygen Isotope Stage [MIS] 5e) and the cooler Penultimate Interglacial (MIS 7a-c). These datasets offer new insights into the palaeoclimatic and palaeoenvironmental conditions under which grey wolves endured climate change in the past, free of anthropogenic influence. And allows the identification of hitherto-hidden ecological stress today, thereby highlighting potential vulnerabilities in modern populations under current and future climate change. 
Type Of Material Database/Collection of data 
Year Produced 2025 
Provided To Others? Yes  
Impact N/A 
URL https://www2.bgs.ac.uk/nationalgeosciencedatacentre/citeddata/catalogue/bfb67ae1-cd30-4e4f-8cb5-aa6c...
 
Title Dental Microwear Texture Analysis (DMTA) of tooth surfaces of fossil wolves from Britain and modern wolves from Poland version 2 
Description Two Excel workbooks reporting dental microwear texture analysis (DMTA) raw data. These are values from scale-sensitive fractal analysis (SSFA), and areal surface texture analysis (based on a suite of parameters, most of which are defined by ISO-25718-2) for dental microwear texture analysis of tooth surfaces of wolves from Britain (MIS Se & MIS 7 a-c) and modern Poland wolves. This study examines the dietary adaptability of European grey wolves using dental microwear texture analysis (DMTA). These datasets offer new insights into the palaeoclimatic and palaeoenvironmental conditions under which grey wolves endured climate change in the past, free of anthropogenic influence. They further allow the identification of hitherto-hidden ecological stress today, thereby highlighting potential vulnerabilities in modern populations under current and future climate change. Tooth surface casts were produced using high-resolution, which is optimised for imaging purposes. Epoxy tooth replicas were scanned with a Sensofar S neox. MountainsMap Imaging Topography was utilised to quantify dental microwear texture parameters from Sensofar scans. 
Type Of Material Database/Collection of data 
Year Produced 2026 
Provided To Others? Yes  
Impact N/A 
URL https://www2.bgs.ac.uk/nationalgeosciencedatacentre/citeddata/catalogue/0aa0e3e8-d668-4dd4-9089-7bd0...
 
Title Measurements of wolf teeth and mandibles 
Description Museums visited (n=13): British Geological Survey, Bristol Museum and art gallery, The Box museum-Plymouth, Museum of Manchester, Natural History Museum, National Museum Wales, Museum of Zoology Cambridge & Sedgwick Resource Centre, Torquay Museum, Creswell Crags Museum, Bucks Museum Resource Centre, Wollaton Hall, Somerset Heritage Centre, Oxford Natural History Museum. We also visited modern wolf collections at (Poland) University of Warsaw in Urwitalt and (Croatia) Faculty of Veterinary Medicine University of Zagreb. A total of 171 first mandibular molars (m1) and 143 second mandibular molars (m2) have been identified as appropriate for this research. These (n=314) teeth represent 221 individual wolves. Wolves were identified from the following marine isotope stages: Holocene, n=25; MIS 2, n=24; MIS 3, n=52; MIS 5, n=62 (substages are 5a, n=25; 5c, n=10; 5e, n=27); MIS 6, n=10; MIS 7, n=26; and 22 from sites that are still being investigated to determine their age. Additionally, samples from contemporary wolves were collected to serve as baseline data. This included first and second molars from modern wolf jaw collections in Poland (n=41 individuals, n=82 teeth) and Croatia (n=111 individuals, n=222 teeth). 
Type Of Material Database/Collection of data 
Year Produced 2024 
Provided To Others? No  
Impact N/A 
 
Title Nature of the Beast: Dental microwear database - digital scans 
Description Each cast has been scanned multiple times with a Sensofar S-Neox confocal microscope. The total area of each scan is approximately 320 x 270 µm taken from the facet on the antero-lingual surface of the hypoconid on the second mandibular molar and the talonid basin of the first mandibular molar. There have been approximately 1860 scans taken. Scans that are considered to most accurately represent antemortem dietary behaviour, specifically those without postmortem damage to the tooth surface, are subsequently measured and data are captured to represent that particular tooth (wolf). Surface files generated by confocal microscopy are processed with MountainsMap® (Digital Surf), designed for analysing surface texture, form, and topography at the microscopic scale. This software generates both Scale-Sensitive Fractal Analysis (SSFA) values and ISO standards that define parameters for characterizing surface roughness. Finally, JMP statistical software is used to evaluate significance of these values and to explain variability. 
Type Of Material Database/Collection of data 
Year Produced 2024 
Provided To Others? No  
Impact N/A 
 
Title Nature of the Beast: Dental microwear database - resin casts 
Description For the purpose of conducting dental microwear texture analysis on wolves, thirteen museums across the United Kingdom were visited to evaluate their collections for the presence of first and second mandibular molars from grey wolf specimens. A total of 171 first mandibular molars (m1) and 143 second mandibular molars (m2) have been identified as appropriate for this research. These (n=314) teeth represent 221 individual wolves. Wolves were identified from the following marine isotope stages: Holocene, n=25; MIS 2, n=24; MIS 3, n=52; MIS 5, n=62 (substages are 5a, n=25; 5c, n=10; 5e, n=27); MIS 6, n=10; MIS 7, n=26; and 22 from sites that are still being investigated to determine their age. Additionally, samples from contemporary wolves were collected to serve as baseline data. This included first and second molars from modern wolf jaw collections in Poland (n=41 individuals, n=82 teeth) and Croatia (n=111 individuals, n=222 teeth). Dental moulds of the occlusal surfaces of mandibular molars (m1&m2) were collected with Regular body polyvinylsiloxane dental impression material (President's Jet, Coltene-Whaledent Corp.). Tooth replicas were then produced with high resolution epoxy, resulting in 618 dental casts (British fossil wolves (n=314); Poland modern wolves (n=82); Croatia modern wolves (n=222)). 
Type Of Material Database/Collection of data 
Year Produced 2024 
Provided To Others? No  
Impact The British Geological Survey published a blog on the use of BGS specimens in the project. The Museum of Somerset made an Instagram post about the sampling of their specimens in the project. 
 
Title Stable isotope sampling and processing 
Description Samples for destructive analysis have been obtained from the following museums: Discover Bucks Museum Resource Centre British Geological Survey Bristol museum and art gallery The Box museum-Plymouth Museum of Manchester National Museum Wales Sedgwick Resource Centre Cambridge Torquay Museum Creswell Crags museum National History Museum-Mammals Department National History Museum-Fossil mammals Department Oxford University Museum of Natural History as of 09/3/26, 450 specimens have been sampled for %N screening and this has resulted in 267 specimens taken forward for collagen extraction. 
Type Of Material Database/Collection of data 
Year Produced 2025 
Provided To Others? No  
Impact The British Geological Survey published a blog to highlight the use of the BGS museum specimens in the research The British Geological Survey published a blog to highlight the use of the Creswell Crags museum specimens in the research 
 
Description EU COST Action "PalaeOpen" 
Organisation Utrecht University
Country Netherlands 
Sector Academic/University 
PI Contribution PalaeOpen has the key objective of advancing understanding of ecosystem responses to climate and land-use changes over decades to millennia and using that information to foster collaboration between palaeoecologists, macroecologists, and conservation practitioners, addressing knowledge gaps and supporting nature conservation. Project data was presented by Danielle Schreve as a keynote lecture at the launch of this EU COST Action programme and she is now leading the Vertebrates sub-group of the COST Action and developing a community of Quaternary vertebrate researchers. The Nature of the Beast research team will be contributing project data (specifically stratified records of wolf and other large carnivores and herbivores, as well as published stable isotope measurements) to the Neotoma paleoecology database (https://www.neotomadb.org/), into which vertebrate records are being deposited. This is an important new development, since the use of such databases by the Quaternary vertebrate community has historically been very poor.
Collaborator Contribution The PalaeOpen management group are providing training for "data stewards" for the Neotoma database. The wider project provides a European (and wider) community of vertebrate workers to share data. Financial support is available to organise and attend workshops.
Impact Database under development
Start Year 2025