Quaternary Science Reviews 98 (2014) 144e165 Contents lists available at ScienceDirect Quaternary Science Reviews journal homepage: www.elsevier.com/locate/quascirev Origin of British and Irish mammals: disparate post-glacial colonisation and species introductions W. Ian Montgomery a, *, Jim Provan a, A. Marshal McCabe b, Derek W. Yalden c a School of Biological Sciences, Queen's University Belfast, MBC, 97 Lisburn Rd., Belfast BT9 7BL, Northern Ireland, UK School of Environmental Sciences, University of Ulster, Coleraine, BT52 1SA, Northern Ireland, UK c Faculty of Life Sciences, University of Manchester, Oxford Rd., Manchester, M13 9PL, UK b a r t i c l e i n f o a b s t r a c t Article history: Received 7 October 2013 Received in revised form 24 March 2014 Accepted 27 May 2014 Available online 27 June 2014 Global climate changes during the Quaternary reveal much about broader evolutionary effects of environmental change. Detailed regional studies reveal how evolutionary lineages and novel communities and ecosystems, emerge through glacial bottlenecks or from refugia. There have been significant advances in benthic imaging and dating, particularly with respect to the movements of the British (Scottish) and Irish ice sheets and associated changes in sea level during and after the Last Glacial Maximum (LGM). Ireland has been isolated as an island for approximately twice as long as Britain with no evidence of any substantial, enduring land bridge between these islands after ca 15 kya. Recent biogeographical studies show that Britain's mammal community is akin to those of southern parts of Scandinavia, The Netherlands and Belgium, but the much lower mammal species richness of Ireland is unique and needs explanation. Here, we consider physiographic, archaeological, phylogeographical i.e. molecular genetic, and biological evidence comprising ecological, behavioural and morphological data, to review how mammal species recolonized western Europe after the LGM with emphasis on Britain and, in particular, Ireland. We focus on why these close neighbours had such different mammal fauna in the early Holocene, the stability of ecosystems after LGM subject to climate change and later species introductions. There is general concordance of archaeological and molecular genetic evidence where data allow some insight into history after the LGM. Phylogeography reveals the process of recolonization, e.g. with respect to source of colonizers and anthropogenic influence, whilst archaeological data reveal timing more precisely through carbon dating and stratigraphy. More representative samples and improved calibration of the ‘molecular clock’ will lead to further insights with regards to the influence of successive glaciations. Species showing greatest morphological, behavioural and ecological divergence in Ireland in comparison to Britain and continental Europe, were also those which arrived in Ireland very early in the Holocene either with or without the assistance of people. Cold tolerant mammal species recolonized quickly after LGM but disappeared, potentially as a result of a short period of rapid warming. Other early arrivals were less cold tolerant and succumbed to the colder conditions during the Younger Dryas or shortly after the start of the Holocene (11.5 kya), or the area of suitable habitat was insufficient to sustain a viable population especially in larger species. Late Pleistocene mammals in Ireland were restricted to those able to colonize up to ca 15 kya, probably originating from adjacent areas of unglaciated Britain and land now below sea level, to the south and west (of Ireland). These few, early colonizers retain genetic diversity which dates from before the LGM. Late Pleistocene Ireland, therefore, had a much depleted complement of mammal species in comparison to Britain. Mammal species, colonising predominantly from southeast and east Europe occupied west Europe only as far as Britain between ca 15 and 8 kya, were excluded from Ireland by the Irish and Celtic Seas. Smaller species in particular failed to colonise Ireland. Britain being isolated as an island from ca. 8 kya has similar species richness and composition to adjacent lowland areas of northwest continental Europe and its mammals almost all show strongest genetic affinity to populations in neighbouring continental Europe with a few retaining genotypes associated with earlier, western lineages. The role of people in the deliberate introduction of mammal species and distinct genotypes is much more significant with regards to Ireland than Britain reflecting the larger species richness of the latter Keywords: Quaternary Holocene Postglacial recolonisation Mammal Archaeology Phylogeography Species introductions British Isles * Corresponding author. E-mail address: i.montgomery@qub.ac.uk (W.I. Montgomery). http://dx.doi.org/10.1016/j.quascirev.2014.05.026 0277-3791/© 2014 Elsevier Ltd. All rights reserved. W.I. Montgomery et al. / Quaternary Science Reviews 98 (2014) 144e165 145 and its more enduring land link with continental Europe. The prime motivation of early people in moving mammals was likely to be resource driven but also potentially cultural; as elsewhere, people exploring uninhabited places introduced species for food and the materials they required to survive. It is possible that the process of introduction of mammals to Ireland commenced during the Mesolithic and accelerated with Neolithic people. Irish populations of these long established, introduced species show some unique genetic variation whilst retaining traces of their origins principally from Britain but in some cases, Scandinavia and Iberia. It is of particular interest that they may retain genetic forms now absent from their source populations. Further species introductions, during the Bronze and late Iron Ages, and Viking and Norman invasions, follow the same pattern but lack the time for genetic divergence from their source populations. Accidental introductions of commensal species show considerable genetic diversity based on numerous translocations along the eastern Atlantic coastline. More recent accidental and deliberate introductions are characterised by a lack of genetic diversity other than that explicable by more than one introduction. The substantial advances in understanding the postglacial origins and genetic diversity of British and Irish mammals, the role of early people in species translocations, and determination of species that are more recently introduced, should inform policy decisions with regards to species and genetic conservation. Conservation should prioritise early, naturally recolonizing species and those brought in by early people reflecting their long association with these islands. These early arrivals in Britain and Ireland and associated islands show genetic diversity that may be of value in mitigating anthropogenic climate change across Europe. In contrast, more recent introductions are likely to disturb ecosystems greatly, lead to loss of diversity and should be controlled. This challenge is more severe in Ireland where the number and proportion of invasive species from the 19th century to the present has been greater than in Britain. © 2014 Elsevier Ltd. All rights reserved. 1. Introduction The dramatic glacial history of the Quaternary is associated with the ebb and flow of northern temperate species; both northern and southern refugia supported isolated populations during interglacials and glacials respectively (Stewart et al., 2010). Refugia are temporary havens which expand bringing together formerly separate lineages that may continue on their route to speciation or alternatively mix, introgress and dissipate re-establishing a single but perhaps more genetically and physiologically diverse population (Provan and Bennett, 2008). The western and eastern coastlines of the land masses of the northern hemisphere are particularly important in this process as not only are these associated with refugia, they are also remote and may have supported isolated populations on islands and peninsula even during extreme environmental conditions (Me'dail and Que'zel, 1997; Hewitt, 1999; Deffontaine et al., 2005). Indeed, regional topography may play a major role in effecting suitable conditions where, more generally none exist, allowing populations to persist at higher latitudes than thought possible (Dobrowski, 2011). Regional variation in landscape, habitat and species interactions are of biogeographical significance in many taxa (Vetaas and FerrerCastan, 2008; Filipe et al., 2010; Fløjgaard et al., 2011; Kissling and Cagan, 2012) and are indicative of how particular taxa might cope with anthropogenic climatic change (Beirer and Brost, 2010; Dormann et al., 2010; Bradbury et al., 2011). The western Atlantic fringe has an excellent environmental record for the Quaternary with detailed vegetation history based on pollen remains in peat and lake sediments (Mitchell and Ryan, 1997; Pilcher and Hall, 2001), dendrochronology (Pilcher et al., 1995; Moir, 2012) and archaeozoology (Woodman et al., 1997; Yalden, 1999). Improved carbon dating has provided further insight into environmental change (Fairbanks et al., 2005; Mellars, 2006) and the introduction of enhanced methods for surveying the continental shelf and coastal waters (see below) has revealed the extent and timing of ice sheet dynamics and evidence of hitherto unsuspected presence of humans and animal species in land exposed during periods of lower sea level during the early Holocene (Gaffney et al., 2009). Further, the application of molecular genetics to biogeography, broadly referred to as phylogeography, is a powerful means of establishing the processes that were responsible for postglacial colonization as well as population genetic structure in ubiquitous species and relatedness amongst fragmented and seemingly isolated populations (Avise, 2000). Here, we review the late Pleistocene and Holocene history of the mammals of Britain and Ireland using archaeological, phylogeographic and ecological evidence in the context of the improved understanding of the physical changes that took place during the latter part of the Quaternary. We re-evaluate the impact on mammal fauna of the earlier isolation of Ireland in comparison to Britain (Mitchell and Ryan, 1997) as well as the role of people in transporting animal species during prehistory (Grayson, 2001), and emphasise the importance of a Europe-wide perspective in identifying potential refugia and reconstructing re-colonisation routes to continental margins. It is concluded that the recent history of mammal introductions into Britain and, in particular, Ireland due to globalisation of trade and lax border control, further marks the beginning of the Anthropocene (Crutzen and Stoermer, 2000) and that the impact of these introductions is yet to be fully appreciated. Whilst the principal aim here is to elucidate the history of the mammalian fauna of Britain and Ireland and to reflect on their future, it is important from legal, political and management perspectives to establish with a high degree of certainty endemic, native or indigenous species and evolutionary significant units (Ryder, 1986; Moritz, 1994; Lindenmayer and Burgman, 2005). Dates are given in kya, thousands of years ago (before 2000AD). We follow Searle's (2008) approach in discriminating early and late natives which arrive naturally, through their own means, in contrast with early and late introductions where people have accidentally or deliberately translocated non-native species. We refer to indigenous species combining the early native (present at start of Holocene, 11.5 kya) and early introduced species (associated with Mesolithic and Neolithic people) which have become part of the ecology of habitats throughout discrete biogeographical units and may have had a significant economic (resource-based) or cultural role. Later species introductions are associated with the Bronze Age, Iron Age, the Romans (in Britain), Vikings, Normans and during industrialisation and the emergence of global trade during the 19the21st centuries AD. 146 W.I. Montgomery et al. / Quaternary Science Reviews 98 (2014) 144e165 2. Current mammal species richness and affinities in northwestern Europe There is a decline in mammal species richness across Europe from the southeast to the northwest (Fløjgaard et al., 2011) which is evident along the continental fringe as well as in Britain and Ireland. In terms of species composition, Ireland and Britain represent distinctive assemblages. The species composition of the mammal community of Britain has affinities with those of the Benelux countries, Denmark and southern Sweden; the less species rich mammal community of Ireland, however, groups with Britain but also with similar impoverished mammal faunas of the western and northern coastline of Norway (Heikinheimo et al., 2007; AarisSorensen, 2009; Svenning et al., 2011). It has been acknowledged historically, that Britain and particularly Ireland (see, for example, Giraldus Cambrensis, Topographica Hibernica, 1187), have fewer mammal species than adjacent areas of continental Europe (Yalden, 1982, 1999; Fairley, 1984). Comparisons are often made with France (Fig.1) but this seems inappropriate given its much greater latitudinal and altitudinal range in comparison with Britain and Ireland (see Heikinheimo et al., 2012). Here, we make comparisons with Belgium and The Netherlands as recolonisation of Britain by mammals is likely to have occurred across what is now the southern North Sea (Gaffney et al., 2009). Britain lacks only a few mammal species across all families in comparison to Belgium and The Netherlands. Ireland, in contrast, is lacking in mammal species across all families with the possible exception of the Artiodactyla. Ireland is disproportionately low in species richness in families comprised of species with smaller body size and ecologically dependent on woodland and forest, namely, many rodents, insectivores and bats (Fig. 1). As rates of dispersal in terrestrial mammals are related to body size (Sutherland et al., 2000) it is possible this pattern conforms to the Littletonian ‘steeplechase’ model proposed by Mitchell (Mitchell and Ryan, 1997). This presumes an earlier isolation of the island of Ireland compared to Britain i.e. many small mammals could not recolonise Europe after the Last Glacial Maximum (LGM) quickly enough to reach Ireland whereas they either remained in Britain or were able to disperse into the latter before it too was isolated. However, this relatively simple model of postglacial recolonisation must be consistent with recent physical, archaeological and genetic data to remain viable. Further, it is necessary to consider the interplay of ecosystem and trophic factors involving climate, topography, plants and consumers (Heikinheimo et al., 2012). 3. Environmental change LGM to Holocene Methodological advances have changed the understanding of the extent of ice sheets around LGM and the process of deglaciation (Bowen et al., 2002; Clark et al., 2009a,b; O'Cofaigh et al., 2010; Clark et al., 2012a,b). The LGM is defined as the most recent interval when ice sheets reached their maximum integrated volume (Mix et al., 2001). Growth of ice sheets from ca 33 kya resulted in LGM from 26 to 19 kya (Clark et al., 2009a,b). Increases in northern summer insolation marked the start of major deglaciations and abrupt rises in sea levels. However, regional variability in deglaciation depended on regional/global climatic change and the sensitivity of individual ice sheets (Cheverill and Thomas, 2010). There was also a thick ice sheet over western Ireland for up to 20,000 years prior to LGM (McCabe and Clarke, 1998). Therefore, careful field assessment of individual ice sheets is required to elucidate icesea-land interactions over the 15,000 years following the LGM. The British and Irish ice sheet (BIIS) extremities are poorly documented but moraine ridges and subglacial bedforms suggest that most of the inner shelf areas were ice covered (Dunlop et al., 'Native' Species by Order and Country 30 25 Fr 20 Bel 15 10 5 Net Br Ir 0 Rodents Lagomorphs Artiodactyls Carnivores Insectivores Bats Fig. 1. Numbers of presumed native or early introduced species in named families of mammals in France, Belgium, Netherlands, Britain and Ireland. Based on IUCN data (http://data.iucn.org/dbtw-wpd/edocs/RL-4-013.pdf). 2010). Ice sheet extents and changes in relative sea levels (RSLs) are related to wider climatic conditions, meltwater pulses and the strength of the thermohaline circulation in the north Atlantic € se et al., 2012). Oxygen isotope varia(McCabe and Clark, 1998; Bo tion identified from Greenland ice cores (NGRIP) is a record of primary production and, hence, is a proxy for average global temperature and associated ecological changes (Birks and Ammann, 2000). Two time periods follow the LGM. The first lasts for 5000e7000 years and is characterised by very rapid changes in ice marginal positions, massive reductions in ice mass, abrupt changes in relative sea level and sediment regimes, and newly emergent surface morphologies. The second is more relaxed with little ice mass remaining, reduced, slower crustal rebound, marked changes in coastal configuration and a variable sea level rise from around 130 m to the present. Most ice sheets attained their maximum extents by 26.5 kya (Clark et al., 2009a,b). Models do not capture the complexity of sea level changes immediately after ice sheet retreat (Shennan et al., 2005, 2006; Brooks et al., 2008) but dated deposits and landforms record high RSL around the decaying ice sheet in northern and western Britain McCabe et al., 2007a,b; McCabe and Williams, 2012). From ca 15 kya, temperature increased but then temperature decreased until ca 12.4 kya (Joris and Weniger, 2000). This colder period, the Younger Dryas, is associated with expansion of the Scottish ice sheet, and continued to ca 11.8 kya with temperature rising quickly to historical levels. The Holocene commenced ca 11.5 kya and is characterised by a very stable climate until the present. The warming period running from ca 15 kya and the subsequent colder period of the Younger Dryas, perhaps just 2500 years in total, potentially played a significant role in determining the future of cold-adapted and then warm-adapted mammals in Britain and Ireland (Sommer and Zachos, 2009). 3.1. Extent and timing of LGM Clark et al. (2009a,b) used over 4000 dates to constrain LGM to between 26.5 and 19 kya with ice sheet sectors attaining maxima at different times. LGM for Ireland was between 28 and 23 kya with similar estimates for BIIS (Clark et al., 2012b) where the main centres of ice dispersion were over the Scottish Highlands and the central and western lowlands of Ireland. The lower mountains on the coastal fringes of Ireland supported valley and corrie glaciers with the exception of an ice cap in the southwest. Satellite imagery combined with dating regional ice sheet events indicate that BIIS was dynamic with millennial time scale changes or less, in centres of ice dispersion and ice mass (McCabe et al., 1998). Ice moved from the terrestrial centres of ice dispersion onto at least the inner W.I. Montgomery et al. / Quaternary Science Reviews 98 (2014) 144e165 continental shelf though off the western coast of Ireland ice margins waxed and waned on centennial timescales (McCabe et al., 2007). Inner and outer shelf moraines remain undated but their ages may be inferred from cross-cutting features in subjacent marine bays (Dunlop et al., 2010). In Britain, the ice limit can be traced from south Wales northeastwards towards Yorkshire (Bowen et al., 2002; Clarke et al., 2012a,b). The Irish Sea ice stream was a main conduit draining the BIIS into the central basin from adjacent areas of Scotland, southwards towards a poorly defined limit in the northern Celtic Sea (Eyles and McCabe, 1989). Ice from Scotland may have extended as far as the outer shelf edge west of the Outer Hebrides (Everest et al., 2013). The BIIS extended northeastwards across the northern North Sea to join the Fenno-Scandinavian ice sheet (Cheverill and Thomas, 2010; Clark et al., 2012a,b). Bradwell et al. (2008) identified the decoupling of Scottish and Fennoscandian ice masses in the northern North Sea Basin from morainic patterns accompanied by catastrophic changes linked to rises in sea levels following LGM. Similar ice advance and sediment inputs followed by retreat occurred around LGM in the low lying coastal areas of eastern England (Wingfield, 1990; Eyles et al., 1994). Grounded sea ice beyond the current southern Irish coastline indicates that the whole island was glaciated during the LGM (O’Cofaigh and Evans, 2001). The excursion of the Irish Sea ice stream southwards of Ireland was shortlived, however, reaching a peak around 24 kya with major deglaciation by 23 kya (O'Cofaigh et al., 2010). Unglaciated, coastal shelf may have been exposed to the south and southwest of Ireland shortly after the latter date. 3.1.1. Retreat of BIIS in the west e formation of the Irish Sea The retreat from LGM on the continental shelf around Ireland is well known being constrained by over 100 14C and cosmogenic dates (McCabe, 2008; Clark et al., 2012b). Early deglaciation began ca 23 kya and continued to 18 kya resulting in the loss of about two thirds of the BIIS (McCabe and Clark, 1998; Clark et al., 2012b; O'Cofaigh et al., 2012). Rates of retreat varied considerably; ice streams lying over the Irish Sea retreated as rapidly as > 140 m per year, whilst inter-stream ice sheets retreated at ca 10 m per year (Clark et al., 2012a,b). High RSL accompanied contraction of ice sheet margins in western Ireland (Thomas and Cheverrill, 2006; Clark et al., 2012b) and in the Irish Sea Basin (Eyles and McCabe, 1989). High RSL in the Irish Sea Basin allowed calving of icebergs leading to deposition of morainal banks and marine muds (e.g. McCabe and O'Cofaigh, 1995; McCabe et al., 2005). This deglacial phase was complete in, at most, a few thousand years (Eyles and McCabe, 1989). Ice loss from the basin allowed the global meltwater pulse which occurred at ca 19 kya, to deposit thick marine muds along the coast of northeast Ireland providing critical field evidence for marine flooding during early deglaciation (Clark et al., 2004). During the Cooley Point Interstadial, ice remained near to basinal margins, but elsewhere, ice lobes advanced marginally (Thomas and Cheverrill, 2007; Van Landeghem et al., 2009). Hence, it is likely that from ca 19 kya, the middle and southern Irish Sea Basin was free from extensive ice masses. Early deglaciation left all of the Irish midlands and most of northern England free of ice. Ice masses in Scotland, however, remained with most margins near to coastal locations and ice readvances probably overstepped earlier ice limits especially in the west (Bradwell et al., 2008). In uplands, such as the Lake District, deglaciation is described as ‘active ice retreat’ as opposed to one of ‘stagnation and wasting’ (Pinson et al., 2013). In Wales, ice which was contiguous with the Irish Sea ice stream contracted from the South Wales, end moraine (Bowen, 1973, 1991). Overall, highland based ice was more robust to climate forcing than the ice located on lowlands and, hence, residual ice masses remained longest in 147 Scotland. Early deglaciation resulted in the loss of about two thirds of the BIIS. Thus, marine seaways developed around almost the entire Irish coast for the first time since LGM (McCabe et al., 2007a,b). Seabed data (Kershaw, 1986) to the west of the Isle of Man, also suggest that the central and southern Irish Sea Basin was a marine highway by ca 21 kya. This is confirmed by the raised, marine muds associated with the meltwater pulse ca 19 kya (Clark et al., 2004). Sea level rises were as high as 50 mm per year (Cronin, 2012) but global eustatic sea levels were low at this time, so that the Irish Sea Basin was deeply, isostatically depressed. The southern basin must still have been isostatically depressed despite meltwater inputs from adjacent terrestrial ice sheets if the northern Irish Sea Basin was fully marine. Undated linear sand ridges on the inner Celtic Sea (Pantin and Evans, 1984) cannot be used to estimate RSL but ice rafted debris derived from the Irish Sea in the Celtic Sea area (Scourse et al., 2009) confirms the continuity of the marine connection into the Irish Sea. The Isle of Lundy, the Bristol Channel and southwest England also appear to have been mostly ice free at the LGM (Rolfe et al., 2012) again suggesting that the Irish Sea ice stream was much less extensive than previously recognised (Bowen et al., 2002). 3.1.2. Major ice sheet readvances in northern Britain Early models of deglaciation of the BIIS inferred monotonic retreat following the LGM (Charlesworth, 1928) but Synge (1968, 1977) and McCabe (1969) proposed shifts in ice dispersion and associated readvances of ice sheets associated with terrestrial, subglacial, bedform overprinting and sediment fluxes towards tidewater margins (McCabe et al., 1987). The Last Glacial termination, however, is characterized by major regional readvances of ice sheet margins(McCabe et al., 1998; Clark et al., 2012b) although the largest (from the Younger Dryas to the early Holocene) was part of a wider global event (Smith et al., 2011, 2012). Ice margins in the northern Irish Sea Basin readvanced to terminal positions on at least three occasions: Clogher Head Stadial (ca 18 kya), the Killard Point Stadial maximum (ca 16.5 kya), and, the North Channel Readvance (15e15.5 kya) (McCabe and Clark, 1998; McCabe and Williams, 2012). The North Channel Readvance blocked the North Channel as far south as Belfast Lough and Stranraer and covered much of north and east Antrim, Ayrshire and as far west as Islay (McCabe and Williams, 2012). This dynamic pattern of expansion and contraction of BIIS is also seen in Scotland. Bradwell et al. (2008) recognise a very similar pattern of ice sheet activity in northwest Scotland where successive retreats and readvances continued until 16 kya and Ballantyne and Stone (2012) demonstrate that ice caps in the northwest highlands and reached fjords during the Older Dryas (ca 14 kya). Interstadial rates of sea level rise are estimated at ca 15 mm per year but during deglaciation these may increase to ca 50 mm per year (Cronin, 2012). RSL change was less in the northern Irish Sea but, in the Celtic Sea, linear sand ridges are thought to have formed when sea level was up to 120 m below present (Lambeck et al., 2002; Scourse et al., 2009). Dating of organic material from glacial deposits in the northern Irish Sea (Clark et al., 2012a,b), combined with the high rate of glacial retreat over water (Cronin, 2012), suggests that the sea maintained contact with the retreating ice sheet until Scottish and Irish ice sheets separated and the Irish Sea was fully formed at ca 16 kya (Clark et al., 2012a,b). McCarroll et al. (2010) highlight the dynamism of the Irish ice sheet, extending to the Scilly Isles, yet failing to reach the Preseli Hills, and retreating from the east coast of Ireland south of the Mourne Mountains by 18.5 kya. The Isle of Lundy and much of the Bristol Channel also appear to have been free of ice during the LGM (Rolfe et al., 2012) further illustrating the spatial complexity of the 148 W.I. Montgomery et al. / Quaternary Science Reviews 98 (2014) 144e165 development and movement of the ice sheet. Hence, it is possible that parts of coastal shelf of Britain and Ireland were ice-free quickly after LGM with southern Ireland, Wales and northern England exposed by ca 18 kya (Clark et al., 2012a,b). In contrast, the North Channel and parts of the inner Malin sea area were filled by readvance ice up to 16 kya (McCabe and Williams, 2012). During these late readvances, iceberg scour marks on the sea bed of the Irish Sea indicate continued discharge events from retreating ice sheet margins. Kelley et al. (2006) dated sea-level lowstand ( 30 m) from Belfast Lough to around 13.5 kya whilst more recent work suggests the general fall in the Irish Sea Basin and elsewhere may have been greater and oldest towards the south (~ 75 m) transforming the broad expanse of the Irish Sea Basin into two muddy, shallow, marine platforms (50e120 km wide) separated by a deep, central tidal trough. This trough could have been no wider than 40e50 km especially to the west of the Isle of Man and St. George's Channel. Thus, there is no evidence of topographically significant morainic ridges which could have provided pathways across the Irish Sea during the lowstand in RSL i.e. there is no support for a land bridge between Ireland and Britain in the Holocene (Mitchell, 1963). Ireland became a separate island no later than 15 kya prior to which the only likely area of contact with Britain was an ice bridge between northeast Ireland and southwest Scotland; all other contact between these islands was likely to have been lost before 19 kya during the earliest stages of deglaciation. 3.1.3. Sea level change in the Celtic Sea and English Channel The English Channel formed in the late Tertiary and periodically invaded by the sea during the Pleistocene (Lagarde et al., 2003). Hence, the English Channel long pre-dates the LGM. The presence of the Hurd Deep and a complex of canyons, palaeorivers and valleys running from west of Callais to Brest (Lagarde et al., 2003) form a significant physical barrier based around a major palaeoriver system called the Fleuve Manche by Bourillet et al. (2003). The origin and topography of the seabed of the English Channel change markedly from the western approaches to the siliciclastic deposits of periglacial and aeolian origin in the east (Reynaud et al., 2003). Modelling suggests that longitudinal ridges formed between 20 and 12 kya from sediments of the estuary of the Fleuve Manche and the Irish Sea ice stream (Scourse et al., 2009). The southern North Sea, Britain and the adjacent areas of present day France, Belgium and The Netherlands, were ice free in the late Pleistocene. Where the English Channel now lies, the Fleuve Manche drained meltwater westwards from the modern day Rhine, Thames, Seine and other rivers (Lericollais et al., 2003). A flattish plain was present along the northern and southern edges of the Fleuve Manche perhaps extending into the coastal plain in the northern part of the modern Bay of Biscay south of Brittany. The eastern English Channel had increasingly incised river valleys which increased depth below present sea level to 70 m. Further west, the Hurd Deep, was a broad expanse of deeper (170 m) water 150 km long by 2e5 km wide (Antoine et al., 2003; Lericolais et al., 2003). The substantial alluvio-glacial deposits in the western Celtic Sea (Scourse et al., 2009) suggest a major delta formation which would have retreated (transgressed) as sea level rose. As sea level rose further so flow rates declined to leave stable sand ridges (Lambeck et al., 2002; Scourse et al., 2009). Toucanne et al. (2008) suggested that runoff carrying glacial material into the Bay of Biscay by the Fleuve Manche also varied with expansion and contraction of the ice sheet. The eastern English Channel, however, reveals complex patterns of barriers and breaching of barriers during a further period of rapid sea level rise to ca 8 kya (Mellett et al., 2012) suggesting that major events in the English Channel progressed from west to east as RSL increased. 3.1.4. Sea level change in the southern North Sea e drowning of Doggerland As RSL increased land in the south of the present North Sea, Doggerland, started to contract, a process possibly exacerbated by a lake of melt-water (Clark et al., 2012a,b). This process of inundation continued rapidly in the first third of the Holocene, Doggerland succumbing to rising sea level ca 7.8 kya (Waller and Long, 2010; Mellet et al., 2012). The well-documented tsunami resulting from the Storegga landslip (8.2 kya) off the south-western coast of Norway, led to 20 m storm surges and whilst it may have made Doggerland uninhabitable by people (Weninger et al., 2008), it would not itself have led to any further rise in sea level (Julian Orford, pers. comm.). The coast of the Bay of Biscay was exposed to similar changes in sea level with evidence of incised rivers and ‘retrogradation of estuarine sediments’ (ca 9e7.5 kya) until they form part of a more recent, wider coastal environment (Estournes et al., 2012). 4. Dispersal barriers during postglacial recolonisation There were many fluctuations in mean global temperatures between 28 kya and 14 kya such that the ice sheets of northern Britain and Ireland advanced and retreated frequently and asynchronously (Clarke et al., 2012a,b). Ice free habitat may have been available in different places at different times and coastal strips may also have facilitated access to refugia for certain (cold-adapted) species, at least for short periods. However, less cold tolerant species would have been restricted to more southerly refugia during colder interludes (Stewart et al., 2010). As temperature rose, populations expanded out of these areas but the Pyrenees, Alps and other more easterly mountain ranges presented significant barriers to northward expansion of populations whilst numerous major rivers including the Danube, Rhine, Oder, Seine and Loire, made progress across western Europe difficult especially for smaller species. The Fleuve Manche would have presented another major physical barrier to land north and west of present day, continental Europe. However, this could be overcome in either of two ways. Firstly, until ca 8 kya it would have been possible for species to migrate further west across the southern North Sea via Doggerland linking Britain, Belgium and The Netherlands. Secondly, as Fleuve Manche surrendered its load in the deltaic region to the far west, there may have been some limited opportunity for even the smallest of terrestrial species to island hop as channels dried up and temporary islands merged with successfully invaded land. This process is perhaps more likely given decrease in flow rates that might have occurred ca 17.5e16 kya (Toucanne et al., 2008) and the protracted period of transgression as sea level rose and glacial material was redistributed. Mitchell's model of recolonisation of Britain and Ireland during the Holocene (Mitchell and Ryan, 1997) is not supported; there was no land bridge during the Holocene and it may not have existed at all or if it did it exist, it was of short duration and was much earlier than envisaged by Mitchell. Latest contact between Britain and Ireland, ca 15 kya, from southeast Ireland to southwest Wales has been proposed (Edwards and Brook, 2008) but this would have been a short duration, narrow, low-lying neck of land and seems increasingly unlikely given the speed at which the Irish Sea opened up after LGM and the southerly movement of icebergs. Between LGM and the isolation of Ireland at ca 15 kya, larger mammals might have been able to gain access across occasional temporary ice sheets forming during colder periods or particularly cold years. The last contact between Britain and continental Europe is also now much better fixed in time and location. The southern part of the North Sea was the last route into Britain until ca 8 kya. W.I. Montgomery et al. / Quaternary Science Reviews 98 (2014) 144e165 149 Table 1 (a) Irish mammal species, (b) British mammal species also found in Ireland, and (c) British mammal species not found in Ireland. Data based on available archaeological information and knowledge of species behaviour and ecology to infer putative means of arrival. Species are arranged in chronological order of appearance in Ireland as the likely older of the two islands (see text). Species found in Britain but not Ireland, are listed separately following the same inferences regarding arrival in Britain. Details relating to material, sites, dating and stratigraphy, and historical records, can be found in van Wijngaarden-Bakker (1974), Stuart and Wijngaarden-Bakker (1985) and Yalden (1999), with additional material from The Irish Quaternary Fauna Project (Woodman et al., 1997), McCormick (1999), Yalden and Kitchener (2008) and Sleeman and Yalden (2009). Further data on particular species are included where available e.g. giant deer, Megaloceros giganteus, (Stuart et al., 2004); fallow deer, Dama dama (Sykes, 2010); red deer, Cervus elaphus (Carden et al., 2012) and the brown bear, Ursus arctos (Edwards et al., 2011). (a) Ireland Early Holocene extinctions Early Holocene survivors, later extinctions, and arrivals Early Holocene bats, later Arrivals Introductions, last Millenium, present Century People, companion, Animals domestic stock Species/taxon giant deer Arctic fox reindeer Collared lemming Mountain/Irish hare Irish stoat grey wolf brown bear red deer wild boar wild cat lynx wood mouse pygmy shrew otter red fox badger pine marten house mouse ship rat Natterer's bat Daubenton's bat Leisler's bat common pipistrelle brown long eared bat whiskered bat lesser horseshoe bat soprano pipistrelle Nathusius' pipistrelle rabbit red squirrel hedgehog fallow deer brown rat European hare Sika deer grey squirrel bank vole American mink greater white toothed shrew Muntjac deer People domestic dog domestic cattle domestic sheep domestic pigs feral goat domestic horse domestic cat Scientific name Megaloceros giganteus Alopex lagopus Rangifer tardanus Dicrostonyx torquatus Lepus timidus hibernia Mustela erminea hibernica Canis lupus Ursus arctos Cervus elaphus Sus scrofa Felis silvestris Lynx lynx Apodemus sylvaticus Sorex minutes Lutra lutra Vulpes vulpes Meles meles Martes martes Mus domesticus Rattus rattus Myotis nattereri Myotis daubentonii Nyctalus leisleri Pipistrellus pipistrellus Plecotus auritus Myotis mystacinus Rhinolophus hipposideros Pipistrellus pygmaeus Pipistrellus nathusii Oryctolagus cuniculus Sciuris vulgaris Erinaceus europaeus Dama dama Rattus norvegicus Lepus europaeus Cervus Nippon Sciuris carolinensis Myodes glareolus Mustela vison Crocidura russula Muntiacus spp. Homo sapiens Canis familiaris Bos taurus Os aries Sus scrofa domesticus Capra hircus Equus ferus caballus Felis cattus Irish archaeological/historical Earliest record/presumed present Latest record/extant* 14.1 kya 13 kya 13 kya 13 kya 28e12.1 kya 27e10.6 kya 13e9.9 kya 13e9.9 kya 11.8 kya or 5 kya ? 9.2 kya 9 kya 8.9 kya 7.6 kya 5 kya 4 kya 3.8 kya 3.8 kya 2.8 kya 2.5 kya Intro 1.7 kya Intro Na Na Na Na Na Na Na Recent discovery Recent discovery 12th C AD Intro. 12th C AD Intro.; Reintro. 18th C AD? 12th C AD Intro. 13th C AD Intro. 18th C AD Intro. 19th C. AD Intro. 19th C. AD Intro. 20th C AD Intro. 20th C AD Intro. 20th C AD Intro. 21st C AD Intro. 21st C AD Intro. 8.3 kya 11.7 kya 13 kya 10.3 kya 10.3 kya * * 17th C AD 5 kya * 13th/14th C AD 3 kya * * * * * * * * * * * * * * * * * * * (17th ext) * * * * * * * * * * * * * * * * * * 6.6 kya 5 kya 5 kya 4.5 kya ? 4 kya ? (b) Britain Early Holocene extinctions Early Holocene survivors, later extinctions, and arrivals Species/taxon giant deer Arctic fox reindeer Arctic lemming mountain hare stoat grey wolf brown bear red deer wild boar Scientific name Megaloceros giganteus Alopex lagopus Rangifer tardanus Dicrostonyx torquatus Lepus timidus Mustela erminea Canis lupus Ursus arctos Cervus elaphus Sus scrofa British archaeological/historical Earliest record/presumed present Latest record/extant* 15.8 kya 13 kya 13 kya 8.3 kya 13 kya 12.9 kya 13e11 kya 13e9.9 kya 13e9.9 kya 13e9.9 kya 13e11 kya 11.7 kya * 10.4 kya * * 17th C AD 4th C AD * 13th/14th C AD (continued on next page) 150 W.I. Montgomery et al. / Quaternary Science Reviews 98 (2014) 144e165 Table 1 (continued ) (b) Britain Early Holocene bats, later arrivals Introductions, last Millenium, present Century People, companion, animals domestic stock Species/taxon wild cat lynx wood mouse pygmy shrew otter red fox badger pine marten house mouse ship rat Natterer's bat Daubenton's bat Leisler's bat common pipistrelle brown long eared bat whiskered bat lesser horseshoe bat soprano pipistrelle Nathusius' pipistrelle rabbit red squirrel hedgehog fallow deer brown rat European hare Sika deer grey squirrel bank vole American mink greater white toothed shrew Muntjac deer people domestic dog domestic cattle domestic sheep domestic pigs feral goat domestic horse domestic cat Scientific name Felis silvestris Lynx lynx Apodemus sylvaticus Sorex minutes Lutra lutra Vulpes vulpes Meles meles Martes martes Mus domesticus Rattus rattus Myotis nattereri Myotis daubentonii Nyctalus leisleri Pipistrellus pipistrellus Plecotus auritus Myotis mystacinus Rhinolophus hipposideros Pipistrellus pygmaeus Pipistrellus nathusii Oryctolagus cuniculus Sciuris vulgaris Erinaceus europaeus Dama dama Rattus norvegicus Lepus europaeus Cervus Nippon Sciuris carolinensis Myodes glareolus Mustela vison Crocidura russula Muntiacus spp. Homo sapiens Canis familiaris Bos taurus Os aries Sus scrofa domesticus Capra hircus Equus ferus caballus Felis cattus British archaeological/historical Earliest record/presumed present Latest record/extant* 9 kya 9.5 kya 9.5 kya 9.9 kya 9.9 kya 12.3 kya 9.9 kya 10 kya 2.5 kya Intro 1.7 kya Intro 9.9 kya 9.9e5 kya 9.9 kya Na 9.9 kya 9.9 kya 6e7 kya Rec discovery Rec discovery 12th C AD Intro. 8.7 kya 9.5 kya 11th C AD Intro. 18th C AD Intro. 2.5 kya Intro. 20th C AD Intro. 19th C AD Intro. 9.9 kya 20th C AD Intro. Abs 20th C AD Intro. 12 kya 9.5 kya 5.5 kya 5.4 kya 5 kya 4.5 kya 3.7 kya 2.5 kya * 6th C AD * * * * * * * * * * * * * * * * * * * * * * * * * * * Abs * * * * * * * * * (c) Britain Species/taxon scientific name Early Holocene extinctions Norway lemming steppe pika root vole wild horse elk beaver aurochs common shrew mole harvest mouse water vole field vole weasel polecat roe deer water shrew hazel dormouse common vole yellownecked mouse Brandt's bat mouse eared bat Bechstein's bat noctule bat grey long eared bat barbastelle bat serotine bat greater horseshoe bat feral ferret Lemmus lemmus Ochotoma pusilla Microtus oeconomus Equus ferus Alces alces Castor fiber Bos primigenius Sorex araneus Talpa europaea Micromys minutus Arvicola terrestis Microtus agrestis Mustela nivalis Mustela putorius Capreolus capreolus Neomys fodiens Muscardinus avellanarius Microtus arvalis Apodemus flavicollis Myotis brandtii Myotis myotis Myotis bechsteinii Nyctalus noctula Plecotus austriacus Barbastella barbastellus Eptesicus serotinus Rhinolophus ferrumequinum Mustela furo Early Holocene survivors, later extinctions, and arrivals Early Holocene bats, later arrivals Introductions, British archaeological/historical Earliest record/presumed present Latest record/extant* 13 kya 13 kya 13 kya 13 kya 13 kya 9 kya 12 kya 11 kya 13 kya 9.9 kya 9.9 kya 9.9 kya 7.5 kya? 11e 5 kya? 10 kya 9 kya? 9 kya EOrk 5.5 kya; Gurnsey 9.9 kya ? 5 kya na na 3.5 kya 9.9 kya na 9.9 kya 3.5 kya 2 kya 12th C AD Intro. 10.4 kya 9.9 kya 9.9 or 5 kya 9.3 kya 3.9 kya 1.0 kya 3.2 kya * * * * * * * * * * * * * * * * * * * * * W.I. Montgomery et al. / Quaternary Science Reviews 98 (2014) 144e165 151 Table 1 (continued ) (c) Britain last Millenium, present Century Species/taxon edible dormouse coypu Chinese water deer scientific name British archaeological/historical Glis glis Myocastor coypus Hydropotes inermis 5. Human history in the Holocene Human migrations and trade, played a significant role in the translocation of many species during the last 10,000 years (Grayson, 2001). Childe (1925) established an outline for successive cultures e Palaeolithic (end ca 10 kya), Mesolithic (end ca 6 kya) and Neolithic (end 4 kya) e emerging across Europe. Whilst there are Palaeolithic remains from Britain (Jacobi and Higham, 2011a,b) there are none from Ireland. Mesolithic hunter gatherers appeared in Ireland ca 9 kya (Mitchell and Ryan, 1997) but may have been in Britain from as early as 11 kya (Conneller et al., 2012). These people migrated across Europe from the southeast and would have travelled by land into Britain and then, by rudimentary dug-out canoes, to Ireland. The first (Neolithic) farmers appear in both islands ca 6.5 kya (Mitchell and Ryan, 1997); if anything, the onset of farming dated by remains of livestock was slightly earlier in Ireland than Britain (Woodman et al., 1997; Yalden, 1999). Neolithic people were distinct genetically from the earlier migrants into Europe (Bramanti et al., 2009), originating in the southeast of Europe and bringing a new culture (Childe, 1925) and languages (Bouckaert et al., 2012). However, they did not replace the earlier hunteregatherers but interbred such that there is a lower presence of ‘farming’ genotypes and a higher presence of ‘hunteregatherer’ genotypes in modern northwest human populations compared to southeast Europe (Currat and Excoffier, 2005; Belle et al., 2006; Skoglund et al., 2012). Neolithic people developed skills in recovering metals from native ores such that the Bronze Age commenced in Britain and Ireland at ca 4.5 kya and Iron Age around 2.7 kya. Britain but not Ireland was occupied by the Romans for ca 500 years from just over 2 kya. The early Christian period to 1.2 kya involved voyages around the west of Europe, trade and monastic settlements. The Vikings contributed further to these maritime peregrinations along the full length of the western coastline of Europe from 1.2 to 1 kya. Continental influence was enhanced by Norman incursions in both Britain and Ireland between 0.9 and 0.8 kya whilst voyages of discovery commenced 0.7e0.5 kya. The Plantation of Ireland and particularly Ulster was 0.3e0.4 kya with an influx of people from England and Scotland. The industrialisation of Britain exceeded that of Ireland and trade directly with overseas colonies and Europe was probably also greater in Britain than Ireland during the 18th and 19th centuries AD, although Dublin was recognised as one of several ‘second cities’ of the United Kingdom as well as the Empire. The last 100 years involved both islands in European and World wars as well the globalisation of trade and overseas leisure travel. Whilst Britain and Ireland have a largely common, shared history (Morgan et al., 2005), their differences may be important biogeographically. There are only slight disparities in the arrival of early people and farming but the 500 year period of Roman occupancy in Britain and their absence from Ireland may be the biggest disparity in the history of these islands as people tend to bring animal and plant species that are important to them when they move to new areas (Grayson, 2001). Britain is also much more the focus of European and world trade than Ireland and supports a much larger, more urban population (Eurostat, 2012) which could Earliest record/presumed present Latest record/extant* 20th C AD Intro. 20th C AD Intro 20th C AD Intro * 1990 AD * increase the rate at which accidental introductions might occur. The far greater economic activity of Britain in comparison to Ireland might also increase the likelihood of importation and eventual ‘naturalisation’ of exotic, non-european species. 6. Archaeology of the mammals of Britain and Ireland Details relating to material, sites, dating and stratigraphy, and historical records, can be found in: van Wijngaarden-Bakker (1974); Stuart and Wijngaarden-Bakker (1985); Yalden (1999); The Irish Quaternary Fauna Project (Woodman et al., 1997); McCormick (1999); Yalden and Kitchener (2008); and Sleeman and Yalden (2009). Further data on particular species are included where available, namely; giant deer, Megaloceros giganteus, (Stuart et al., 2004); fallow deer, Dama dama (Sykes, 2010); red deer, Cervus elaphus (Carden et al., 2012); and the brown bear, Ursus arctos (Edwards et al., 2011). Archaeological data are used to establish the earliest known time of arrival of a species in Britain and Ireland and, where relevant, the latest record of a mammal species is used to establish the earliest date after which it was extinct (Table 1). Dates are based on calibrated carbon dating, comparable stratigraphy and historical records. Domestic species are also considered where they indicate the arrival of farming and had a significant ecological impact in Britain and Ireland. There are few data for bats in the British record and none at all in the Irish. Bats are an important component of temperate woodland communities and capable of long distance dispersal over water as well as land. Hence, their time of arrival during postglacial recolonization has been set at when suitable habitat became available. Many British and Irish bats favour woodland, rivers and lake edge (Walsh and Harris, 1996; Vaughan et al., 1997; Warrant et al., 2000; Russ and Montgomery, 2002) and make use of natural roosts in tree holes (Boonman, 2000; Ruczynski and Bodanowicz, 2005; Spada et al., 2008) and natural swarming sites such as caves (Parsons et al., 2003). Such conditions would have been spreading northwards across Britain and Ireland (Mitchell and Ryan, 1997; Whitehouse, 2006) from ca 9.5 kya and, hence, this date is taken as the time of first arrival of all bats on both islands except where there are data to the contrary. Vera (2000) has argued that the early Holocene forest would not have been continuous but contained gaps comprising lower, more open vegetation. Mitchell British and Irish species by taxon 25 20 15 10 5 0 Insectivora Chiroptera Carnivora Lagomorpha Rodentia Artiodactyla Fig. 2. Numbers of British (black) and Irish (white) mammal species of named taxa. 152 W.I. Montgomery et al. / Quaternary Science Reviews 98 (2014) 144e165 Species by Origin and Island 25 20 15 10 5 0 EH Ext EH Sur E Bat L Bat E Econ Acc In Del In Fig. 3. Numbers of British and Irish mammal species by putative origin based on archaeological material and historical records. EH early Holocene; Ext extinct; Sur surviving; E. Bat early bat; L Bat Late bat; E. Econ early economic; Acc In accidental introduction; Del In deliberate introduction. Origin of British and Irish mammals 60 50 40 30 20 10 0 Pre Man Meso/Neo/Brz Commensal Norman Late intro mode of arrival Fig. 4. Numbers of British and Irish mammal species by putative origin based on date of archaeological material and historical records. Pre Man before arrival of people during the Holocene; Meso/Neo/Brz arrival at time of Mesolithic, Neolithic and Bronze age people; Commensal arrival of commensal rodents; Norman arrival with Norman conquest; Late intro introduction from mid 19th century onwards. (2005), however, argues that forests were opened up only with the arrival of livestock. Whitehouse and Smith (2010) present convincing evidence that limited areas of open vegetation were present during the early Holocene although forests became more closed towards the onset of the Neolithic. Regardless of these details in relation to the homogeneity of forest cover, many temperate bats forage along ecotones such as lake, river and forest edges and it is difficult to believe that bats arriving in Ireland and Britain early in the Holocene did not find abundant suitable habitat. Britain has five bat species that are more recent arrivals, all with southern distributions, and Ireland, just one, again suggesting bats were present from the early Holocene in both Britain and Ireland. Comparison of the total mammal species in Britain and Ireland (Fig. 2) suggests that the latter enjoys parity only with respect to lagomorphs and is particularly deficient relative to Britain in rodents and bats. When British and Irish mammal species are arranged by putative time of appearance (Table 1, Fig. 3), the major disparity is early with regard to both those species which survived the end of the Pleistocene or arrived early in the Holocene. The importance of the late Pleistocene, early Holocene transition in characterizing the British and Irish mammals is even more dramatic if species are separated along a crude time line starting with those that arrived before people, around the same time as the Mesolithic and Neolithic peoples and the Bronze Age, and later arrivals (Fig. 4). Figs. 3 and 4 also suggest that in proportion, but not number of species, Ireland has more species with potential economic value as food or for their pelt, arriving contemporaneously with early people. Similarly, Ireland is subject to proportionally greater impact by more introduced species in historical times. Hence, archaeological data are consistent with early people introducing mammal species they needed into Ireland much as early settlers brought plants and animals they needed to islands in the Pacific Ocean (Semah and Detroit, 2006; Dening, 2007; Larson et al., 2007a,b; Fitzpatrick and Callaghan, 2009; Fall and Drezner, 2011) and Mediterranean islands (Schule, 1993; Van de Noort, 2003; Farr, 2006; Dubey et al., 2007; Masseti, 2009; Vigne et al., 2009; Knapp, 2010; Rowley-Conwy, 2011). Despite having a much smaller human population throughout the Holocene and a much lower level of world trade than Britain (Eurostat, 2012), Ireland has acquired just as many introduced species. 7. Morphometric analyses Morphometric analyses are used to identify relationships between populations within species based on phenotypic characteristics including qualitative and quantitative cranial features and dentition (Cardini and O'Higgins, 2004; Murphy et al., 2006; Wroe and Milne, 2007; Monteiro and Nogueira, 2011). Although used less frequently in the molecular genetic era, morphometric analyses in conjunction with genetic analyses may help determine differences in the origin of populations and the evolutionary processes that lead to reproductive isolation and eventual speciation or introgression and enhanced genetic diversity within species (Edwards et al., 2011; Carden et al., 2012; Tougard et al., 2013). There are also generalizations made with respect to latitude and climate and variation in body mass, Bergmann's Rule (Ashton et al., 2000; Meiri and Dayan, 2003), aspects of shape (Jungers et al., 1995; Barrow and Macleod, 2008; McGuire, 2010) and colouration, Golger's Rule (Stoner et al., 2003a, 2003b; Lai et al., 2008; Elton et al., 2010; Kamilar and Bradley, 2011) or with discontinuities in physical and biological constraints as in island populations in comparison to continental populations of the same species (Adler and Levins, 1994; Michaux et al., 2002; Lomolino, 2005; Dawson and Milne, 2012). However, there may be as many exceptions to these generalizations as species (Meiri et al., 2004, 2006). A review of morphological and morphometric data relating to British and Irish populations making comparisons between continental and island populations, suggests that in most British and Irish mammals there is much more variation within a population or biogeographical zone or continuous variation along a geographical cline across Europe, than consistent, discrete differences among continental Europe, Britain and Ireland. Most British and Irish species are indistinguishable morphologically from each other and from continental populations. Perhaps nine species are worth considering as having unique island forms in Britain and/or Ireland and, hence, have been in either island sufficiently long to show local adaptation and divergence from ancestral stock. These are described here in order of decreasing effect based largely on Harris and Yalden (2008). The Irish (mountain) hare Lepus timidus hibernicus is larger than its Scottish counterpart, has a redder coat colour, completely white tail and rarely moults to white in winter (Barrett-Hamilton and Hinton, 1910). Neil Reid (pers. comm.) has also shown that lower jaw shape of Irish hare is different from other mountain hares. It also exploits a wider range of habitats including lower land down to sea level and is more dependent on grasses for forage (Whelan, 1985; Dingerkus and Montgomery, 2001). The Irish stoat Mustela erminea hibernica is smaller than its British counterpart but larger than continental stoats (Fairley, 1981; McDonald, 2002). The Irish form also has a distinctive irregular line along its flank where darker dorsal fur abuts white ventral fur. The Irish otter Lutra lutra roensis shows greater sexual dimorphism in body size than British W.I. Montgomery et al. / Quaternary Science Reviews 98 (2014) 144e165 otters from which it can also be discriminated using cranial criteria (Lynch et al., 1996). Irish otters have darker fur than British ones which have darker fur than continental ones (Dadd, 1970). The badger Meles meles in Ireland is lighter than in Britain (Lynch et al., 1993) and British badgers have larger skulls than continental ones (Lynch et al., 1997). Similarly, British pine martens Martes martes are larger and have larger skulls than continental forms (BarretteHamilton and Hinton, 1910; Miller, 1912; Jeffries and Critchley, 1994). British water voles Arvicola terrestris are larger than their continental counterparts but Scottish water voles are smaller than English ones (Corbet, 1970). Many red squirrels in Britain and Ireland have ‘bleached’ ear tuffs and tail, a characteristic which has been used to place them in a separate sub-species Sciuris vulgaris leucorus (Harris and Yalden, 2008). Skulls of red and roe deer, C. elaphus and Capreolus capreolus, from Britain, are distinguishable from continental conspecifics (Lowe and Gardner, 1974; Hewison, 1997) and presumed ancient forms of red deer from Kerry (Ireland) can be distinguished from Scottish conspecifics using craniometrics (Carden et al., 2012). A number of other species show morphological traits on some smaller islands around Britain: greater white-toothed shrew Crocidura russula from Guernsey are smaller than continental conspecifics (Delany and Healy, 1966); the extinct St Kilda house mouse Mus musculus muralis was notably larger than others and sufficiently unique to enjoy sub-specific status (BarretteHamilton, 1899); and, common (Orkney) vole Microtus arvalis orcadensis/sandayensis is larger than continental forms (Corbet, 1986). Greater horseshoe bat Rhinolophus ferrumequinum and harvest mouse Micromys minutus in Britain are smaller in body size than some continental populations (Harris and Yalden, 2008). 8. Phylogeography of mammals of Britain and Ireland There has been a surge in phylogeographical information regarding the spatial relationships of British and Irish mammals since the late 1990s due to: greater availability of mitochondrial and nuclear markers (Carvahlo, 1998; Thomson et al., 2010); useful ancient DNA sequences (Willerslev and Cooper, 2005); noninvasive methods using hair and other tissue; PCR to amplify small amounts of DNA (Swenson et al., 2011; Rodgers and Janecka, 2013); and, advanced genomic (McCormick et al., 2013) and statistical methods (Jako et al., 2009; Ho and Shapiro, 2011). The judicious application of the ‘molecular clock’ (Ho et al., 2005, 2011) has shed light on the timing of recolonisation after LGM (time of lineages separating or date of last common ancestor) and, indeed, throughout the Quaternary. Published and unpublished studies (successfully completed doctoral theses and externally evaluated reports to Governments) yielded useful data for 40 extant or extinct mammal species from Britain and Ireland. Whilst almost all studies had continental samples, there were fewer reporting data from Britain and Ireland such that useful studies were available for 27 species from Britain and Ireland and a further 14 from Britain alone. Most studies (37) drew on recent material only; seven used both ancient and recent DNA and two used only ancient DNA. Overall, 43 studies reported data based on mitochondrial DNA and 23 on nuclear DNA. Twenty three used both mitochondrial and nuclear DNA and 29 studies used two or more markers. The most commonly used markers were Cyt b (24 samples) and microsatellites (24 samples). Additional common markers were from the Control Region of mitochondrial genome and sequences from the X and Y chromosomes. Phylogeographical studies are vulnerable to sampling difficulties, inconsistency and variation in conditions deployed in 153 resolution of markers, scoring, and reporting (Estoup et al., 2002; Morin et al., 2004; Toews and Brelsford, 2012), whilst the ‘molecular clock’ is sensitive to mutation rates used (Galtier et al., 2009; Lanfear et al., 2010), such that comparisons across a large number of studies on different species using different markers and conducted by different laboratories, can only discern major spatial and temporal signals during shorter and more recent evolutionary periods. This review focuses on to what extent phylogeography reflects the archaeological and historical records and also how well it reveals mechanisms of recolonisation that are consistent with recent ideas in physiography and environmental change during the Pleistocene and Holocene. There are recurring phylogeographical features which suggest that recolonisation of Europe by different mammal species during the late Pleistocene and the Holocene was affected by natural and anthropogenic influences, in a similar manner. (1) Earlier oscillations of ice during the Quaternary affected mammal phylogeography across Europe as well as LGM e.g. Apodemus spp. (Michaux et al., 2003, 2004), Irish hare (Hughes et al., 2009), stoat (Martinkova et al., 2007), pine marten (Ruiz-Gonzalez et al., 2013), greater (Flanders et al., 2009) and lesser (Dool et al. in prep.) horseshoe bats, and red fox (Edwards et al., 2012). (2) There is evidence of replacement of earlier lineages by later ones e.g. stoat (Martinkova et al., 2007), wolf (Pilot et al., 2010), and pygmy shrew, common shrew and bank vole (Searle et al., 2009). (3) Phylogeographical patterns in commensal (Rajabi-Mahon et al., 2007; Searle et al., 2008), domestic (Larson et al., 2007) and other mammal species (McDevitt et al., 2009; Stamatis et al., 2009), to a greater or lesser degree, parallel changes in human populations and migration (Currat and Excoffier, 2005; Skoglund et al., 2012) or trade (Mitchell and Ryan, 1997). (4) Genetic affinities are present between samples from Britain and Ireland in brown bear (Edwards et al., 2011), red deer (Carden et al., 2012), pine marten (Kyle et al., 2003), badger (Pope et al., 2006), otter (Finnegan and O'Neill, 2010), hedgehog (Hewitt, 1999) and pygmy shrew (McDevitt et al., 2011). (5) Less anticipated links between samples from Britain and/or Ireland and parts of Scandinavia are evident in badger (O'Meara et al., 2012; Frantz et al., 2014), pygmy shrew (McDevitt et al., 2011), weasel (Lebarbenchon et al., 2010), Orkney vole (Haynes et al., 2003), water vole (Piertney et al., 2005), house mouse (Searle et al., 2008) and recent populations of red squirrel (Hale et al., 2004; Finnegan et al., 2008). (6) There are genetic associations involving Britain, Ireland and Spain (Andorra) in pine marten (Davison et al., 2001; Ruiz-Gonzalez et al., 2013), badger (O'Meara et al., 2012), pygmy shrew (Mascheretti et al., 2003) and weasel (Lebarbenchon et al., 2010). The most effective way of identifying common patterns and interspecific differences is to consider how consistent these empirical studies are with hypotheses which make predictions regarding variation in genetic variation among samples from different locations, in this case, particularly, continental Europe, Britain and Ireland. Here, six non-mutually exclusive hypotheses are outlined on which predictions are made with respect to genetic diversity (columns 1 and 2, Table 2). After examination of the available data conclusions are made regarding consistency and inconsistency of each species with these predictions (columns 3 and 4, Table 2). In some cases, there is strong evidence for consistency or inconsistency but in some species there are few data which have been used with caution and only where these data are persuasive. Consistency with observations does not in any sense verify a hypothesis: future data could come to light that might be inconsistent with a hypothesis. Here, species are treated as wholly independent: congeneric species and other close relatives often show different phylogeographic patterns whilst distantly related mammals may show similarities. 154 W.I. Montgomery et al. / Quaternary Science Reviews 98 (2014) 144e165 Table 2 Six non-mutually exclusive hypotheses (Column 1) are outlined on which predictions (Column 2) are made with respect to genetic diversity of mammal species occurring in Britain and/or Ireland. All available genetic data are used to assess consistency (Column 3) an inconsistency (Column 4) of mammal species with these predictions (columns 3 and 4). Hypotheses Predictions Consistent species 1-5 1. Postglacial colonization of Ireland was from Britain and in turn Britain derived its mammals from continental Europe Genetic diversity of continent populations is greater than those of Britain which in turn is greater than those of Ireland with few unique forms in the latter pygmy shrew , lesser horseshoe bat6, Natterer's bat7-9, whiskered bat10, common pipistrelle11,12, soprano pipistrelle11,12 2. Mammal populations of Ireland and/or Britain reflect one or more translocation by people pygmy shrew1-5, greater white toothed shrew36 hedgehog37,38, brown hare39, red squirrel 18,19, bank vole24, red deer26,28, Sika deer26, pine marten29,30,40, badger41,31, 3. Postglacial Irish populations not derived from Britain but directly from a continental or other refugium Genetic diversity in Irish and/or British populations very limited in comparison to continental ones suggesting strong founder effect; conversely, admixtures of diverse origin suggesting more than one translocation Irish populations markedly divergent genetically from British ones possibly with continental associations 4. Irish populations of ancient origin predating LGM surviving in ‘cryptic’ refugia i.e. refugia exposed by lower RSL Irish populations many unique alleles and/or haplotypes not shared with populations in Britain or continental Europe Leisler's bat14, Daubenton's bat13, Irish hare17, stoat34, brown bear42,43 5. Postglacial Irish mammals derived from refugia in south and west of Europe. Mammal species that occur in Britain species but are not indigenous to Ireland originate from south east or eastern European refugia (a) Genetic association of Irish samples with samples from south and west of Europe suggesting recolonisation predominantly via western routes (b) Strong genetic links between British and eastern European samples in mammals absent from Ireland 6.Postglacial recolonisation of Britain and Ireland from southern refugia without any indication of additional refugia in northern, central or western Europe Genetic links only from British and Irish samples to southern refugia (a) hedgehog37,38, pygmy shrew1-5, lesser horseshoe bat6, common pipistrelle11,12, pine marten29,30,40, badger44,41,31 (b) mole47,48, common shrew49,3, greater horseshoe bat50,51, brown hare52,39, bank vole53, field vole5456 , water vole57-59, yellow-necked mouse60,20, weasel61,62, roe deer63, aurochs64, lynx65, arctic fox66 mole47,48, hedgehog37,38, common shrew67,3, whiskered bat10, lesser horseshoe bat6, soprano pipistrelle11,12, brown hare52,39, bank vole 58,59,68, field vole54-56, Orkney vole69, wood mouse70,20,21,71,22, wild boar72, red deer26-28, pine marten29,30,40, red squirrel19, weasel61,62 pygmy shrew1-5, Leisler's bat14, Daubenton's bat13, Irish hare15-17, house mouse23, pine marten29,30,40, otter33,76, stoat34, Brown bear42,43 Inconsistent species Daubenton's bat13, Leisler's bat14, Irish hare15-17, red squirrel18,19, wood mouse20-22, house mouse23, bank vole24, red deer25-28, pine marten29,30, badger31,32, otter33,76, stoat34, red fox35 lesser horseshoe bat6, whiskered bat10, Daubenton's bat13, Natterer's bat9, Leisler's bat14, common pipistrelle11,12, soprano pipistrelle11,12, Irish hare15,17, wood mouse21,22, house mouse23, otter33,76, stoat34, red fox35 Hedgehog37,38, lesser horseshoe bat6, whiskered bat10, Natterer's bat9, common pipistrelle11,12, soprano pipistrelle11,12, bank vole24, red squirrel19, red deer26,28, Sika deer26, badger44,41,32, red fox35,45 pygmy shrew1-5, hedgehog37,38, lesser horseshoe bat6, whiskered bat10, Natterer's bat9, common pipistrelle11,12, soprano pipistrelle11,12, red squirrel19, bank vole24, Sika deer26, pine marten29,30,40, badger44,41,31, otter33,46, red fox35,45 (a) Leisler's bat14, Natterer's bat7,8, Irish hare15,17, Sika deer26, otter33,46, stoat34, red fox35,45 (b) soprano pipistrelle11,12 Daubenton's bat13, Leisler's bat14, Natterer's bat7,8, greater horseshoe bat50,51, common pipistrelle11,12, Irish hare15,17, water vole57, house mouse73,23, roe deer63, Sika deer26, aurochs64, otter33,46, badger31, stoat34, lynx74,65, brown bear42,43, wolf75, red fox35,45, arctic fox66 1 Mascheretti et al., 2003; 2McDevitt et al., 2009b; 3Searle et al., 2009; 4McDevitt et al., 2011; 5Vega et al., 2010; 6Dool et al. in prep; 7Salicini et al., 2011; 8Salicini et al., 2013; Scott, 2012; 10Boston et al., 2011; 11Hulva et al., 2010; 12Boston et al. in prep; 13Atterby et al., 2010; 14Boston et al. in revision; 15Hamill et al., 2006; 16Melo-Ferreira et al., 2007; 17Hughes et al., 2009; 18Hale et al., 2004; 19Finnegan et al., 2008; 20Michaux et al., 2005; 21Berckmoes et al., 2005; 22Booth et al., 2009; 23Jones et al., 2011, 2012; 24Stuart et al., 2007; 25Hmwe et al., 2006; 26McDevitt et al., 2009a; 27Skog et al., 2009; 28Carden et al., 2012; 29Davison et al., 2001; 30Kyle et al., 2003; 31O'Meara et al., 2012; 32Frantz et al., 2014; 33Finnegan and Neill 2010; 34Martinkova et al., 2007; 35Edwards et al., 2012; 36Allan McDevitt pers., comm; 37Hewitt 1999; 38Seddon et al., 2001; 39Stamatis et al., 2009; 40Ruiz-Gonzalez et al., 2013; 41Pope et al., 2006; 42Davison et al., 2011; 43Edwards et al., 2011; 44Marmi et al., 2006; 45Kutschera et al., 2013; 46Mucci et al., 2010; 47 Tryfonopoulis et al., 2009; 48Colangelo et al., 2010; 49Yannic et al., 2008; 50Rossiter et al., 2007; 51Flanders et al., 2009; 52Kasapidis et al., 2005; 53Deffontaine et al., 2005; 54 Jaarola and Searle 2002; 55Jaarola and Searle 2004; 56Herman and Searle 2011; 57Piertney et al., 2005; 58Kotlik et al., 2006; 59Wojcik et al., 2010; 60Michaux et al., 2004; 61 Lebarbenchon et al., 2010; 62McDevitt et al., 2012; 63Lorenzeni and Lovaria 2006; 64Edwards et al., 2007; 65Schmidt et al., 2011; 66Dalen et al., 2007; 67White and Searle, 2008; 68Deffontaine et al., 2009; 69Haynes et al., 2003; 70Michaux et al., 2003; 71Dubey et al., 2008; 72Scandura et al., 2008, 2011; 73Searle et al., 2008; 74Hellborg et al., 2002; 75Pilot et al., 2010; 76Ferrando et al., 2004. 9 Consistency and inconsistency with hypotheses 1e6 are also summarised for mammal species that occurred or occur in Ireland and Britain or in Britain alone in Table 3 wherein species have been placed in putative date of arrival in Ireland as: pre-Holocene, Mesolithic, Neolithic, Bronze Age, Norman or Recent (<150 ya). These ascriptions are based on genetic, archaeological, biological and historical information and, hence, it is not intended as an exhaustive treatment of all British and Irish mammal species. Time from most recent common ancestor (TMRCA), divergence time or time of expansion of a lineage (Arbogast et al., 2002; Ricklefs, 2007; Liu et al., 2009) are illustrated for the last 100,000 years and last million years in Fig. 5a and b respectively. These figures illustrate 95% limits of the estimates of divergence time with species arranged from least to oldest TMRCA or expansion time. It is clear that divergence times in many species is broadly contemporaneous with environmental changes during the period following the LGM whilst others suggest that isolation and divergence occurred much earlier in the Quaternary e.g. Apodemus spp. Some, notably Irish hare and stoat, show TMRCA estimates that predate LGM which are consistent with early origins in comparison with pygmy shrew and W.I. Montgomery et al. / Quaternary Science Reviews 98 (2014) 144e165 155 Table 3 Irish and British mammal species reflecting consistency or inconsistency with hypotheses 1e6 (see Table 2) regarding genetic diversity in phylogeographic studies. Putative origins are based on tests of all phylogeograhic hypotheses where sufficient data are available, and known and inferred presence in Ireland based on archaeology, ecology and historical records. Hypotheses 1e6 are arranged from left to right on basis of inferred length of presence with Ireland from pre- LGM to present. Natterer's bat and, particularly, house mouse. Whilst the spread of TMRCA values is such that it is difficult to determine whether divergence occurred before or after e.g. arrival of people, there is general concordance between temporal and spatial phylogeography and with archaeological and historical records. However, precision of estimate is generally poor reflecting problems in estimating and applying appropriate rates of mutation at this shallow level of evolution (Ho et al., 2005, 2011). 9. Chronology of mammals in Britain and Ireland since LGM Combining physiographic, archaeological, morphological and genetic evidence allows tentative conclusions regarding the arrival of the Holocene mammal fauna in Britain and Ireland from the LGM to the present. This chronology and understanding of the processes at work are imprecise and speculative but it is an attempt to accommodate the balance of recent evidence as fully as possible. The major stages are: (1) LGM to isolation of Ireland at ca 15 kya when a subarctic mammal fauna would have been present across the region; (2) temporary warming to 14 kya when more coldadapted species declined or were lost from the islands arising along southwestern and western parts of the Atlantic coast of Europe, whilst warm-adapted species continued to arrive in Britain across land in the southern part of the present North Sea; (3) temporary cooling during the Younger Dryas to ca 12 kya when ranges of warm-adapted species contracted and these species disappeared completely from Ireland; (4) further colonization of Britain from 11.5 kya to ca 8 kya by warm-adapted species arriving by the remaining land in the southern part of the North Sea before this was submerged by rising sea level; and, (5) deliberate and accidental introductions of mammal species by people arriving in Britain and Ireland in search of uninhabited land, trade or conquest. Deliberate introductions were most likely for economic reasons related to food supply and land management but some species have been transported for cultural reasons. Accidental introductions most likely reflect close association with livestock or, latterly, horticulture. These introductions occurred around five main periods during the Holocene; (a) with Mesolithic people, the first 156 W.I. Montgomery et al. / Quaternary Science Reviews 98 (2014) 144e165 kya Mus domesticus Nyctalus leisleri Cervus elaphus Mustela erminea Sorex minutus Myotis nattereri Myotis mystacinus Bos primigenius Microtus agrestis Mustela nivalis Sus scrofa Lepus timidus/hibernia Rhinolophus ferrumequinum Martes martes Myodes glareolus Vulpes vulpes Rhinolophus hipposideros 100 90 80 70 60 50 40 30 20 10 0 Mya Mus domesticus Sorex minutus Myotis nattereri Nyctalus leisleri Cervus elaphus Mustela erminea Myotis mystacinus Sus scrofa Mustela nivalis Bos primigenius Pipistrellus pygmaeus Microtus agrestis Rhinolophus ferrumequinum Pipistrellus pipistrellus Vulpes vulpes Martes martes Lepus timidus/hibernia Rhinolophus hipposideros Myodes glareolus Ursus arctos/maritimus Lepus europaeus Capreolus capreolus Microtus arvalis Sorex araneus Canis lupus Talpa europaea Apodemus flavicollis Apodemus sylvaticus Erinaceus europaeus 1.0 0.9 0.8 0.7 0.6 0.5 0.4 0.3 0.2 0.1 0.0 >1Mya >1Mya >1Mya >1Mya Fig. 5. a Times of divergence (95% bounds) of lineages in British and Irish species based on the molecular clock from 100kybp to present. Black indicates species of uncertain origin and shades of grey indicate lineages within species. Figure 5b Times of divergence (95% bounds) of lineages in British and Irish species based on the molecular clock from 1 Mybp to present. Shades of grey indicate lineages within species. colonizers; (b) with Neolithic, Bronze and Iron age people who brought agriculture, technology and built permanent buildings suitable for commensal species; (c) the 500 year period of Roman domination in Britain commencing just before 2 kya and forming part of an Empire with extensive trade focused in the south of Europe; (d) late 1st millennium and early 2nd millennium AD corresponding to Viking and Norman conquests; and, (e) late 2nd millennium and early 3rd millennium AD corresponding to a major increase in globalization of trade in natural resources, goods and services. (1) LGM to ca 15 kya. More southerly parts of Britain and subsequently Ireland, and cryptic refugia in the Bay of Biscay, Celtic Sea and western continental shelf, could have supported a sub-arctic or tundra mammal community comprising Norway and collared lemming, stoat, arctic fox, and reindeer. Brown bear could have survived close to open water and exploited seasonal berries, salmonid fish arriving to spawn and ground nesting birds but it has been argued that they would have been unable to sustain a viable population in Ireland through LGM (Leonard et al., 2013). Irish hare and wolf might have occurred along southern and western fringes where conditions were less harsh, the latter exploiting smaller mammals and following reindeer during seasonal migrations. It is probable that rapidly rising temperature around 22e20 kya made this ecosystem unstable due to repeated thaws and refreezing blocking access to fodder resulting in reduced population size in multiannual species and possible eventual extinction of Norway lemming and arctic fox in the west where melt-freeze events were more likely (Hof et al., 2012; Schmidt et al., 2012; Hansen et al., 2013). These conditions could also have made much of western Europe unsuitable for giant deer which was absent from Britain and Ireland until 15.8 kya and 14 kya respectively (Stuart et al., 2004). Red deer and perhaps pygmy shrew may have arrived in Britain and Ireland during the period of rapid warming after LGM, moving to more southerly refugia as the temperature dropped before rising again leading to the isolation of Ireland as sea level rose. (2) Warming 16 kyae14 kya. Cold adapted mammals in Ireland would have been particularly vulnerable to rising temperatures that followed the formation of the Irish Sea. Arctic fox may have been lost finally from Ireland at this time leaving wolf and bear as the main predators with reindeer, giant deer and perhaps red deer as larger prey, the latter if not already present entering across the remaining narrows and shallows of the Irish Sea. The 1800 year disparity in the earliest date for giant deer in Britain and Ireland and the isolation of the latter ca 15 kya, suggests that they did not simply recolonise from continental Europe to Britain to Ireland. The delay in recolonisation of Ireland may reflect the gradual reduction of suitable habitat adjacent to this island as sea level rose and expansion of suitable habitat on the island as temperature increased. Although there is nothing obvious in the spatial W.I. Montgomery et al. / Quaternary Science Reviews 98 (2014) 144e165 and temporal variability of giant deer morphology in Britain and Ireland (Lister, 1994), it is possible that like the stoat (Martinkova et al., 2007) and Irish hare (Hughes et al., 2009), giant deer recolonised Ireland and Britain independently. Hares and collared lemming would have sustained stoat in Ireland at this time whilst otter could have started to use more southerly coasts and rivers. In contrast, Britain had more northerly and higher altitudes than Ireland affording some refuge from rising temperatures. Root vole associated with wetter tundra conditions and still occurring in the Netherlands and steppe pika could have entered Britain at this time. Later, warm-adapted lineages and species entered Britain crossing Doggerland from adjacent areas of the continent. This early British, open ground small mammal fauna may have been supplemented by field vole and common shrew. (3) The Younger Dryas. This period of cooling immediately before the onset of the Holocene might have led to the final demise of the brown bear in Ireland, if still present, which without a link to continental Europe could not sustain a population of a large carnivore at a time of very low productivity. Similarly, red deer may have succumbed, leaving Ireland to a dwindling population of giant deer, reindeer and presumably wolf, Irish hare, collared lemming and stoat. Giant deer barely survived to the start of the Holocene in either Ireland or Britain, with latest material dating to 11.7 kya (Stuart et al., 2004). There is also evidence that in Ireland, giant deer as large, generalist herbivores would have been highly vulnerable to starvation as primary productivity decreased during the cooler years of the Younger Dryas (Chritz et al., 2009). Otter might also have contracted its range as a response to these colder conditions. Some southerly areas may have retained small populations of pygmy shrew at this time and the colder conditions in Britain might have delayed further encroachment of warm-adapted species. Absence of forest cover, in particular, would have limited Britain's mammal fauna in the Younger Dryas such that it might not have been that different from Ireland with some additional species and repletion from continental Europe. Hence, cold-adapted species such as both lemming species and arctic fox in Britain managed to survive to the start of the Holocene. (4) Holocene, 11.5 kya to ca 8 kya. Temperatures rose rapidly during the early Holocene. It is likely that these conditions led to the demise of the giant deer, reindeer and lemming species comprising the remains of the relatively coldadapted mammals of both Britain and Ireland. In Ireland, wolf may have clung on as the numbers of larger herbivores dwindled and subsisted thereafter on Irish hares, ground nesting birds and perhaps berries and fruits. Stoat presumably also depended on Irish hares whilst otters are fish dependent and could have expanded their range once again. Pygmy shrews in Ireland, if present at the start of the Holocene, would have colonized from one or more small areas to occupy the whole island, losing considerable genetic richness in the process. In contrast, Britain was still accessible to warm adapted species from the south such that once tree cover developed a succession of insectivores (mole, water shrew), rodents (wood mice, red squirrel), artiodactyls (roe deer) and carnivores (fox, pine marten, badger, weasel). Despite the simultaneous development of tree cover this did not happen in the early Holocene island of Ireland. Bats, notably common pipistrelle, Leislers' and Daubenton's, could get to Ireland and presumably did so exploiting suitable habitat and natural roost sites at much the same time as bats arrived in Britain. 157 (5) Species introductions. (a) Mesolithic. The first travellers to Britain and Ireland found very different landscapes and mammal fauna. It is probable that Ireland could only be accessed by estuaries, river valleys and sheltered coastal loughs due to a near complete cover of forest and offered limited species to hunt. People in rudimentary craft could travel between Britain and Ireland quickly and frequently and so could make return visits. Hence, introductions of species lacking in the latter, especially young, hand-reared individuals, could be made. Wild boar provided a source of food as well as a means of opening up forest whilst lynx and wildcat would provide fur either as clothing or comfort in shelters. Early travellers to Ireland may also have transported smaller mammals unwittingly: wood mice may survive for two days without food (Montgomery and Woods, 1986) and could have arrived amongst the baggage of early people in the bottom of a canoe, and pygmy shrew, if absent from Ireland during the early Holocene, must also have made this journey though as a very small insectivore needing to eat frequently to meet its metabolic needs (Meharg et al., 1990), its chance of successfully completing a boat trip of several hours must have been very small. In contrast, Britain had a wide selection of potential sources of food amongst its richer mammal fauna, and remained in direct contact to continental Europe which provided yet further resources. (b) Neolithic, and Bronze and Iron Ages. Red deer was probably absent from Ireland when people first arrived but would have been a mainstay of Mesolithic huntergatherers in Britain and continental Europe. Notwithstanding the problems of successfully transporting and introducing young deer, people would have been anxious to do this because of the food deer provide as well as skins, bone and antlers used in making tools. By Neolithic times forest was opening up and suitable habitat for red deer would have been widespread. Genetic and fossil evidence suggests that red deer reappeared in Ireland in the early Neolithic. Badger and red fox, and then pine marten appeared in Ireland during Bronze and Iron Ages, respectively, adding additional food resources and materials derived from fur and skins. Yellow-necked mice appear in Britain in the Neolithic whilst European hare and greater horseshoe bat appear in the late Iron Age. The former was almost certainly introduced for hunting but the latter could have been a natural range expansion. During the Iron Age, house mice appeared in both Britain and Ireland reflecting its commensalism and westwards migration with people, agriculture and more permanent buildings. (c) Roman Britain. The commensal rodent species, the black rat, arrived in both Britain and Ireland during the Roman occupation of the former. Considering the level of trade and human movements between Britain and southern Europe during this 500 year period, it is surprising that more mammal species were not introduced to Britain at this time. This may be due to the relatively rich mammal fauna already present in Britain such that there was limited human motivation as well as greater competition from indigenous species towards any accidental introductions. The Romans may have been instrumental in the translocation of novel genetic material rather than actually introducing species. However, it seems clear that the Romans introduced fallow deer into Britain ca. 2 kya 158 W.I. Montgomery et al. / Quaternary Science Reviews 98 (2014) 144e165 only for it to die out as Roman occupation came to an end (Sykes, 2010). (d) Viking voyages, Norman conquests. Similarly, the peregrinations of the Vikings along European coastlines did not result in any introductions of new species into either Britain or Ireland but probably did result in movement of genetic material amongst isolated populations of existing species. In contrast, the Normans created much more permanent settlements throughout both islands and, apparently, were responsible for the introduction of hedgehog, rabbit, red squirrel and fallow deer into Ireland, and rabbit and the return of the fallow deer into Britain. In this regard, the Normans were introducing their accustomed food species, culture and technology rather than individual mammal species. (e) Recent, 19the21st centuries. Only one species, Nathusius' pipistrelle bat, has arrived in Ireland and Britain by natural means in recent history (Lundy et al., 2010). In contrast, seven mammal species have been introduced accidentally or deliberately into Ireland, namely: European hare, grey squirrel, Sika deer, bank vole, American mink, greater white toothed shrew, and muntjac deer. Two species, roe deer and musk rat became established in particular places in Ireland but were removed by shooting and trapping, respectively (Fairley, 1984, 2001). Recently, the hazel dormouse has been confirmed as present in a small area (ca 30 km2) of Co.Kildare, Ireland, and has been estimated as present for at least 6 years (Sheehy and Lawton, in press) and roe deer were photographed in south Armagh (Catherine Bertrand, pers.comm.). Seven mammal species have also been introduced into Britain during this period, namely: grey squirrel, Sika deer, American mink, Chinese water deer, muntjac deer, edible dormouse and coypu which has been eradicated. As in Ireland, musk rat was established in Britain but trapped out. Red-necked wallaby was established in the Peak District but has also died out (Yalden, 2013). Thus, the rate of recent introductions into Ireland with viable populations, relative to number of mammal species already present, is much higher than that of Britain despite a much greater level of trade and human traffic in the latter. It is also noteworthy that the persisting, introduced mammal species new to Ireland include four from Europe and five orders, whilst those introduced into Britain originated from four continents and three orders, with only one species from Europe. Ireland continues to attract species introductions with a greater taxonomic diversity and has a greater dependency on European species as a source of introductions than Britain which has had an influx of species from further afield and introductions are more uniform taxonomically. 10. Discussion The critical periods in the history of British and Irish mammals are between the LGM and Ireland, then Britain, becoming islands, the climatic changes immediately preceding the Holocene, and the phases in human migration and trade. There is now a wellestablished, post-glacial record indicating that Ireland emerged from the ice sheets as an island no later than 15 kya making it approximately twice the age of Britain which lost contact with continental Europe with a further rise in sea level ca 8 kya. Between these two critical events, climate warmed possibly accounting for loss of cold-adapted mammal species and then cooled sharply during the Younger Dryas compromising warm-adapted species, before rising to the predominantly warmer, stable conditions of the Holocene ca 11.5 kya. Mammals in Ireland which initially had a more restricted land area and lower primary production, would have been more stressed by these contrary climatic events during the first few thousand years of the island's existence than the same species might have experienced in Britain. Thus, the latter not only emerged into the Holocene with a larger complement of species than Ireland, re-colonisation from the southwest and southeast of Europe continued for a further period of nearly 4000 years. It now seems highly likely that there was no Holocene land bridge between Britain and Ireland so that the timing of the sequence of events outlined by Mitchell and Ryan (1997) is no longer tenable. However, isolation of one more distant island and then another, more proximal island from their principal source of colonising species during global change in sea level effected by glaciation, is a global biogeographic pattern. This is the largest factor influencing biogeographical patterns of west Europe and its offshore islands during the Holocene. It is now evident how extreme this process has been in regard to the great disparity in the terrestrial mammals of Ireland in comparison to Britain and neighbouring areas of continental Europe. The fact that Ireland has been an island much longer than suspected, the relatively rapid oscillations in temperature leading to the Holocene, and perhaps the limited land area of Ireland, have all contributed to the its limited mammal species richness at the start of the Holocene. Paradoxically, these factors may also have contributed to the survival of early genotypes and greater genetic differentiation within Ireland such that its mammal fauna is unique at the community, species and intraspecific levels. Uncertainties in the physical history of Britain and Ireland after the LGM surround the location, extent and duration of land below current sea level around Ireland and Britain, and whether any of these areas were sufficiently free of both ice and sea to be refugia during southwards advances of the BIIS. ‘Cryptic’ refugia are now speculated on widely and have been proposed e.g. in northern part of Bay of Biscay and off the southern coast of Ireland. Such refugia may have been limited in both area and duration much influenced by local advances and retreats of ice. One approach has been to hindcast suitable habitat based on current species distributions and a suite of physical climatic data. Applying such a palaeodistribution modelling approach to mammals that occurred in Ireland by the middle part of the Holocene and hindcasting to 21 kya, suggests that stoat, red deer, Irish (mountain) hare, Leisler's bat and pygmy shrew all had suitable habitat to the west and south of Ireland as well as in the northern part of the Bay of Biscay, whereas badger, pine marten, brown bear, Natterer's bat and lynx lacked suitable habitat off the present day Irish coastline and had limited suitable habitat in the Bay of Biscay (Fig.6). Whilst there is no proof that either land to the west and south of Ireland or in the northern part of the Bay of Biscay acted as glacial refugium, these palaeoreconstructions of suitability, broadly complement genetic phylogeographic evidence regarding origin of genetic diversity in these species in Ireland e.g. in the stoat and hare, in comparison to the badger and pine marten, and is consistent with absences and reappearances in the Irish archaeological record e.g. in brown bear and lynx. Further, Beatty and Provan (2013) have demonstrated the importance of northern part of the Bay of Biscay in explaining the distribution of the Lusitanian plant species. Whilst more information has accrued in recent years, there remain uncertainties with respect to gaps in archaeological evidence. Material is generally sparse and past observer bias against smaller species, differential preservation, loss of context and possible incursions often confound interpretation. Irish archaeology is biased towards human settlements which seldom allow W.I. Montgomery et al. / Quaternary Science Reviews 98 (2014) 144e165 Mustela erminea Meles meles Cervus elaphus Lepus timidus Martes martes Ursus arctos Nyctalus leisleri Myotis nattereri 159 Sorex minutus Lynx lynx Fig. 6. Maximum Entropy models of distributions of selected species projected to 21kya indicating areas with suitable climatic conditions which could have acted as ‘cryptic’ refugia. reconstruction of animal communities or ecosystems. Ireland has a shortage of dry caves with undisturbed archaeology in comparison to Britain and those investigated in the past focussed on larger species. Archaeologically interesting British caves, in contrast, have yielded a wide range of smaller mammal species including bats. Such material could unleash the power of carbon dating to calibrate divergence times identified through molecular analyses. Similarly, molecular-genetic analyses are not a panacea. Genetic data can mislead e.g. nuclear DNA with copies of mitochondrial genes, and sampling is not always representative or sufficiently widespread to allow insights across the full range of a species. There is also a bias towards particular loci as well as extant material whilst ancient DNA which proves highly valuable in exploring phylogeographic patterns, is not always available or is scant. Methodological and observer biases in scoring sequences can confound even well designed studies. The full potential of the molecular clock is yet to be realised as it is very susceptible to variation in rates of mutation between loci, lineages and in deep versus shallow phylogeny. How useful molecular clock methods will be in discrimination of TMRCA during the Holocene or even historical times remains to be seen. Despite the shortcomings outlined above, the combined power of improved carbon dating and molecular genetics placed in the context of a better understanding of the timing of climatic and environmental changes during the late Quaternary and Holocene has led to improved understanding and more certainty of the postglacial history of the mammals of Britain and Ireland summarised in Fig.7. Conclusions arrived at through the more conventional approach of archaeology and palaeoecology on the one hand, and moleculargenetic phylogeography on the other, are mostly congruent, but their true value is most evident where archaeological and genetic approaches are combined and phylogeographic inferences are based on data from both extant and ancient DNA and modern dating procedures. Genetic analyses have revealed some unique genetic lineages amongst British and Irish mammals. Long regarded as an example of species ‘impoverishment’, Ireland, contributes to the overall genetic diversity of European mammals. Indeed, some of these unique forms may be invaluable in adaptation to rapidly changing environmental conditions. Phylogeographical data have confirmed that the absence of mammal species from Ireland reflects its earlier isolation and that people were directly or indirectly instrumental in the introduction of other mammal species from their first arrival. Similarly, biogeographic and phylogeographic analyses suggest that Britain is not that different with respect to the history of its mammals from neighbouring, lowland, continental northwest Europe. With the exception of the Romans in Britain, successive waves of human migrants and trade introduced species deliberately or accidentally to Ireland from Britain and directly from parts of continental Europe. More recently, exotic mammal species have been introduced to both islands from much further afield. Alien, invasive species (AIS) have accumulated globally and in both Britain and Ireland more rapidly over the last 100 years than the last 10,000 years. The impact of AIS is likely to be proportionately greater in Ireland as a smaller island with a more limited indigenous mammal fauna. Whilst data on the impact of AIS are scarce several species are believed to have significant ecological effects, namely: mink, grey squirrel, sika deer, muntjac, and bank vole and greater white toothed shrew in either Britain or Ireland or both. Hybridisation involving AIS has been confirmed in sika and red deer and brown and Irish hare in Ireland. Hence, the revelation of the unique genetic diversity of the mammals of Britain and Ireland, is offset by the major threat to their survival due to AIS. Management of AIS has generally taken the form of a species by species approach and often involves no action. This approach needs urgent re-evaluation, particularly, in Ireland. The origin of British and Irish mammals should also inform conservation policies to prioritise species that show unique genetic variation reflecting survival through LGM or arrival with the early people of these islands. 160 W.I. Montgomery et al. / Quaternary Science Reviews 98 (2014) 144e165 Fig. 7. Map illustrating the major physical challenges afecting mammal recolonisation of northern and western Europe. Red circles indicate location of accepted refugia during LGM: 1. Iberia. 2. Northern Italy. 3. Balkans. 4. Carpathia. White circles indicate inferred or ‘cryptic’ refugia on the eastern Atlantic shelf: A. Shannon sea area off southwest Ireland; B. Fastnet sea area, Celtic Sea. C. Armoricain Shelf, Bay of Biscay. Major barriers to dispersal are indicated: mountain ranges in yellow, rivers in light blue, coastal seas in dark blue. Large blue arrows indicate putative routes of reinvasion from Britain to Ireland prior to 15kya, and continental Europe to Britain prior to 7.8 kya. 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