Origin of British and Irish mammals: disparate post-glacial colonisation and species introductions

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 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.

The chronostratigraphy of the Haua Fteah cave (Cyrenaica, northeast Libya)

The 1950s excavations by Charles McBurney in the Haua Fteah, a large karstic cave on the coast of northeast Libya, revealed a deep sequence of human occupation. Most subsequent research on North African prehistory refers to his discoveries and interpretations, but the chronology of its archaeological and geological sequences has been based on very early age determinations. This paper reports on the initial results of a comprehensive multi-method dating program undertaken as part of new work at the site, involving radiocarbon dating of charcoal, land snails and marine shell, cryptotephra investigations, optically stimulated luminescence (OSL) dating of sediments, and electron spin resonance (ESR) dating of tooth enamel. The dating samples were collected from the newly exposed and cleaned faces of the upper 7.5m of the ~14.0m-deep McBurney trench, which contain six of the seven major cultural phases that he identified. Despite problems of sediment transport and reworking, using a Bayesian statistical model the new dating program establishes a robust framework for the five major lithostratigraphic units identified in the stratigraphic succession, and for the major cultural units. The age of two anatomically modern human mandibles found by McBurney in Layer XXXIII near the base of his Levalloiso-Mousterian phase can now be estimated to between 73 and 65ka (thousands of years ago) at the 95.4% confidence level, within Marine Isotope Stage (MIS) 4. McBurney's Layer XXV, associated with Upper Palaeolithic Dabban blade industries, has a clear stratigraphic relationship with Campanian Ignimbrite tephra. Microlithic Oranian technologies developed following the climax of the Last Glacial Maximum and the more microlithic Capsian in the Younger Dryas. Neolithic pottery and perhaps domestic livestock were used in the cave from the mid Holocene but there is no certain evidence for plant cultivation until the Graeco-Roman period. © 2013 Elsevier Ltd.

New age estimates and climatostratigraphic correlations for the Borrobol and Penifiler Tephras: evidence from Abernethy Forest, Scotland

The emerging tephrostratigraphy of NW Europe spanning the last termination (ca. 15–9 ka) provides the potential for synchronizing marine, ice-core and terrestrial records, but is currently compromised by stratigraphic complications, geochemical ambiguity and imprecise age estimates for some layers. Here we present new tephrostratigraphic, radiocarbon and chironomid-based
palaeotemperature data from Abernethy Forest, Scotland, that refine the ages and stratigraphic positions of the Borrobol and Penifiler tephras. The Borrobol Tephra (14.14–13.95 cal ka BP) was deposited in a relatively warm period equated with Greenland Interstadial sub-stage GI-1e. The younger Penifiler Tephra (14.09–13.65 cal ka BP) is closely associated with a cold oscillation equated with GI-
1d. We also present evidence for a previously undescribed tephra layer that has a major-element chemical signature identical to the Vedde Ash. It is associated with the warming trend at the end of the Younger Dryas, and dates between 11.79 and 11.20 cal ka BP.

Detection of Lateglacial distal tephra layers in the Netherlands

Three distal tephra layers or cryptotephras have been detected within a sedimentary sequence from the Netherlands that spans the last glacial-interglacial transition. Geochemical analyses identify one as the Vedde Ash, which represents the southernmost discovery of this mid-Younger Dryas tephra so far. This tephra was found as a distinct horizon in three different cores sampled within the basin. The remaining two tephras have not been geochemically ‘fingerprinted’, partly due to low concentrations and uneven distributions of shards within the sequences sampled. Nevertheless, there is the potential for tracing these tephra layers throughout the Netherlands and into other parts of continental Europe. Accordingly, the possibilities for precise correlation of Dutch palaeoenvironmental records with other continental, marine and ice-core records from the North Atlantic region are highlighted.

Evolving southwest African response to abrupt deglacial North Atlantic climate change events

Climate change during the last deglaciation was strongly influenced by the „bipolar seesaw‟, producing antiphase climate responses between the North and South Atlantic. However, mounting evidence demands refinements of this model, with the occurrence of abrupt events in southern low to mid latitudes occurring in-phase with North Atlantic climate. Improved constraints on the north-south phasing and spatial extent of these events are therefore critical to
understanding the mechanisms that propagate abrupt events within the climate system. We present a 19,400 year multi-proxy record of climate change obtained from a rock hyrax midden in southernmost Africa. Arid anomalies in phase with the Younger Dryas and 8.2 ka events are apparent, indicating a clear shift in the influence of the bipolar seesaw, which diminished as the Earth warmed, and was succeeded after ~14.6 ka by the emergence of a dominant interhemispheric atmospheric teleconnection.

A multi-proxy chronological reconstruction of palaeosalinity in the eastern Limfjord, Denmark

The Jutland peninsula in northern Denmark is home to the Limfjord, one of the largest estuarine bodies of water in the region. Human inhabitance of the Limfjord’s surrounding coastlines stretches back further than 7,800 cal BP, with anthropogenic influence on the landscape beginning approximately 6,000 cal BP. Understanding how the Limfjord as a system has changed throughout time is useful in comprehending subsistence patterns and anthropogenic influence. This research is part of a larger project aimed at discerning subsistence patterns and environmental change in the region. Following the Younger Dryas, as the Fennoscandian ice sheet began to melt, Denmark experienced isostatic rebound, which contributed to the complex sea level history in the region. Between ice melt and isostatic rebound, the Jutland peninsula experienced many transgression and regression events. Connections to surrounding seas have shifted throughout time, with most attention focused on the western connection of the Limfjord with the North Sea, which has experienced numerous closures and subsequent re-openings throughout the Holocene. Furthermore, the Limfjord-North Sea connection has been the focal point of research because of the west to east water flow in the system, and the present day higher salinity in the west compared to the east. Little to no consideration has been paid to the influence of the Kattegat and Baltic on the Limfjord until now. A 10m sediment core was taken from Sebbersund (near Nibe, Limfjord), along the connection between the Limfjord and the Kattegat in the east to understand how the eastern part of the system has changed and differed from changes observed in the west. The Sebbersund sequence spans a majority of the Holocene, from 9600 cal BP to 1030 cal BP, determined via radiocarbon dating of terrestrial macrofossils and bulk sediment. Over this time period palaeoenvironmental conditions were reconstructed through the use of geochemical analyses (13C, 15N, C:N), physical sediment analyses, dinoflagellate cyst abundances and molluscan analyses. apart from two instances of low salinity, one at the top and one at the bottom of the core, the sequence has a strong marine signal for a majority of the Holocene. Radiocarbon dating of bulk sediment samples showed the presence of old carbon in the system, creating an age offset between 1,300 ± 200 and 2,800 ± 200 calibrated 14C years compared to the age-depth curve based on the terrestrial macrofossils. This finding, along with the strong marine influence in the system, discerned through geochemical data, dinoflagellate cyst and mollusc counts, is important for obtaining accurate radiocarbon ages in the region and stresses the importance of understanding both the marine and freshwater reservoir effects. The marine dominance in the eastern Limfjord differs from the west, which is characterized by a number of freshwater events when the North Sea connection was closed off during the Holocene. The eastern connection was open to the Kattegat throughout a large portion of the Holocene, with influx of open ocean water entering the system during periods of higher sea level.

Late-Glacial and early Holocene marine 14C reservoir ages off North Iceland (16-9 cal kyr BP)

Tephrochronological age models and 48 14C age determinations on molluscs and foraminifera (planktonic and benthic) are applied for the calculation of marine 14C reservoir age variability during a time period covering the Heinrich event H1 to early Holocene (16–9 cal kyr BP). Our data source consists of four high-resolution marine sediment cores (HM107-04, HM107-05, MD99-2271, MD99-2275) from the North Icelandic shelf. The marine reservoir age (ΔR) is found to be extremely variable, ranging from 385 to 1065 14C years. Extreme ΔR values occur at the end of H1, with values around 1000 14C years (~15 cal kyr BP), probably due to reduced northward flow of well-ventilated subtropical surface waters and a southward expansion of polar waters, as well as an expansion of sea ice limiting air-sea gas exchange. With the onset of the Bølling-Allerød interstadial, the ΔR values decrease towards 0 14C years suggesting a more vigorous North Atlantic Current and an active meridional overturning circulation system. During the Younger Dryas stadial, ΔR values are consistently around 700 14C years suggesting e renewed expansion of polar waters and a weakened meridional overtuning circulation. Interestingly, ΔR values remain high (~200 14C years) at the onset of the Holocene suggesting continued high influence of polar waters. Subsequently, ΔR values rapidly decrease to ~¬ 250 14C years around 11 cal kyr BP, indicating increased air-sea CO2 exchange with the coeval atmosphere. The ΔR values average around 0 14C years from around 10.5 to 9.0 cal kyr BP.

Interpreting the response of a dryland river system to Late Quaternary climate change

A U-series calcrete chronology has been constructed for three Late Quaternary terrace units, termed the D1, D2 and D3 terraces in age descending order, from the Rio Aguas river system of the Sorbas basin, southeast Spain. The D1 terrace formed between 30,300 +/- 4400 year BP and 12,140 +/- 360 year BP, correlating well with the Last Glacial Maximum when rates of sediment supply would have increased greatly, because of higher rates of weathering, reduced vegetation cover and weak soil development. The D2 terrace formed between 12,800 +/- 1100 year BP and 9,600 +/- 530 year BP, correlating well with the Younger Dryas event. The D3 terrace could only be poorly constrained to the early Holocene and no unequivocal cause could be assigned to this period of aggradation. The sedimentology and geomorphology of the D2 terrace suggests, however, that the aggradation of this unit was a response to diapirism/karstic processes occurring within the underlying Messinian gypsum strata and the subsequent damming of the Aguas system. Therefore, despite its coincident occurrence with the Younger Dryas, aggradation of the D2 terrace is unrelated to climate change. The style of this response, controlled predominantly by the characteristics of the underlying bedrock, makes correlating the terrace record of the Aguas with other systems in the Mediterranean unreliable. This study, therefore, highlights the problems of correlating fluvial sequences in regions of variable tectonics, climatic history and bedrock geology and emphasises the need to properly understand the main controls on individual fluvial systems before any attempt is made to correlate their depositional histories. (C) 2004 Elsevier Ltd. All rights reserved.

Vegetation development and human occupation in the Damascus region of southwestern Syria from the Late Pleistocene to Holocene

Rockshelter Baaz in the Damascus region of Syria provided a variety of botanical remains from the Late Pleistocene and Early Holocene period. These remains provide new information about the vegetation evolution in this region. The earliest occupational levels correspond with a moisture peak during the Late Pleistocene, between ca. 34-32 kyr b.p., when pine expanded. The next occupations took place during extreme arid conditions, ca. 23-21 kyr b.p., and probably during the Last Glacial Maximum when a steppe vegetation was established. The occupation level of the Younger Dryas, represented by Natufian remains, suggests that the area had been covered by almond-pistachio steppe, similar to later periods of the Early Holocene, and was probably located just outside the range of dense wild cereal stands. There is no drastic impact of the Younger Dryas visible on the vegetation in the botanical remains. The lack of fruits and seeds at Baaz indicates that the site was more likely to have been a temporary hunting post rather than a plant processing site for much of its history. It is ideally suited to this purpose because of its location over the Jaba'deen Pass and the associated springs. However, archaeological remains from the Natufian period, suggest that the site was more permanently occupied during this time.

The sea-level conundrum: case studies from palaeo-archives

Uncertainties in sea-level projections for the 21st century have focused ice sheet modelling efforts to include the processes that are thought to be contributing to the recently observed rapid changes at ice sheet margins. This effort is still in its infancy, however, leaving us unable to make reliable predictions of ice sheet responses to a warming climate if such glacier accelerations were to increase in size and frequency. The geological record, however, has long identified examples of nonlinear ice sheet response to climate forcing (Shackleton NJ, Opdyke ND. 1973. Oxygen isotope and paleomagnetic stratigraphy of equatorial Pacific core V28–239, late Pliocene to latest Pleistocene. Geological Society of America Memoirs145: 449–464; Fairbanks RG. 1989. A 17,000 year glacio-eustatic sea level record: influence of glacial melting rates on the Younger Dryas event and deep ocean circulation. Nature342: 637–642; Bard E, Hamelin B, Arnold M, Montaggioni L, Cabioch G, Faure G, Rougerie F. 1996. Sea level record from Tahiti corals and the timing of deglacial meltwater discharge. Nature382: 241–244), thus suggesting an alternative strategy for constraining the rate and magnitude of sea-level change that we might expect by the end of this century. Copyright © 2009 John Wiley & Sons, Ltd.

Wildfire responses to abrupt climate change in North America

It is widely accepted, based on data from the last few decades and on model simulations, that anthropogenic climate change will cause increased fire activity. However, less attention has been paid to the relationship between abrupt climate changes and heightened fire activity in the paleorecord. We use 35 charcoal and pollen records to assess how fire regimes in North America changed during the last glacial–interglacial transition (15 to 10 ka), a time of large and rapid climate changes. We also test the hypothesis that a comet impact initiated continental-scale wildfires at 12.9 ka; the data do not support this idea, nor are continent-wide fires indicated at any time during deglaciation. There are, however, clear links between large climate changes and fire activity. Biomass burning gradually increased from the glacial period to the beginning of the Younger Dryas. Although there are changes in biomass burning during the Younger Dryas, there is no systematic trend. There is a further increase in biomass burning after the Younger Dryas. Intervals of rapid climate change at 13.9, 13.2, and 11.7 ka are marked by large increases in fire activity. The timing of changes in fire is not coincident with changes in human population density or the timing of the extinction of the megafauna. Although these factors could have contributed to fire-regime changes at individual sites or at specific times, the charcoal data indicate an important role for climate, and particularly rapid climate change, in determining broad-scale levels of fire activity.

Vegetation and environmental dynamics in the southern Black Sea region since 18 kyr BP derived from the marine core 22-GC3

Sediments from the Black Sea, a region historically dominated by forests and steppe landscapes, are a valuable source of detailed information on the changes in regional terrestrial and aquatic environments at decadal to millennial scales. Here we present multi-proxy environmental records (pollen, dinoflagellate cysts, Ca, Ti and oxygen isotope data) from the uppermost 305 cm of the core 22-GC3 (42°13.53′N, 36°29.55′E) collected from a water depth of 838 m in the southern part of the Black Sea in 2007. The records span the last ~ 18 kyr (all ages are given in cal kyr BP). The pollen data reveal the dominance of the Artemisia-steppe in the region, suggesting rather dry/cold environments ~ 18–14.5 kyr BP. Warming/humidity increase during melt-water pulses (~ 16.1–14.5 kyr BP), indicated by δ18O records from the 22-GC3 core sediment and from the Sofular Cave stalagmite, is expressed in more negative δ13C values from the Sofular Cave, usually interpreted as the spreading of C3 plants. The records representing the interstadial complex (~ 14.5–12.9 kyr BP) show an increase in temperature and moisture, indicated by forest development, increased primary productivity and reduced surface run-off, whereas the switch from primary terrigenous to primary authigenic Ca origin occurs ~ 500 yr later. The Younger Dryas cooling is clearly demonstrated by more negative δ13C values from the Sofular Cave and a reduction of pines. The early Holocene (11.7–8.5 kyr BP) interval reveals relatively dry conditions compared to the mostly moist and warm middle Holocene (8.5–5 kyr BP), which is characterized by the establishment of the species-rich warm mixed and temperate deciduous forests in the low elevation belt, temperate deciduous beech-hornbeam forests in the middle and cool conifer forest in upper mountain belt. The border between the early and middle Holocene in the vegetation records coincides with the opening of the Mediterranean corridor at ~ 8.3 kyr BP, as indicated by a marked change in the dinocyst assemblages and in the sediment lithology. Changes in the pollen assemblages indicate a reduction in forest cover after ~ 5 kyr BP, which was likely caused by increased anthropogenic pressure on the regional vegetation.

The coupled δ13C-radiocarbon systematics of three Late Glacial/early Holocene speleothems: insights into soil and caveprocesses at climatic transitions

The coupled δ13C-radiocarbon systematics of threeEuropean stalagmites deposited during the Late Glacial and early Holocene were investigated to understand better how the carbon isotope systematics of speleothems respond to climate transitions. The emphasis is on understanding how speleothems may record climate-driven changes in the proportions of biogenic (soil carbon) and limestone bedrock derived carbon. At two of the three sites, the combined δ13C and 14C data argue against greater inputs of limestone carbon as the sole cause of the observed shift to higher δ13C during the cold Younger Dryas. In these stalagmites (GAR-01 from La Garma cave, N. Spain and So-1 from Sofular cave, Turkey), the combined changes in δ13C and initial 14C activities suggest enhanced decomposition of old stored, more recalcitrant, soil carbon at the onset of the warmer early Holocene. Alternative explanations involving gradual temporal changes between open- and closed-system behaviour during the Late Glacial are difficult to reconcile with observed changes in speleothem δ13C and the growth rates. In contrast, a stalagmite from Pindal cave (N. Spain) indicates an abrupt change in carbon inputs linked to local hydrological and disequilibrium isotope fractionation effects, rather than climate change. For the first time, it is shown that while the initial 14C activities of all three stalagmites broadly follow the contemporaneous atmospheric 14C trends (the Younger Dryas atmospheric 14C anomaly can be clearly discerned), subtle changes in speleothem initial 14C activities are linked to climate-driven changes in soil carbon turnover at a climate transition.

Fire history and the Global Charcoal Database: a new tool for hypothesis testing and data exploration

Version 1 of the Global Charcoal Database is now available for regional fire history reconstructions, data exploration, hypothesis testing, and evaluation of coupled climate–vegetation–fire model simulations. The charcoal database contains over 400 radiocarbon-dated records that document changes in charcoal abundance during the Late Quaternary. The aim of this public database is to stimulate cross-disciplinary research in fire sciences targeted at an increased understanding of the controls and impacts of natural and anthropogenic fire regimes on centennial-to-orbital timescales. We describe here the data standardization techniques for comparing multiple types of sedimentary charcoal records. Version 1 of the Global Charcoal Database has been used to characterize global and regional patterns in fire activity since the last glacial maximum. Recent studies using the charcoal database have explored the relation between climate and fire during periods of rapid climate change, including evidence of fire activity during the Younger Dryas Chronozone, and during the past two millennia.

Deglacial changes of the southern margin of the southern westerly winds revealed by terrestrial records from SW Patagonia (52 degrees S)

Much of the ongoing discussion regarding synchrony or bipolar asynchrony of paleoclimate events has centered on the timing and structure of the last glacial termination in the southern mid- latitudes, in particular the southwestern Patagonian region (50�e55�S). Its location adjacent to the Drake Passage andnear the southern margin of the southern westerly winds (SWW) allows examining the postulated links between the Southern Oceane SWW coupled system and tmospheric CO2 variations through the last glacial termination. Results from two sites located in the Última Esperanza area (52�S) allow us to infer SWW-driven changes in hydrologic balance during this critical time interval. These findings indicate peatland development under temperate/wet conditions between 14,600 and 14,900 cal yr BP, followed by cooling and a lake transgressive phase that led to a shallow lake during the early part of the Antarctic Cold Reversal (ACR, 13,600-14,600 cal yr BP), followed in turn by a deeper lake and modest warming during Younger Dryas time (YD, 11,800-13,000 cal yr BP), superseded by terrestrialization and forest expansion at the beginning of the Holocene. We propose that the SWW (i) strengthened and shifted northward during ACR time causing a precipitation rise in northwestern and southwestern Patagonia coeval with mid- and high-latitude cooling and a halt in the deglacial atmospheric CO2 rise; (ii) shifted southward during YD time causing a precipitation decline/increase in NW/SW Patagonia, respectively, high-latitude warming, and invigorated CO2 release from the Southern Ocean; (iii) became weaker between 10,000 and 11,500 cal yr BP causing a precipitation decline throughout Patagonia, concurrent with peak mid- and high-latitude temperatures and atmospheric CO2 concentrations.