Quaternary Science Reviews 98 (2014) 144e165
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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.
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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
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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. Smaller blue arrows indicate putative
dispersal from cryptic refugia. White arrows indicate main routes of recolonisation from western, southern and eastern continental Europe.
Acknowledegments
We are grateful to many scientists who have made major contributions to the study of the physiography, archaeology, history
and phylogeography of British and Irish mammals. This paper is
dedicated to the memory of Derek Yalden.
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