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MAMMAL COMMUNICATIONS Volume 7 ISSN 2056-872X (Online) 17 Page Number Hedgehog Road Casualties in Great Britain MAMMAL COMMUNICATIONS Volume 7 ISSN 2056-872X (online) 17 page number Hedgehog road casualties in Great Britain © Spindrift Images A.C. Kitchener, V.E. Szabo, M. Buckley, Y. van den Hurk, I. Mainland, M. Carruthers, C. MacKay & B.A. Frasier First records of grey whale from Scotland Mammal Communications First records of grey whale, Eschrichtius robustus, from Scotland A.C. Kitchener1*, V.E. Szabo2, M. Buckley3, Y. van den Hurk4, I. Mainland5, M. Carruthers5, C. MacKay5 & B.A. Frasier6 ABSTRACT The grey whale, Eschrichtius robustus, occurred in the Atlantic Ocean until c.300 years ago, having been extirpated probably owing to hunting by humans. Three records of fossil grey whale bones were known previously from Cornwall and Devon in the 18th and 19th centuries. Morphological identification to species of cetacean remains from palaeontological and archaeological sites is problematic, because the bones are often fragmentary or have been modified by humans. The application of ancient DNA analysis and collagen fingerprinting (ZooMS) allows the identification of fragmentary and modified cetacean bone to species and/or generic level, thus offering new opportunities to understand the exploitation of cetaceans by human coastal communities. Analyses of cetacean bone from Orkney and Shetland have identified for the first time the presence of grey whale at archaeological sites in Scotland, extending the geographical range of records in Britain. These findings confirm for the first time that the seas around Scotland were part of the range of the grey whale in the eastern North Atlantic, thereby linking records of this species (from southern Scandinavia, the Northern Isles of and southwest Britain, the southern North Sea and Netherlands, to the Iberian Peninsula and Morocco) over a similar latitudinal range that the species still occupies in the Pacific Ocean. INTRODUCTION The grey whale, Eschrichtius robustus, once inhabited the species was extirpated from the North Atlantic within both the Pacific and Atlantic Oceans. Originally described the last 300 years, probably because of human hunting from fossil remains from Gräso, Sweden by Lilljeborg (Mead & Mitchell 1984; Bryant 1995, Kitchener et al. (1861), the species has since been extirpated from the 2008). In the past decade, there has been a considerable North Atlantic and is today found only in the Pacific increase in the numbers of grey whale fossils discovered Ocean (Jefferson et al. 2015), although two vagrants on both sides of the Atlantic Ocean, with a particular hot have been recorded recently in the Atlantic (Scheinin et spot in the southern North Sea off the coast of the al. 2011; Elwen & Gridley 2013) and a further live animal Netherlands, and it has now been confirmed at several was recorded off Rabat, Morocco in March 2021. archaeological sites in the Mediterranean and elsewhere Radiocarbon dating and historical accounts suggest that in Western Europe (Alter et al. 2015; Rodrigues et al. 1Department of Natural Sciences, National Museums Scotland, Chambers Street, Edinburgh EH1 1JF, UK and Institute of Geography, School of Geosciences, University of Edinburgh, Drummond Street, Edinburgh EH8 9XP, UK 2Department of History, Western Carolina University, Cullowhee, North Carolina, 28723, USA 3School of Natural Sciences, Manchester Institute of Biotechnology, University of Manchester, 131 Princess Street, Manchester, M1 7DN, UK 4University of Groningen, Groningen Institute of Archaeology, Poststraat 6, 9712 ER, Groningen, The Netherlands 5Archaeology Institute, University of the Highlands and Islands, Orkney UHI, Kirkwall KW15 1LX, UK 6Saint Mary's University, Biology Department, 923 Robie Street, Halifax, Nova Scotia, Canada B3H 3C3 * corresponding author [email protected] Key words: Key words: Eschrichtius robustus, grey whale, fossil, archaeology, aDNA, collagen fingerprinting Full citation: Kitchener, A.C., Szabo, V.E., Buckley, M., van den Hurk, Y., Mainland, I., Carruthers, M., MacKay, C. & Frasier, B.A. (2021) First records of grey whale, Eschrichtius robustus, from Scotland. Mammal Communications 7: 17-28, London 18 First records of grey whale from Scotland Mammal Communications 2018; van den Hurk 2020). The presence of a breeding identify a wide array of cetacean species, including the population in the North Atlantic has not been larger mysticete whales, which are the most difficult to demonstrated unequivocally. Evidence for a persistent identify morphologically in archaeological contexts population with annual migrations between feeding and (Buckley et al. 2014; Hufthammer et al. 2018; van den breeding grounds, rather than the presence of occasional Hurk et al. 2021). Collagen preservation is often much vagrants from the Pacific Ocean, is supported by genetic better than aDNA and occurs in conditions in which aDNA differences between Atlantic and Pacific grey whales, and is rapidly degraded. Collagen fingerprinting is also the increasing number of fossil and archaeological relatively inexpensive compared with aDNA analysis and Atlantic records that span around 60,000 years over a does not require such stringent laboratory conditions. wide latitudinal range (Alter et al. 2015; Rodrigues et al. However, collagen fingerprinting may limit the level of 2018). Although grey whale fossil or archaeological taxonomic identification to family or genus, especially for remains have been reported from a few sites in southwest closely related species. However, this is not the case for England (Bryant 1995, Kitchener et al. 2008), no grey mysticete whales, except for the family Balaenidae, which whale fossil or archaeological remains have yet been includes the right whales, Eubalaena spp. and bowhead reported from Scotland or elsewhere in the British Isles. whale, Balaena mysticetus (Buckley et al. 2014, Whale skeletal remains occur frequently at archaeological Hufthammer et al. 2018). sites, but they can be difficult to identify owing to As part of the interdisciplinary NSF-funded project, fragmentation or modification. This often results in a lack Assessing the Distribution and Variability of Marine of identification or mis-identification (Buckley et al. 2014; Mammals through Archaeology, Ancient DNA, and Hufthammer et al. 2018). Analysis of ancient DNA History in the North Atlantic (henceforth Norse North (aDNA) is increasingly being used to identify Atlantic Marine Mammals), archaeological cetacean archaeological bone fragments to species and can offer bones from several sites in Orkney and Shetland were additional information, such as sex and aspects of sampled for aDNA analyses and collagen fingerprinting, population genetics. These data inform the interpretation in order to identify them to species level and provide a of skeletal remains within an archaeological context, but more complete and accurate assessment of the utilisation also provide information on aspects of species of cetaceans in the Norse period. Here we report on five demographics over time. However, aDNA may be poorly bones of grey whale from three sites in Orkney and one or not preserved and can be relatively expensive to site in Shetland that were identified using a combination extract and analyse, requiring uncontaminated laboratory of aDNA analysis and collagen fingerprinting. These are conditions for successful extraction. In recent years the first records from Scotland and the first confirmed collagen fingerprinting (ZooMS) has been developed to records from archaeological sites in Britain. MATERIALS AND METHODS As part of the continuing Norse North Atlantic Marine from compact bone (Qiagen, Mississauga, ON). In all Mammals project, 109 cetacean bones from steps of DNA analysis (e.g., DNA extraction, archaeological sites were selected for aDNA analysis and amplification, sequencing), negative (or ‘null’) controls collagen fingerprinting from the collections of National were included as a means to detect potential DNA Museums Scotland and Orkney Museums, and material contamination. from continuing excavations of the UHI Archaeology We examined for the presence of analysable DNA by Institute in Orkney. This sample set is comprised primarily amplifying and sequencing a ~445 basepair (bp) fragment of bone fragments that are not identifiable to species of the cytochrome b gene (using primers CBCet4F (5’ using traditional morphological methods. To date, these ACA TGG ACT TCA ACC ATG AC 3’) and CBCet5R (5’ bones comprise specimens that have been sampled and CTC AGA ATG ATA TTT GTC CTC AGG 3’). PCR analysed from 11 sites in Orkney and Shetland, Scotland. product was evaluated for quantity and quality using An additional bone (Specimen 5) was later added to the agarose electrophoresis, then prepared for sequencing, current study. Although all 110 specimens were submitted cycle sequenced, and size separated and visualised on for aDNA (except for Specimen 5) and collagen an ABI 3500xl Genetic Analyzer (Applied Biosystems) fingerprinting, only results pertaining to identified grey after McLeod et al. (2014). All samples were amplified in whale identifications are discussed further. duplicate and then each duplicate was sequenced in both directions (e.g., with both primers of the primer pair used Ancient DNA analysis for amplification). Sequences were then examined All 109 specimens (excluding Specimen 5) underwent visually using 4Peaks v1.8 (Nucleobytes.com) and edited bone tissue collection/subsampling, aDNA handling and and aligned in MEGAX v10.2.4 (Kumar et al. 2018). DNA extraction as per Rastogi et al.
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