ARTICLE IN PRESS

Quaternary Science Reviews 25 (2006) 1755–1789

The Holocene British and Irish ancient forest fossil fauna: implications for forest history, and faunal colonisation

Nicki J. Whitehouse

Palaeoecology Centre, School of Archaeology and Palaeoecology, Queen’s University Belfast, Belfast BT7 1NN, Northern Ireland, UK

Received 15 June 2004; accepted 10 August 2005

Abstract

This paper presents a new review of our knowledge of the ancient forest beetle fauna from Holocene archaeological and palaeoecological sites in Great Britain and Ireland. It examines the colonisation, dispersal and decline of beetle species, highlighting the scale and nature of human activities in the shaping of the landscape of the British Isles. In particular, the paper discusses effects upon the fauna, and examines in detail the fossil record from the Humberhead Levels, eastern England. It discusses the local extirpation of up to 40 species in Britain and 15 species in Ireland. An evaluation of the timing of extirpations is made, suggesting that many species in Britain disappear from the fossil record between ca 3000 and 1000 cal BC (ca 5000–3000 cal BP), although some taxa may well have survived until considerably later. In Ireland, there are two distinct trends, with a group of species which seem to be absent after ca 2000 cal BC (ca 4000 cal BP) and a further group which survives until at least as late as the medieval period. The final clearance of the Irish landscape over the last few hundred years was so dramatic, however, that some species which are not especially unusual in a British context were decimated. Reasons behind the extirpation of taxa are examined in detail, and include a combination of forest clearance and human activities, isolation of populations, lack of temporal continuity of habitats, edaphic and competition factors affecting distribution of host trees (particularly ), lack of forest fires and a decline in open forest systems. The role of climate change in extirpations is also evaluated. Consideration is given to the significance of these specialised ancient forest inhabitants in Ireland in the absence of an early Holocene land-bridge which suggests that colonisation was aided by other mechanisms, such as human activities and wood rafting. Finally, the paper discusses the Continental origins of the British and Irish fauna and its hosts and the role played by European glacial refugia. r 2006 Elsevier Ltd. All rights reserved.

1. Introduction seen perhaps most dramatically in areas such as New Zealand, where the environmental impact of settlement It is observable that not only the moose and the wolf caused faunal extirpation, dramatic changes in erosion disappear before the civilised man, but even many rates, sedimentation and deforestation (e.g. McGlone, species of , such as the black fly and the almost 1983; Martin, 1984; McGlone and Wilmshurst, 1999). In microscopic ‘‘no-see-em’’. How imperfect a notion have contrast, the European lands from which many of the we commonly of what was the actual condition of the settlers originated had already been significantly modified place where we dwell, three centuries ago! Henry David and shaped by millennia of cultural activities, but where Thoreau, January 29, 1856 (Homan, 1991, p. 132). the effects of human impact had had no less of a dramatic Henry Thoreau’s comments relate to some of the impact—albeit over a more protracted period of time. enormous changes to the American flora and fauna which The study of sub-fossil has highlighted the scale arose within a few centuries of the arrival of Europeans. of these environmental impacts in , particularly The palaeoecological record covering this period highlights over the last 5000 years of the Holocene (Buckland and the dramatic environmental impact of such settlers (e.g. Coope, 1991). Transitions from forested to cleared land- Baker et al., 1996; Schwert, 1996; Bradbury et al., 2004), scape (termed ‘‘culture steppe’’ by Hammond (1974)) have been observed from several sites (e.g. Girling, 1985, 1989; E-mail address: [email protected]. Dinnin, 1997). Much of this record has been obtained via

0277-3791/$ - see front matter r 2006 Elsevier Ltd. All rights reserved. doi:10.1016/j.quascirev.2006.01.010 ARTICLE IN PRESS 1756 N.J. Whitehouse / Quaternary Science Reviews 25 (2006) 1755–1789 fossil beetles from archaeological investigations, from both geographical focus of the paper is Great Britain and rural and urban archaeological sites (e.g. Girling, 1976, Ireland, where there has been a long tradition of 1979; Hall and Kenward, 1980; Robinson, 1991, 2000; palaeoentomology (particularly in the former rather than Kenward and Hall, 1995; Hall et al., 2003). There have also the latter) and a correspondingly good data set, although been palaeoecological studies which have examined ‘‘nat- reference is also made to research carried out elsewhere. ural’’ deposits, indirectly associated with archaeological Figs. 1 and 2 show the location of sites discussed. features, but which carry a record of human impact on the Coleoptera nomenclature follows Lucht (1987), whilst landscape. The anaerobic conditions ensure excellent plant nomenclature follows Stace (1991). Information preservation of fossils whilst the often rapid build-up of on insect biology has been obtained from the entomo- deposits provide good temporal resolution. Floodplain logical database BUGS (Buckland et al., 1997, www. deposits, for instance, are immensely valuable, as they bugs2000.org), whilst distributional information has been provide records of fluvial histories, floodplain evolution largely obtained by a search through specialised catalogues and change, vegetation composition and structure as well housed in the Royal Entomological Society’s library, as the nature and scale of human impact (e.g. Osborne, London. Where necessary, reference is made to the current 1988, 1995; Dinnin, 1997; Andrieu-Ponel et al., 2000; threatened status of some of the beetles discussed. These Plunkett et al., 2003; Smith and Howard, 2004). Bogs and are species which are included in the Nature Conservancy fens are also rich sources of fossil insect assemblages, Council’s British Red Data Books: 2, Insects (Shirt, 1987). although fen peats tend to be richer in insect remains than This is a comprehensive statement on the status of most acid peats (e.g. Buckland, 1979; Klinger et al., 1990; Elias, threatened insects in Great Britain. The book contains 1994; Roper, 1996; Whitehouse, 1997a, 2004; Ponel et al., three major categories based upon degrees of threat, RDB 2001; Marra, 2003). Results of such investigations highlight (Red Data Book) 1, 2 and 3 (respectively, endangered the enormous changes effected upon the British and Irish vulnerable and rare). A further classification of Notable faunas, particularly since the onset of agriculture. More- species (A and B) (Hyman, 1992, 1994) is also used. over, they also provide an insight into the rapid colonisa- Notable A species are those which do not fall within RDB tion and expansion of these islands by insects as the ice categories but which are none-the-less uncommon and are sheets retreated. thought to occur in 30 or fewer 10 km2 of the National This paper presents a new contribution towards our Grid, whilst Notable B taxa are those which are thought to understanding of the colonisation, dispersal and decline of insect species and what this can tell us about the nature and role of human activities in the shaping of the European landscape and their effects upon the insect fauna. In particular, the contraction of old forest communities and species (Urwaldtiere) is examined in detail. This contribu- tion augments discussions originally highlighted by Os- borne (1965, 1997), Buckland and Kenward (1973), Buckland (1979), Girling (1982, 1985), Whitehouse (1997a, b), Dinnin and Sadler (1999) and Smith and Gransmoor Humberhead Levels Whitehouse (2005), and brings together information Wood Hall published in a wide variety of sources which will be of Chelford Dimlington interest not only to palaeoecologists but also researchers Bole Ings interested in Holocene forest history and ecology. It differs from previous accounts by considering previously pub- Rodbaston Wilsford lished (Coope et al., 1979; Reilly, 1996, 2003; Caseldine et West Bromich Lea Marston al., 2001; Whitehouse, in press) and unpublished Irish research (e.g. Rogers, 2004). It also discusses in detail the Upton Warren biology of saproxylic species (species where ‘‘the immature Dorney stages develop in some part of the wood-decay succession Runnymede or on the products of it’’, Alexander, 2002, p. 10) and Sweet Track Hampstead causes for the dramatic changes seen amongst the fauna, Heath including climatic factors and their wider significance. It Hollywell Coobe also provides distributional and habitat information of individual species which have not previously been pub- lished and adds to the last major review of these species 0 100 200 km undertaken by Buckland (1979). Finally, the paper addresses issues concerned with the mechanisms behind the initial colonisation of insect biota, particularly to Fig. 1. Map of England and Wales, showing location of sites discussed in Ireland, and considers the Continental origins of taxa. The the text. ARTICLE IN PRESS N.J. Whitehouse / Quaternary Science Reviews 25 (2006) 1755–1789 1757

appropriate to report these as cal BC dates in the same manner.

Deer Park Farms 2. The ancient forest fossil beetle record Sluggan Bog Ballymacombs Belfast More 2.1. The development of Urwald and the expansion of the Strangford beetle fauna Aghnadarragh Lough The Lateglacial period (ca 13,000–10,000 BP) is impor- Drumurcher tant in any consideration of the beetle fauna of the present interglacial. Hammond (1974) suggests that as much as half Corlea of the present-day fauna had already established itself Dublin before major afforestation. Investigations by Coope (1979) indicate that thermophilous species arrived extremely rapidly during the early phases of warming. Southern Derryville parts of Britain acted as a glacial refuge for at least some cold-adapted species since these areas were not covered by ice sheets. Southern and westerly areas of Ireland are likely Shortalstown to have similarly acted as refugia (Praeger, 1932). This story may be seen in the arrival of thermophilous taxa as 0 100 200 km they re-expanded during periods of climatic amelioration (Interstadials) during the last glaciation (Devensian in Britain and Midlandian in Ireland), such as assemblages from Chelford, Cheshire (Coope, 1959), Upton Warren, Fig. 2. Map of Ireland, showing location of sites discussed in the text. Worcestershire (Coope et al., 1961), and Aghnadarragh, Co. Antrim (McCabe et al., 1987). In contrast, faunas from occur between 31 and 100 10 km2 of the National Grid. Dimlington, Yorkshire, provide an insight into the very There is no equivalent Red Data Book for Irish species so limited range of cold-adapted taxa present during the the British version is applied to species, whilst acknowl- coldest parts of the last glaciation (Penny et al., 1969). edging that their status in Ireland may be different. One The same pattern of expansion and contraction is seen point worth remembering is that species that are rare today during the Lateglacial, but is often observable in far greater may not have been so in the past, and perhaps vice versa detail and at a greater resolution than is possible with (Buckland and Dinnin, 1993). Species which are today earlier deposits. Coope (1979) highlights the waves of cold- ‘‘locally extirpated’’ from Britain are identified by * in adapted taxa, followed by warmth-loving species during front of the species name, whilst species which are the early part of the Interstadial (13,200–12,200 BP) extirpated in Ireland are identified by the prefix ].Insome followed by cooler taxa during the latter parts of the cases, we know that taxa are extirpated from both Ireland Interstadial (12,200–11,000 BP), with cold taxa re-invading and Britain (such taxa are prefixed by m). However, the during the Younger Dryas interval (11,000–10,000 BP). limited number of investigations in Ireland means it is This pattern is repeated at numerous sites across the British often hard to assume a species that is present in the British Isles. In some southerly locations, an insect fauna fossil record was present in Ireland in the past and vice associated with light forest had already started to colonise versa, so the absence of this symbol should not be some locations shortly after 13,000 BP, with the arrival of extrapolated in any meaningful way. the (scolytid), ratzeburgi at Holocene dates are expressed as BC/AD for dendro- Holywell Coombe, on the south-east coast of England chronological dates (followed by the appropriate biblio- (Coope, 1998). Its presence along with other species graphic citation) and cal BC/AD for radiocarbon dates associated with birch, and other trees through the (range calculated to 95% level confidence and calibrated Interstadial indicate suitable climatic conditions for light using INTCAL 1986 calibration curve (Pearson et al., forest. They apparently did not survive the cold of the 1986), following recent recommendations by Reimer et al. Younger Dryas, however, retreating southwards. (2002). The use of precise calendrical dendrochronological Despite the wealth of Lateglacial fossil insect investiga- chronologies in the paper precludes the use of radiocarbon tions carried out over the last 50 years or so, remarkably dates (BP) dates throughout, a preferable option for many little is known about the early Holocene fossil insect readers, particularly those without an archaeological record, a period which would clearly benefit from further background. However, where appropriate, calibrated BP investigation (Whitehouse and Smith, 2004). During the dates are also included in brackets. Where reference is early phases of the present interglacial, rapid climatic made to Lateglacial and earlier records, uncalibrated warming enabled tree taxa to re-colonise Ireland and radiocarbon dates (BP) are used, since it would not be Britain from their glacial refugia (cf Bennett et al., 1991; ARTICLE IN PRESS 1758 N.J. Whitehouse / Quaternary Science Reviews 25 (2006) 1755–1789

Willis, 1996; Stewart and Lister, 2001). Each taxon 1997; Coope, 1998). It is difficult to know exactly when displayed individual patterns of arrival, rates of spread, these species arrived because of the lack of palaeoentomo- directions, timings and attainment of distributional limits logical research covering the earlier part of the Holocene (Birks, 1989). These ‘‘wildwoods’’ (also known as Urwald) (cf Osborne, 1972, 1974, 1980; Dinnin, 1997; Coope, 1998). appear to have been structurally complex and rich The lack of research is related to lack of research interest, ecosystems characterised by significant numbers of large, rather than lack of suitable sites, although there is mixed long-lived trees, in which dead wood was a major availability of study material from this period. Some areas, component of the habitat (Buckland and Dinnin, 1993; such as Ireland, have a relative abundance of sites covering Smith and Whitehouse, 2005). The structure of these forest this period, whilst in other locations material of this date ecosystems has recently been the subject of fierce discus- are more difficult to obtain and may be buried beneath sion, led by the ecologist Frans Vera (2000), who has agricultural soils. questioned many assumptions about the natural ecological Deposits at Holywell Coombe in Kent, England, provide state of ‘‘wildwood’’. In particular, he suggests that the role a valuable record including the Lateglacial and first of large herbivores has been seriously underestimated and millennium of the Holocene (Coope, 1998). The record that they played an important part in maintaining from this site is especially interesting as it allows us to trace substantial open areas in the forests of primeval Europe. the arrival of species associated with high forest (wild- This idea challenges the widespread belief that the ‘‘wild- wood) in a southerly area of England. Deposits spanning woods’’ of Ireland, Britain and the rest of Europe were a the first thousand years of the Holocene indicate a dense, continuous canopy (cf Rackham, 1986; Peterken, community of beetles associated with open ground and 1996). Vera suggests they may have been similar in light forest. By 8630–8280 cal BC (10,580–10,230 cal BP, structure to wood pasture and parkland, an idea which 9230775 BP, Q-2710) a suite of specialist species had has support amongst entomologists (e.g. Harding and already arrived, including the jewel beetle (Buprestidae) Rose, 1986; Alexander, 2002), but generated intense Melasis buprestoides (Notable B), false click beetles discussion and debate amongst conservationists, biologists () Eucnemis capucina (RDB 1) and Dirhagus and palaeoecologists (e.g. Blumer, 2002; Eriksson et al., pygmaeus (RDB 3), and the long-horn beetle (Cerambyci- 2002; Sutherland, 2002; Svenning, 2002; Bradshaw dae) Anaglypus mysticus (Notable B)—all dependant upon et al., 2003; Kirby, 2003, 2004; Rackham, 2003; Bakker rotting wood and dry heart-wood of deciduous trees— et al., 2004; Bradshaw and Hannon, 2004; Whitehouse and whilst the non-British () mRhyncolus Smith, 2004; Birks, 2005; Mitchell, 2005; Moore, 2005), not elongatus, is normally associated with dead and rotting all of whom agree with his perspective. wood of conifers, although in this context may have been Whatever the role of grazing in maintaining living in deciduous wood (see below for discussion on its forest structure, it is likely that these forests were subject to biology). The recovery of specimens of Dirhagus pygmaeus a range of natural disturbances (e.g. forest fires, storm in the uppermost samples from the Lateglacial site at damage, insect and damage, floods), creating open, Gransmoor, Yorkshire (Walker et al., 1993), and its sun-exposed places with a considerable amount of dying recovery in Interstadial deposits by Ashworth (1973) at and dead wood, and semi-permanent open spaces (Bhiry Rodbaston Hall, Staffordshire, suggests it may well have and Filion, 1996; Whitehouse, 2000; Bradshaw and been part of the early colonising fauna. Indeed, its presence Hannon, 2004; Smith and Whitehouse, 2005). Such in both Interstadial and very early Holocene deposits small-scale disturbances would have maintained a highly suggests that it might have survived locally in warm micro- diverse faunal community (cf Kaila et al., 1997), maintain- climatic pockets during the Younger Dryas (cf Rundgren ing an open canopy in places, creating clearings, and and Ingo´lfsson, 1999; Stewart and Lister, 2001)and helping to generate a build-up of fuel upon the forest floor, highlights the swiftness with which species arrive in on occasion making the forest more susceptible to fire response to warming. (Danks and Foottit, 1989). Early Holocene and later By 500 years into the Holocene, a forest fauna had sediments often contain abundant charcoal indicating already established itself in the English Midlands at Lea frequent fires (Patterson et al., 1987; Patterson and Marsdon, Warwickshire, suggesting that development Backman, 1988; Huntley, 1993). These natural processes towards closed forest ecosystem was well underway would have created clearings, long before human activities (Osborne, 1974, 1997). Birch and willow appear to have had any impact upon the forest. been abundant, and there were species associated with These wildwoods supported a distinctive invertebrate mature dead wood, such as the (Lucanidae) population of Urwaldtiere (cf Palm, 1951; undisturbed, old cylindricum, cylindrical bark beetle (Colydi- growth relict forest taxa). Pearson (1966) and later dae) Ditoma crenata and long-horn beetles such as Hammond (1974) suggested that the insect fauna had mordax and Aromia moscata (Notable B). As achieved its present characteristics by ca 6000 cal BC (ca tree species expanded from Europe and habitats became 8000 cal BP). More recent research clearly indicates that more diverse, the associated forest fauna moved north- many elements of the thermophilous forest fauna had wards over the next few thousand years. This included a established themselves well before this period (e.g. Dinnin, diverse range of specialist saproxylics that no longer live in ARTICLE IN PRESS N.J. Whitehouse / Quaternary Science Reviews 25 (2006) 1755–1789 1759

Britain (Table 1). By about ca 6000 cal BC (ca 8000 cal BP), undertaken by Rogers (2004) on inter-tidal peats in increased diversification of saproxylic communities seems Strangford Lough, Co. Down, dated between 7320– to have occurred (Dinnin and Sadler, 1999). These 7070 cal BC (9270–9020 cal BP, 8173734 BP, UB-4587) increases were probably caused by the availability of the and 7030–6600 cal BC (8980–8550 cal BP, 7894735 BP, full range of forest habitats at different successional stages UB-4581) (McErlean et al., 2002) indicate a pine fauna and the arrival of the highly specialised thermophilous including the non-Irish bark beetles ]Hylastes ater, ]H. Urwaldtiere from their mainland European refugia (cf angustatus and weevil ]Rhyncolus ater. All species are still Bennett et al., 1991). present on the British list, the former two living in southern In Ireland, the story is rather less well-known and it is England, whilst the latter is mostly confined to the unclear when wood-dependant taxa arrived, due to the lack Caledonian forests in Scotland. The lack of high forest of investigations covering this earlier period. New work taxa associated with deciduous woods is noticeable, but

Table 1 Known pre-Linnean Holocene Coleoptera extirpated from Great Britain (England, Scotland, and Wales), listing species in taxonomic order together with information on earliest and latest calibrated date in the fossil record

Species Earliest fossil record Most recent fossil record

Carabidae: Chlaenius sulcicollis (Payk.) 4040–3820 cal BC 4040–3820 cal BC Carabidae: Oodes gracilis Vil. 4040–3820 cal BC 4040–3820 cal BC Gyrinidae: Gyrinus colymbus Er. 1300 cal AD 1300 cal AD Rhysodidae: Rhysodes sulcatus (F.)] 8565–7965 cal BC 1390–1060 cal BC Staphylinidae: Micropeplus caelatus Er. Early Holocene 4th century cal AD Pselaphidae: Batrisus formicarius Aube.# 43360–2920 cal BC 43360–2920 cal BC Buprestidae: Buprestis rustica L.# 2921–1489 BCa 2921–1489 BCa Elateridae: Porthmidius austriacus (Schrk.)# Mid-Holocene Mid-Holocene Eucnemidae: Isorhipis melasoides (Cast.)# 1390–1060 cal BC 1390–1060 cal BC Eucnemidae: Dromaeolus barnabita (Villa)] 7480–5050 cal BC 44300–4100 cal BC : laniarius Ill. 1610–1300 cal BC 1610–1300 cal BC Ostomidae: Zimioma grossum (l.)] 44350–4170 cal BC 1390–1060 cal BC Ostomidae: Temnochila coerula Westw.] 43350–3100 cal BC 43350–3100 cal BC Ostomidae: Tenebrioides fuscus Goeze] 41320–1030 cal BC 41320–1030 cal BC Cucujidae: Cryptolestes ? corticinus Er.] 43350–3100 cal BC 42000 cal BC Cucujidae: Airaphilus elongatus (Gyll.) 5000–4000 cal BC 4th century AD Cucujidae: Prostomis mandibularis (F.)] 4030–3820 cal BC 1390–1060 cal BC Chryptophagidae: Leucohimatium sp. 9250–8250 cal BC 9250–8250 cal BC Chryptophagidae:Caenoscelis subdeplanata Bris.] 42820–2660 cal BC 42820–2660 cal BC Colydiidae: Pycnomerus terebrans (Ol.)] 4030–3820 cal BC 3444–2921 BCa Colydiidae: Bothrideres contractus F.] 4030–3820 cal BC 3444–2445 BCa Endomycidae: cruciata (Schall.]) 3444–2445 BCa 1390–1060 cal BC Cisidae: Rhopalodontus bauderi Ab]. 1390–1060 cal BC 1390–1060 cal BC Anthicidae: Anthicus gracilis (Panz.) 4030–3820 cal BC 415–380 cal BC Anobiidae: borealis Israelsson] 43350–3100 cal BC 43350–3100 cal BC Scarabaeidae: Onthophagus fracticornis Preys. Early Holocene 2nd century AD Scarabaeidae: Aphodius quadriguttatus (Hbst.) 1610–1300 cal AD 1610–1300 cal AD Lucanidae: caraboides (L.)] Mid Holocene Early 19th century? Cerambycidae: cerdo L.] Mid-Holocene 2500–1750 cal BC Cerambyc: Hesperophanes fasciculatus (Fald.)] 2nd century AD 2nd century AD~ Curculionidae: Cathormiocerus validiscapus Roug. 7580–7080 cal BC 7580–7080 cal BC Curculionidae: Cyphocleonus trisulcatus (Hbst.) Lateglacial Interstadial 9250–8250 cal BC Curculionidae: auratus /bacchus (L.) 7480–5050 cal BC 7480–5050 cal BC Curculionidae: Rhyncolus elongatus Gyll.] 8630–8280 cal BC 2921–2445 BCa Curculionidae: Rhyncolus sculpturatus ‘Walt]. 43350–3100 cal BC 2921–2445 BCa Curculionidae: Rhyncolus punctulatus Boh.] 43350–3100 cal BC 1520–1310 cal BC Curculionidae: Rhyncolus strangulatus Perr.] Mid-Holocene Mid-Holocene Curculionidae: Pissodes gyllenhali (Gyll.)] 43350–3100 cal BC 43350–3100 cal BC Curculionidae: Acalles sp. 1390–1060 cal BC 1390–1060 cal BC

Please note that the timing and dates of extinctions are poorly known; last dates in the fossil record may not bear particularly good relationships to individual species’ extinction dates. This table is an updated and modified version of one originally published by Buckland and Dinnin (1993) and modified by Dinnin and Sadler (1999). This list does not include recent extinctions. Data from: Duffy (1968); Osborne (1972, 1974, 1995); Harding and Plant (1978); Buckland (1979); Girling (1980, 1982, 1984a, 1985, 1989); Whitehead (1989); Robinson (1991, 2000); Roper (1996, 1993); Whitehouse (1993, 1997a, b, 1998, 2004); Dinnin (1997); Wagner (1997); Coope (1998); Hughes et al. (2000); and Boswijk and Whitehouse (2002). Symbols: saproxylic species: ]; probably imported in furniture during the Roman period: ~. aDendrochronological dates. ARTICLE IN PRESS 1760 N.J. Whitehouse / Quaternary Science Reviews 25 (2006) 1755–1789 their absence is much more likely to be related to edaphic suggest that a significant reduction in primary forest had conditions associated with the study area and may imply occurred, with many Urwald assemblages showing the little about the wider environment. effects of forest clearance (Osborne, 1965). The forest A further site investigated that falls within this earlier insect fauna was increasingly being replaced by elements of period is Sluggan Bog, Co. Antrim. This raised bog ‘‘culture-steppe’’—insects associated with cleared land- contains an important Lateglacial and Holocene palaeoe- scapes and pasture. Significant in-roads had also been cological record (Smith and Goddard, 1991; Pilcher et al., made into the forested areas throughout much of Europe. 1995; Lowe et al., 2004). Radiocarbon and dendrochrono- By 2000 to 1000 years ago, it is doubtful whether anything logical dates indicate that around 6300–5500 cal BC (ca more than patches of secondary forests remained in the 8250–7450 cal BP) and again at 3350–3250 BC (ca 5300– lowlands, over most of the Mediterranean and parts of 5200 cal BP, dendrochronological date), pine forest invaded Western Europe accessible to cultivation (Greig, 1982; the surface of the bog (Pilcher et al., 1995). Insect material Speight, 1989). By the onset of the scientific period when associated with the first pine layer has been studied by Linnaeus started to give animals and plants Latin names, Whitehouse (in press). Many of the species are typical of Europe’s forest areas had been reduced to their smallest lowland raised bogs, but there is also a community extent since the last glaciation. Today, woodland cover in characteristic of areas of old pine forest, including three Britain is reduced to ca 8% of the landscape, over half of species which today are not included on the Irish or British which is in plantations of non-native tree species (Warren lists, the Urwaldrelikt species mR. elongatus, mRhyncolus and Key, 1991), although there has been a recent modest sculpturatus and the rare mBothrideres contractus. The expansion of lowland woodland composed of native tree former two species have also been recovered associated with species as a consequence of agricultural reform (Dennis, pine trees from Ballymacombs More (Whitehouse, in 1997). In Ireland, woodland habitat is even more im- press). From a biogeographic perspective, it is interesting poverished, with just 5% of the countryside covered with that the fauna shows similarities with those from Inter- trees (Neeson, 1997), just 1% consisting of native wood- stadial deposits at nearby Aghnadarragh, Co. Antrim, land (Pilcher and Hall, 2001). dated to 448,180 BP (SRR-2949) (McCabe et al., 1987). As early as the Mesolithic archaeological period Here, taxa characteristic of raised peats and a pine fauna (between 8000 and 4200 cal BC or 10,000– and 6200 cal BP) were recovered, including the non-Irish mR. elongatus and there is evidence for the disturbance of primary forest in mR. strangulatus. This suggests that the former species is the palynological and charcoal record (e.g. Smith, 1970, part of the original Irish fauna, which re-assembled itself 1981; Williams, 1985; Edwards, 1996, 1999; Simmons, with climate warming. New research by the author is 1996, 2001). Tipping (1994, p. 16), however, rightly points currently re-evaluating the significance of this fauna in the out that at least some identified episodes of ‘‘Mesolithic light of new material from this site. activity’’ may have been generated by autogenic forest Overall, the re-expansion of thermophilous and forest processes; moreover, the recent recognition of climatic taxa seen in the early Holocene is no more than a coming ‘‘events’’ during the early Holocene suggest that some of together of a configuration of species that had occurred these disturbances could be vegetational responses to many times previously (Buckland and Dinnin, 1993), climatic fluctuations. Millennial-scale climate variability although each interglacial appears to have had its own resulting from solar forcing (Bond et al., 1997, 2001) and peculiarities (e.g. Coope, 1990). By ca 5000–4000 cal BC (ca additional rapid, short-term, climatic shifts occurred 7000–6000 cal BP) forests had reached their maxima, with during this early part of the Holocene (Broecker et al., extensive tracts of lime-dominated forest in lowland areas 1989; Clark et al., 2001). Climate variability affected forests of England and Wales, oak in the north and west, giving in several different ways, including making them more way to birch and pine in the north of Scotland (Bennett, susceptible to disturbance and changing the frequency and 1988). In Ireland, oak, elm and pine were well established intensity of forest fires (Bradshaw and Hannon, 1992). by about 9000 years ago, with pine the most common There is far clearer evidence for human disturbance taxon particularly on infertile sites in the west, and oak and during the Neolithic, starting in 4200 cal BC (ca elm in the fertile regions, such as the south-east and north- 6200 cal BP), when we know from the archaeological east. By about 6000 cal BC (ca 8000 cal BP) had record that humans were starting to clear the landscape, spread through most of central and southern areas, but also managing forest areas through coppicing and particularly in areas prone to waterlogging or flooding pollarding (Rackham, 1986; Rasmussen, 1990). These (Watts, 1985). Hazel forest and scrub remained important changes can be observed in detail through the fossil beetle in all areas (Pilcher and Hall, 2001). record from several archaeological sites. For instance, at West Heath, Hampstead, London, Mesolithic deposits 2.2. From Urwald to culture steppe: forest clearance and its indicate a largely undisturbed landscape, with a diverse effects upon the forest fauna range of old growth forest species (Urwaldtiere). These communities included the lime-feeding bark beetle Erno- By about the mid-late Holocene ca 2500–800 cal BC (ca porus caucasicus (RDB1) and two taxa which no longer live 4000–2700 cal BP), both the pollen and beetle records in Britain, the cylindrical bark beetle *Pycnomerus ARTICLE IN PRESS N.J. Whitehouse / Quaternary Science Reviews 25 (2006) 1755–1789 1761 tenebrans, a species which displays a very scarce, sporadic slant of these investigations, it is very difficult to get any distribution in mainland Europe, where it is considered a view of how faunas changed over time, leaving just snap- relict species of old forest, and the false click beetle shots of different areas and periods. Nevertheless, a *Isorhipis melasoides another ancient forest inhabitant growing number of species have been recovered which often found in dry beech (Reitter, 1911). Within a relatively are not on the current Irish list (Table 2). short period of time, perhaps 500–1000 years, the beetles Investigations associated with a timber/brushwood indicate a shift to a chiefly managed landscape (Girling, trackway at Corlea in Co. Longford (Corlea 9), dated to 1989). Similarly, at Runnymede, Surrey, a series of ca 3580 cal BC (ca 5530 cal BP) (Caseldine et al., 2001) investigations cover a sequence of at least 6000 years, one revealed a disappointing forest fossil insect fauna, apart of the longest Holocene insect sequences studied in Britain from the weevil ]R. ater a species that probably came onto (Robinson, 2000). The Mesolithic deposits (ca 7000– the site in structural wood. Although not on the present 5000 cal BC or 9000–7000 cal BP) until the late Neolithic Irish list (Anderson et al., 1997), this species is still found (ca 4250–3650 cal BC or 6200–5600 cal BP, 51007100 BP, commonly in Britain especially in the pine forests of BM-3039, Needham, 2000) are characterised by an insect Scotland. It is not likely to indicate extant Urwald forest, fauna with a strong Urwald element. By the Late Bronze but may indicate good quality secondary forest in the area. Age the insect fauna reflects a largely agricultural land- In contrast, investigations undertaken at Derryville Bog, scape (Robinson, 1991, 2000). Deposits at Dorney, Co. Tipperary, associated with a Bronze Age trackway and Oxfordshire, similarly indicate clearance of floodplain dated to ca 1600 cal BC (ca 3600 cal BP) (Caseldine et al., forest by this period (Parker and Robinson, 2003). At 2001), revealed a varied beetle fauna associated with wood. Wilsford, in the British Midlands, at ca 1880–1430 cal BC This included several Urwaldrelikt species, including (ca 3830–3380 cal BP, 3330790 BP, NPL-74) Osborne mProstomis mandibularis which is locally extinct in Ireland (1969) records a fauna that suggests grazing animals in and Britain, ]Dirhagus pygmaeus (RDB3), ]Teredus an almost tree-less landscape. These transitions from cylindricus (RDB 1) and ]Phloeophagus lignarius none of forested to cleared landscape have been observed at several which appear on the present Irish list (Anderson et al., other sites in Britain (e.g. Dinnin, 1997; Whitehouse, 1998) 1997). Presumably, the beetles were transported onto the and (e.g. Andrieu-Ponel and Ponel, 1999; Ponel site via wood used in the trackway from nearby forest, et al., 2001) with the result that by about 2500 years ago apparently old growth in character (e.g. mProstomis mature forest beetle species appear to represent an mandibularis and ]T. cylindricus) and suggesting surviving insignificant faunal element. undisturbed ‘‘wildwood’’ in this area (Caseldine et al., In Ireland, very few sites have been investigated and 2001). published which cover the temporal span required to see Research by Reilly (2003) on 13th century AD Anglo- the changes over this crucial period. Eileen Reilly has done Norman deposits at Back Lane in Dublin is rather the majority of this work (e.g. Reilly, 1996, 2003; Caseldine informative. Beetles recovered from house floors and et al., 2001), mostly associated with the analysis of structural timbers included large numbers of wood- archaeological material. Because of the archaeological dependant taxa, together with a number of rare and locally

Table 2 Known pre-Linnean Holocene Coleoptera extirpated from Ireland, listing species in taxonomic order together with information on earliest and latest date in the fossil record

Species Earliest fossil record Most recent fossil record Status in Britain

Hydrophilidae: Cercyon depressus Steph. 13th century cal AD 413th century cal AD Notable Eucnemidae: Dirhagus pygmeus (F.)] Post 1200 BCa Post 1200 BCa RDB 3 Cucujidae: Prostomis mandibularis (F.)] 160679 and 159079BCa 160679 and 159079BCa Extirpated Colydiidae: Teredus cylindricus Ol.] 160679 and 159079BCa 160679 and 159079BCa RDB 1 Colydiidae: Bothrideres contractus F.] 6500–6000 cal BC 6500–6000 cal BC Extirpated Tenebrionidae: Blaps leithifera Marsham 11th century cal AD 11th century cal AD Cerambycidae: scalaris (L.)] 13th century cal AD 13th century cal AD Notable A Scolytidae: Scolytus mali (Bech.)] 7320–6600 cal BC 13th century cal AD Notable B Scolytidae: Hylastes ater (F.)] 7320–6600 cal BC 7320–6600 cal BC Scolytidae: Hylastes angustatus (Hbst)] 7320–6600 cal BC 7320–6600 cal BC Scolytidae: Leperisinus orni (Fuchs.)] 13th century cal AD 13th century cal AD Notable B Scolytidae: Acrantus vittatus] 13th century cal AD 13th century cal AD Curculionidae: Rhyncolus elongatus Gyll.] 6500–6000 cal BC 2000–3000 BCa Extirpated Curculionidae: Rhyncolus sculpturatus Walt] 6500–6000 cal BC 2000–3000 BCa Extirpated Curculionidae: Rhyncolus ater (L.)] 7320–6600 cal BC 13th century AD

Limitations as per Table 1. Data from: Caseldine et al. (2001); O’Connor (1979); Reilly (1996, 2003); Rogers (2004); Whitehouse (in press). Symbols: saproxylic species: ]. ARTICLE IN PRESS 1762 N.J. Whitehouse / Quaternary Science Reviews 25 (2006) 1755–1789 extinct taxa. Three different species of bark beetle where structures on urban sites. The woodland element, however, recovered (]Leperisinus orni (Notable B), ]Scolytus mali was very small, with no ancient woodland species and only (Notable B), and ]Acrantus vittatus), the long-horn beetle a small number of more ubiquitous species such as the ]Saperda scalaris, found mainly in dead and dying wood-borers Anobium punctatum and planus deciduous trees, usually of large girth (Palm, 1959) and present. These findings hint that the survival of ancient the weevil ]R. ater which is usually associated with dead woodland may have varied considerably across the pine and oak wood. None are recorded today in Ireland, country, although more sites would need to be investigated although they still live in Britain, ranging from relatively to ascertain these variations. common to rare or endangered. Reilly (2003) suggests the species were probably transported into Dublin in structural 2.3. The effects of clearance: a regional case study from the timber or firewood and indicate that good-quality wood- humberhead levels land was still present within the catchment areas of the city during the Medieval period. These findings are in contrast It is possible to examine the nature of the faunal changes to those of an early Christian ‘‘rath’’ site (a protected discussed above in greater detail, through a regional habitation or fort typical of this period) in Co. Antrim palaeoentomological investigation into the effects of forest (Kenward and Allison, 1994). Here, the insect assemblage clearance in the Humberhead Levels (Whitehouse, 1998). was remarkably similar to those found in and around This region (Fig. 3) has been exceptionally well studied for

ary Estu ber use um r O H Goole ve Ri Rawcliffe e Air River er Old course of Swinefleet Riv River Don Dutch

Cowick Old course of Goole Fields River Went Old course of River Trent Swinefleet River Don ent r W Rive Warping New Turnbrigg Drain Junction Dike Canal THORNE Sutton Common MOORS Eastoft

Thorne Crowle and Don forth tain anal River S by C ead Stainforth K Scunthorpe Old course of River Don Folly Drain Hatfield Chase Hatfield Sandoft Lindholme Old course of Island River Idle 0 5 km SOUTH HATFIELD MOORS YORKSHIRE 30 Epworth N

30 Wroot Isle of Axholme ne 15 30

River Tor 30 15

Rossington Bridge Bykers Dike Idle River MistertonCarr

Fig. 3. Map of Humberhead Levels, showing sites discussed in text. Re-drawn from an original by P.I. Buckland.r Thorne and Hatfield Moors Conservation Forum. ARTICLE IN PRESS N.J. Whitehouse / Quaternary Science Reviews 25 (2006) 1755–1789 1763 its insect fauna, particularly the raised mires of Thorne and Medieval and post-Medieval deposits at Cowick (Hayfield Hatfield Moors. Several other sites in the region have also and Greig, 1989) and Woodhall (Wagner, 1997) all provide been studied, mainly floodplain deposits at Misterton Carr, additional valuable information. on the Idle floodplain (Osborne, 1978; Whitehouse, 1998), Thorne and Hatfield Moors have been the subject of at Rossington Bridge on the Torne floodplain (White- extensive archaeological and palaeoecological investiga- house, 1998) and at Sandtoft on the Don (Buckland and tions (Smith, 1958, 1985, 2002; Dinnin et al., 1997; Sadler, 1985). Table 3 shows the list of extirpated species Boswijk, 1998, 2003; Boswijk et al., 2001; Chapman and recovered from all sites in the region. A further site to the Gearey, 2003; Van de Noort, 2004) coupled with extensive south, at Bole Ings on the Trent floodplain (Dinnin, 1997; research into their rich and diverse fossil beetle assemblages Brayshay and Dinnin, 1999), the Iron Age deposits at (Buckland and Kenward, 1973; Buckland, 1979; White- Sutton Common (Roper and Whitehouse, 1997) and house, 1993, 1997a, b, 1998, 2000, 2004; Roper, 1996, 1993;

Table 3 List of extirpated species recovered from the deposits investigated in the Humberhead Levels

Species TM HM ROS MISS Reference Other records

]Pterostichus O Whitehouse and Eversham None angustatusa (unclear (2002) status with regard to its ‘‘extinction’’ Rhysodes sulcatus O Buckland (1979) Shustoke, Warks (Kelly and Osborne, 1965); Somerset (Girling, 1980); North Sea Floor, E. Anglia (Blair, 1935); Cumbria (Hughes et al., 2000). ]Buprestis rusticab O Skidmore (unpubl.) None Isorhipis melasoides O ? OOBuckland (1979); West Heath Spa, Herts (Girling, 1989); Somerset Whitehouse (2004); Osborne (Girling, 1980). (1978) Zimioma grossum O Buckland (1979) Runnymede, Surrey (Robinson, 1991). ]Temnochila coerula O Roper (1996) and None Whitehouse (1997a) Tenebrioides? fuscus O Whitehouse (2004) None ]Cryptolestes OO Whitehouse (1997a, 2004). None corticinus Pycnomerus tenebrans O Whitehouse (1998) West Heath Spa, Herts (Girling, 1989); Minsterley, Salops (Osborne, 1972); Stileway, Somerset (Girling, 1985). Prostomis OO Buckland (1979); Boswijk Sweet Track, Somerset (Girling, 1984a); mandibularis and Whitehouse (2002) Derryville Ireland (Caseldine et al., 2001). Bothrideres contractus O Whitehouse (1998) Stileway, Somerset (Girling, 1985); Sluggan Bog (Whitehouse, in press) Mycetina cruciata O Buckland (1979); None Whitehouse (1998) Rhopalodontus bauderi O Buckland (1979) None ] Stagetus borealisc O Whitehouse (1993, 1998) None Caenoscelis O Roper (1996) None subdeplanatad Acalles sp. O Buckland (1979) None ]Rhyncolus elongatus OO Whitehouse (1997a); Roper Worldsend, Salops. (Osborne, 1972); Hollywell (1996); Whitehouse (1998, Coombe, Kent (Coope, 1998); Sluggan Bog, and 2004). Ballymacombes More, Co. Antrim (Whitehouse, in press) ]R. punctulatus OOO Whitehouse (1997a, 1998, Stileway, Somerset (Girling, 1985) 2004) ]R. sculpturatus OO Whitehouse (1993, 1998, Sluggan Bog and Ballymacombes More, Co. 2004); Boswijk and Antrim (Whitehouse, in press) Whitehouse (2002) ]Pissodes gyllenhali O Whitehouse (1997a) None

TM ¼ Thorne Moors; HM ¼ Hatfield Moors; ROS ¼ Rossington; MISS ¼ Misterton Carr. ] ¼ Pinus-loving species. aSpecies is on the current British list but is commonly believed to be an introduction earlier this century. Since this ground beetle is unlikely to have been overlooked during collecting, it is likely that it became extinct until its re-introduction. bPeter Skidmore recovered B. rustica from a pupal chamber in a pine whilst fieldwalking. cOriginally identified as ‘‘Species A’’ by Whitehouse (1993). dThis species was added to British list by Johnson (1966). Became extirpated before its ‘‘re-introduction’’? ARTICLE IN PRESS 1764 N.J. Whitehouse / Quaternary Science Reviews 25 (2006) 1755–1789

Boswijk and Whitehouse, 2002; Smith and Whitehouse, proportion of saproxylics, including a considerable number 2005). They represent probably one of the most intensely of rare and extirpated Coleoptera. In total, a further six studied sites in the world for their fossil beetle record and species were added to the list of locally extirpated taxa arguably the best examples of fossil Urwald faunas in the recovered from Thorne Moors, four of which had never Northern Hemisphere. There are probably very few, if any, been previously found in British contexts. Additionally, a comparable modern European sites still containing such a good proportion of the saproxylic species were associated valuable ancient forest fauna. The sites represent examples with pine woodland (e.g. locally extinct *Temnochila of a habitat which has virtually disappeared from Ireland coerulea,*Pissodes gyllenhali,*Cryptolestes corticinus, and Britain—the mire woodland (Smith and Whitehouse, Whitehouse, 1997b). The abundance of insects associated 2005). The sites also represent the most temporally recent with dead and decaying timber suggests an area of old post-glacial sites that display diverse elements of this growth forest, with mature and older trees which included Urwaldrelikt fauna, suggesting that undisturbed forest a diverse and well-developed range of types of wood decay. persisted in this area of wetland for a considerable period, A further site was examined by Whitehouse (1998) and long after forest had been cleared elsewhere. The Moors Boswijk et al. (2001) who studied the transition from oak- were located in a vast area of agriculturally un-usable to pine-dominated woodland on the northern side of wetland which probably ensured the survival of its ancient Thorne Moors. Throughout both phases there were many forest and associated fauna, effectively creating an faunal elements associated with mature, ancient woodland ‘‘ecological island’’ (cf MacArthur and Wilson, 1967), such as *Pycnomerus tenebrans, Teredus cylindricus (RDB which was buffered from external environmental influ- 1), *Mycetina cruciata, mainly characterised by species ences. It is probable that similar ‘‘islands’’ existed else- associated with oak, with some birch and alder associates. where, particularly on areas of developing wetlands such as By the next phase, however, there was a marked increase in in the Somerset Levels, Cheshire Mosses in England and pine-loving species and a corresponding decline in decid- Irish Midlands bogs. Their island status, however, whilst uous-wood associates. protecting them from outside influences also had the effect On Hatfield Moors, the faunal record spans the period of isolating their faunas, creating an ecological fragment from 3618 BC (dendrochronological date, Boswijk and with no opportunity for expansion (cf Ashworth, 2001; Whitehouse, 2002, ca 5568 cal BP) to 560–680 cal AD (ca Opdam and Wascher, 2004). 1390-1270 cal BP, 1405745 BP, SRR-6120, Whitehouse, On Thorne Moors, four fossil insect sampling locations 2004), including seven non-British beetle species, all have been studied (Buckland, 1979; Roper, 1996, 1993; saproxylics today restricted to isolated enclaves within Whitehouse, 1997a, 1998), with five on Hatfield Moors mature forests of mainland Europe (Table 3). A discussion (Whitehouse, 2004). The faunal record for Thorne Moors of their significance is provided by Whitehouse (2004).In ranges in date from ca 3400 BC (dendrochronological date, contrast to Thorne Moors, there appears to have been a Boswijk, 1998) to 1420–1260 cal BC (3370–3210, 30607 very limited earlier oak woodland phase at Hatfield Moors, 65 BP, CAR-180) (Buckland, 1979). Plant macrofossil dated dendrochronologically to 3618–3408 BC, succeeded evidence and dendrochronological investigations of the by a pine-heath woodland, between 2921 and 2445 BC (ca mire trees illustrate that after an earlier phase of oak 4871–4395 cal BP, Boswijk and Whitehouse, 2002). Strati- woodland, lasting between 3777–3017 BC (dendrochrono- graphic evidence from elsewhere on the site indicates that logical dates, Boswijk, 1998, ca 5727–4967 cal BP), pine and that pine continued growing on the site during periods of birch became the dominant tree species, particularly over mire dryness. On the southern margins a remnant of pine- the period 2921–1489 BC (dendrochronological dates, heath community survived until ca 560–680 cal AD (ca Boswijk, 1998, ca 4871–3439 cal BP). However, on the 1390–1270 cal BP, 1405745 BP, SRR-6120) (Whitehouse, edges of the mire, areas of oak woodland remained 2004). Evidence from further afield, from sequences important until 1420–1260 cal BC (ca 3370–3210 cal BP, examined at Rossington (Whitehouse, 1998) and Misterton 3060765 BP, CAR-180) (Buckland, 1979), where a pre- Carr (Osborne, 1978; Whitehouse, 1998), indicates that historic timber trackway was also constructed, presumably pine-woodland heath continued to be an important habitat to allow people to cross the bog and make use of the bog on the higher sandy areas above the floodplain. resources. Buckland (1979) examined beetles from the Thus, an impressive range of beetle species associated basal peat, finding a diverse range of rare and locally with oak- and pine-dominated woodland have been extinct taxa, including seven species no longer found in recovered from both sites. The forest fauna is dominated Britain. One of the most frequent fossils from the samples by two categories of saproxylics: those that are predators was the flat bark beetle mProstomis mandibularis,an of primary invaders of freshly dead trees or wood and Urwaldrelikt today virtually restricted to isolated popula- species that live in well-rotted, dead, fungoid wood. There tions within the Central European forests (Palm, 1959; is also the notable presence of species apparently associated Horion, 1960). In 1993, research carried out by Roper with burnt woodland, particularly pinewood (cf White- (1996, 1993) and Whitehouse (1993, 1997a) from deposits house, 2000). Thorne and Hatfield Moors together contain dated to 3350–3100 cal BC (ca 5300–5050 cal BP, 45157 the highest number of extirpations of any site in the British 70 BP, CAR-232, Smith, 1985, 2002) indicated a high Isles, with 18 species of nationally extinct Urwaldrelikt ARTICLE IN PRESS N.J. Whitehouse / Quaternary Science Reviews 25 (2006) 1755–1789 1765 beetles as well as 49 rare or threatened wood associates a few sites, probably reflects the decline of its host within (Buckland, 1979; Whitehouse, 1998; Smith and White- the landscape due to preferential use of this tree for house, 2005). As Smith and Whitehouse (2005) discuss, it is wood and fodder. Lime, together with elm, is palatable likely that Thorne and Hatfield Moors have produced such to stock—the bark and leaves were once widely used large numbers of extinct species due to a combination of in Europe as fodder, and the decline of these trees may factors that make them exceptional (e.g. significant be related to this (Turner, 1962). Other trees may have amounts of deadwood because of the rising water table been converted to coppice or pollards, which inhibits (Buckland, 1979); large amounts of dead pine wood flowering and could at least partially explain the lime (Whitehouse, 1997b). These conditions have ensured an palynological decline, although the fact that its beetle, outstanding entomofaunal record from the two sites, but it Ernoporus caucasicus, appears to disappear from the is perhaps worth highlighting that investigations of mire fossil record by the Early Bronze Age, suggests the decline woodlands at Sluggan Bog and Ballymacombs More, Co. is a real one rather than a palynological ‘‘event’’. It is Antrim (Whitehouse, in press) have yielded similar likely that by this time the frequency of the tree had numbers of species to those from Thorne and Hatfield already been severely reduced (Buckland and Dinnin, Moors. This suggests the sites may not be particularly 1993). The evidence from the Humberhead Levels fits into unusual within the wider context of mire woodlands, but this pattern. are clearly somewhat unusual compared with other forest In addition to clearance of lime forest, beetle faunas types, as discussed more fully in Smith and Whitehouse indicate that forest clearance had intensified by ca (2005). 1500 cal BC (ca 3400 cal BP) on the Torne floodplain, The mire assemblages from Thorne and Hatfield Moors although patches of primary forest remained (Whitehouse, are notable for their absence of beetles associated with lime 1998). The evidence indicates areas of grassland, with (Buckland, 1979), which was an important component species which are often associated with waste, cultivated within British lowland forests of this period (Greig, 1982). and open ground and an absence of tree-associated beetles; Lime is absent from pollen records and seems to have never the change in composition of aquatic beetle faunas also reached Ireland (Birks, 1989). Oak-alder forest, itself suggest that higher amounts of sediment were entering the largely the result of human interference in the succession, river system, possibly caused by clearance and agricultural largely took over as the most frequent forest type there- activities upriver (Whitehouse, 1998). This is in contrast to after, at least in many areas of lowland Britain. One bark nearby Thorne and Hatfield Moors, where Urwald forest beetle in particular, Ernoporus caucasicus (RDB 1), is persisted at least until 1420–1260 cal BC (ca 3370– found in most mid-Holocene forest insect assemblages 3210 cal BP, 3060765 BP, CAR-180, Buckland, 1979). By (Buckland and Dinnin, 1993; Dinnin, 1997). However, it the Roman period (1st century AD, ca 1900 cal BP) the has been found in roughly contemporaneous floodplain fossil insect and pollen evidence indicates a substantially deposits from Thorne Waterside, a few kilometres west cleared landscape, with few areas of forest and no insects of Thorne (43020–2620 cal BC, ca 4970–4570 cal BP, associated with ancient woodland. Pasture areas are 42307100 BP, BIRM-358) (Buckland, 1979), and is indicated by insects associated with grazing animals and abundant in the samples from Misterton Carr in the Idle grassland. Changes amongst the aquatic beetles suggest floodplain, south of Hatfield Moors (43300-2700 cal BC, increased levels of eutrophication and sedimentation in the ca 5250–4650 cal BP, 43307100 BP, BIRM-359) (Buckland river Torne and its catchment. and Dolby, 1973; Osborne, 1978). Ernoporus caucasicus is Similar environmental changes are also evident in the also present in the Torne floodplain at Rossington at about assemblages from Misterton Carr on the Idle (Osborne, this time, but disappears sometime after ca 1690– 1978; Whitehouse, 1998). Here, the landscape appears to 1450 cal BC (ca 3640–3400 cal BP, 3290745 BP, SRR- have been subject to less intensive clearance at least until 6131), which is roughly contemporary with Turner’s the latter parts of the prehistoric period; the lack of a (1962) primary lime decline (ca 1750–1100 cal BC, ca proper chronology for the upper part of the sequence and 3700–3050 cal BP, 31707115 BP, Q-481) evident in pollen the limited nature of the faunal list makes it difficult to be diagrams of this time. In contrast, at Bole Ings, in the Trent more precise about the nature of clearance (Osborne, 1978; valley, a few kilometres south of Misterton Carr, this bark Whitehouse, 1998). A site to the west of Thorne Moors, at beetle seems to disappear sometime after ca 2140– Sutton Common, ca 750–390 cal BC (ca 2700–2340 cal BP, 1740 cal BC (ca 4090–3690 cal BP, 3570770 BP, BETA- 2370750 BP, GU-5524) indicates a largely cleared land- 75271) (Dinnin, 1997), suggesting lime may have been scape, with little evidence for primary forest (Parker cleared from the landscape in the Trent valley earlier than Pearson, 1997; Roper and Whitehouse, 1997). Further in the Humberhead Levels. south, in the Trent valley, pollen and insect evidence The present and former distribution of this bark beetle suggests that relatively undisturbed forest survived on the suggests that it is a relic of the formerly dominant lime- marginal floodplain soils until sometime between 2140– forest across much of lowland southern England and the 2000 cal BC and 41040–800 cal BC (ca 4090–3950 cal BP, Midlands (Buckland and Dinnin, 1993). Its decline, from a 3570770 BP, BETA-75271 and ca 2990–2750 cal BP, former widespread distribution to its present occurrence on 2750760 BP, BETA-75269) (Brayshay and Dinnin, 1999). ARTICLE IN PRESS 1766 N.J. Whitehouse / Quaternary Science Reviews 25 (2006) 1755–1789

Buckland (1979) considered that by the end of the 3. Discussion Roman period, in the 4th century AD (ca 1600 cal BP), primary forest on all but the most marginal land had been 3.1. When did species become extirpated? cleared for agriculture, an impression which is corrobo- rated by extensive field system crop-marks (Riley, 1980). Based on the present data, it is difficult to know when Buckland and Sadler’s (1985) comments concerning the many species disappear from the fossil record. Super- Roman insect faunas from Sandtoft are revealing: ficially, many of the last fossil dates (Table 1) indicate that species were still present within the British landscape sometime between ca 3000 cal BC and 1000 cal BC (ca By the Roman period areas of river bank must have 5000–3000 cal BP), after which many seem to disappear differed little from the over-managed species-poor from the fossil record. Many of these last dates, however, fringes of many fenland streams of the present dayythe come from isolated locations such as Thorne and Hatfield evidence combines to imply a mixed farming economy in Moors, so it is hard to know how widespread many of a wholly cleared landscape, perhaps not too dissimilar these species still were in the wider landscape. Moreover, from the modern one (Buckland and Sadler, 1985, these dates probably reflect the larger number of natural p. 246). deposits studied (where one would best expect to have an insight into forest faunas) from this period compared with During the later Roman period the Torne, Idle and Don subsequent periods; this is compounded by the difficulty of floodplains become increasingly waterlogged, with the finding suitable deposits covering the last 2000 years. Even eventual deposition of alluvium, partially in response to where these exist, few have been studied. We should be higher sea-levels (Dinnin and Lillie, 1995; Long et al., open to the possibility that at least some of these species 1998), but perhaps also because of increased soil erosion may have persisted in certain areas until relatively recently. due to agricultural intensification and innovation (cf Such an example is the long-horn beetle Cerambyx cerdo Buckland and Sadler, 1985). Effects of deforestation whose current status is ‘‘X’’ (extinct) in Britain. Several resulted in changes of sedimentation in many major rivers, fossil records of this species indicate that it was present in and drastically altered communities of water and riparian Britain about 4000 years ago (Duffy, 1968; Harding and beetles (Osborne, 1988; Smith, 2001). Similar destabilisa- Plant, 1978). Duffy (1968) reviewed the records for this tion events are known from other sites in Britain (e.g. Kelly species and noted that it was recorded by several and Osborne, 1965; Lambrick and Robinson, 1979; Coleopterists in the earlier part of the 19th century, Needham, 1985, 1992), including the Trent Valley (Dinnin, suggesting that it may have managed to survive in some 1997), varying in date from area to area, in accordance areas until then. Declines in its records during the 19th with soil and vegetation type, and degree of landscape use. century suggest that its population was contracting rather Despite the impression of a largely cleared landscape in than expanding at that time. Moreover, no thorough the Humberhead Levels by the end of the Roman period, review of old collections has been carried out to investigate excavations at the Medieval moated manor house at whether other similar early British records of the extirpated Cowick, (moat dug AD 1322–1324, ca 628–630 cal BP) species exist. Even within major institutions, cataloguing of 5 km west of Thorne Moors, suggests that a relic pine and insect collections is still in progress and there are smaller, ancient forest fauna survived into the medieval period in a lesser-known collections owned by small institutions and landscape of pasture woodland (Hayfield and Greig, 1989). private individuals that would merit specialist attention. Wagner (1997) recovered several ancient woodland species Indeed, many old species records were dismissed by later such as Teredus cylindricus (RDB 1) and the pine-loving R. coleopterists, being regarded as ‘‘European’’ rather than ater from another moated site, Wood Hall, Womersley, British records and were thus never included within North Yorkshire, 15 km west of Cowick in samples modern check-lists (Osborne, 1997). Such assumptions spanning ca 1400 AD to the 18th century AD (ca may be incorrect if viewed in conjunction with the fossil 550–250 cal BP). The area was protected by Forest Laws record. and suggests that despite widespread clearance in the In Ireland, the story may be slightly different—albeit Roman period, vestiges of ancient woodland remained in based on very limited research and likely to be modified in the Humberhead Levels, raising the possibility that other years to come. The data presented in Table 2 show two ‘‘extinct’’ taxa may have survived in refugia into this period distinct trends: species which are absent after ca (Whitehouse, 2004). 2000 cal BC (ca 4000 cal BP) and a further group which The Humberhead Levels represents one of the best survives until at least the 13th century AD. This may studied areas of the British Isles for its fossil beetle record. indicate a phased process, whereby some specialised species Research undertaken on sites over the last few decades were lost sometime during prehistory, followed by a further highlights the history of woodland clearance in this region, loss during the Medieval and post-Medieval period, hinting a story which was repeated in many other areas, although that some areas of good quality forest survived into this each region is likely to have been subject to distinct period. The more unusual Urwaldtiere are not in evidence differences. by the later date, but the palaeoecological record covering ARTICLE IN PRESS N.J. Whitehouse / Quaternary Science Reviews 25 (2006) 1755–1789 1767 this period is poorly studied at present and is dominated by (cf Szwecki, 1987). For instance, Szwecki (1987) notes that investigations associated with urban deposits, where one the number of trees inhabited by saproxylic insects in 150 might not expect to find such species (e.g. Reilly, 2003). year-old oak forest is a third of that recorded from 220–350 That some taxa are being recovered in such material is all year old ones, with the lowest number of xylophages the more surprising, and suggests certain species’ ability to recorded on living trees between 120–150 years old (15.5%). make use of some anthropogenic habitats in some cases (e.g. wattle) and/or their transportation into the area with 3.4. The importance of micro-habitats imported wood from hinterlands. Besides the quality of the decaying trunks, various 3.2. Why did species become extirpated? environmental factors appear to affect the occurrence of both common and rare saproxylic species (Kaila et al., The biology of many of these saproxylics is considered in 1997). Kaila et al. (1994) for instance, demonstrate strong detail below, together with some other factors that may associations between the presence of saproxylic beetles in have affected these species’ biogeography. A consideration un-logged forests in and the colonisation of birch of the role of climate change in extirpations is also made. by specific species of polypore fungi. They concluded that Saproxylic species in primeval forests feed on decaying the Polyporus species decaying the wood are important for wood or on the fungi responsible for its decomposition individual saproxylic beetle species, and that there are (Va¨isainen et al., 1993). Two characteristics of old-growth different specialised beetle assemblages associated with forests are important for many specialist species: coarse trunks attacked by different Polyporus fungi (Kaila et al., woody debris and large, mature trees. There has been a 1994)—*Zimioma grossum, one of the extirpated species resurgence of interest in these saproxylics in recent years, recovered from Thorne Moors (Buckland, 1979), is a taxon particularly in Fennoscandia, where there has been concern which seems to be tied to a particular Polyporus about the status of many of these saproxylics due to forest (Juha Siitonen, pers. comm., 1998). The availability of clearance and the loss of natural disturbance regimes (e.g. decaying wood in sun-exposed positions appears to be Helio¨vaara and Va¨isa¨nen, 1984; Bistro¨mandVa¨isa¨nen, critical to many species (Kaila et al., 1997) and may explain 1988; Va¨isainen et al., 1993; Kaila et al., 1994, 1997; Siitonen the importance of forest fires in creating such habitat. and Martikainen, 1994; Haila et al., 1995; Martikainen Lichen species also appear to be essential to certain faunal et al., 1996; Niemela¨,1997; Ranius and Jansson, 2000). elements (Spence et al., 1997); the loss of lichen affects the whole ecosystem through the removal of an important 3.3. The importance of dead wood and disturbances component from the food chain and loss of insulation on the forest floor. Old growth forests are characterised by large volumes of dead, standing and fallen trees as well as old living trees. In 3.5. Forest history and management some cases, the volume of this wood can be equal to that of living trees (e.g. the great forest of Kubany Wald, Forest history may have been decisive in the survival of Schattawa, ) (Elton, 1966). These forests are subject many of these invertebrates. Va¨isainen et al. (1993) suggest to a range of natural disturbances, with a considerable that if the forest has lost its primeval character due to amount of dying and dead wood. Patches emerge locally human activities, then the fauna may be irretrievably lost quite frequently and regionally may always be present, with during this period of forest change. In many cases, species moderately small inter-patch distances (Kaila et al., 1997). are not able to re-colonise forests: The role of these natural disturbances is often highly Although old-growth specialists may occur in low important in these forests. numbers in the managed forest, perhaps maintained by The abundance and availability of dead woody debris is immigrants from the old-growth sources, these sink a key component of the primeval forest, supporting many populations probably disappear if cut-overs become too of the extirpated and rare species such as those recovered in large, hampering movement of individuals between the the fossil record. This wood-decomposing system has preferred old-growth fragments (Niemela¨, 1997, p. 607). exceptionally high species diversity (Bistro¨mandVa¨isa¨nen, 1988); for instance, Elton (1966), pointed out that if dead Fennoscandian researchers have attempted to quantify wood is removed from the forest ecosystem, the system the effect of clearance and management of forests on loses a fifth or more of its invertebrate fauna, although that saproxylics and suggest that of the ca 1200 threatened figure is now thought to be considerably higher (Keith species in , over half are sensitive to both thinning Alexander, pers. comm., 2005). and clear-cutting of the forest (Niemela¨, 1997). Studies in Many of the rare and extirpated species recovered in the Finland (Va¨isainen et al., 1993; Kaila et al., 1997) suggest fossil record are associated with very mature trees, so the that whilst there may be little difference in species richness extirpations and contractions in species’ range indicate that between old growth and managed forests, the number of not only dead wood was being removed from forests, but rare species is much higher in old-growth forest than in that trees were not allowed to reach full maturity managed forest (Niemela¨, 1997), and furthermore that the ARTICLE IN PRESS 1768 N.J. Whitehouse / Quaternary Science Reviews 25 (2006) 1755–1789 specialist species of old-growth forest do not survive in the species is, in general, poorly known, and most probably managed forests (Bistro¨mandVa¨isa¨nen, 1988). Indeed, varies remarkably between species (Kaila et al., 1997). amongst the abundant saproxylic species, there appear to Much of the available evidence supports the view that they be few species in common between managed and unma- have very limited powers of dispersal (Wormwell, 1977; naged forests (Va¨isainen et al., 1993) and it seems that Warren and Key, 1991). It is unlikely that habitat managed forests are unable to maintain the original fauna availability was a problem in primeval forest conditions, of primeval forests. They conclude that ‘‘The real primeval where there was almost continuous forest cover and an species seem to be prisoners of the existing remnants of abundance of dead wood, since suitable hosts may have primeval forests’’ (Va¨isainen et al., 1993, p. 107) and that been present within a distance of a few metres (Warren and these species are likely to disappear, especially in small Key, 1991). Under such conditions, there was presumably forest fragments. little selection pressure to evolve strong mobility (McLean Thus, temporal continuity and stability of habitat may and Speight, 1993), where more-or-less continual habitat be a significant component in the development and would have been available. The poor mobility of many of maintenance of many of these saproxylic communities these species is likely to have been an important factor in (Spence et al., 1997). This might explain the longevity of their decline and extirpation in the British Isles. the records from Thorne and Hatfield Moors, where However, the fossil insect evidence indicates that this is continuity of habitat appears to ensure at least some of not the whole story. One has to ask how forest Coleoptera, these species’ survival longer than elsewhere. with apparently low dispersal abilities, appear so promptly in the early Holocene fossil record. Despite their ‘‘low 3.6. Effects of clearance and forest fragmentation on powers of dispersal’’, components of the ‘‘wildwood’’ saproxylics fauna were able to reach northern England and Scotland and expand into Ireland in the early part of the Holocene. In an ancient forest, organic matter and nutrients It seems that as long as Urwald forest existed, mobility and accumulate in the standing crop of trees and shrubs and dispersal of these saproxylics was not a major problem. in the organic debris of the forest floor. All this debris acts Indeed, when forest has reached an appropriate age, as an insulating layer (Bistro¨m and Va¨isa¨nen, 1988). colonisation may occur easily from adjacent old forest Clearance has the effect of creating an exposed habitat, (Elton, 1966) and species may well have made good use of with increased temperature and decreased moisture and other modes of dispersal, such as wood rafting (e.g. shade (Atlegrim and Sjo¨berg, 1995). Several species Johansen and Hytteborn, 2001). Moreover, it raises apparently cannot live in the dry, open conditions of questions about the location of the refugia of many species, managed forests, and require the presence of stable air a point which is returned to below. humidity, protection from sunlight, wind and particularly Once the continuity of forests was broken through drought. Such factors are also likely to decrease the widespread deforestation, however, many species were no survival of invertebrate larvae (Atlegrim and Sjo¨berg, longer able to expand to other patches of habitat and 1995), and will also affect the wood-decomposing processes became isolated within small biogeographic islands, such as attributable to fungi, and consequently the food quality of on the wetlands of the Humberhead Levels. Even where many invertebrates (Va¨isainen et al., 1993). regenerated woodland created linking corridors between Geographical and structural forest fragmentation can areas of ‘‘wildwood’’, species were apparently not able to have a range of effects upon forest invertebrates. For re-colonise forest fragments. It seems that not only do instance, instead of a decline, species richness may actually species require corridors of habitat to re-expand, but that increase in small forest fragments (Niemela¨, 1997), the quality of these linking forests is also important, although the duration and quality of this ‘‘richness’’ is needing to be of the appropriate ‘‘wildwood’’ longevity and unclear. Geographical isolation, however, usually reduces required structure. Much of the Fennoscandian literature species richness as some species are poor dispersers or indicates that once a forest has lost its primeval character, restricted to particular habitat types. Isolation can cause species are unable to re-colonise from nearby patches and extirpation of a species, or genetic changes and shifts in the fauna may be irretrievably lost (Bistro¨mandVa¨isa¨nen, habitat quality (Dennis, 1997). It seems likely that forest 1988; Va¨isainen et al., 1993; Siitonen and Martikainen, fragmentation and the concurrent isolation of many species 1994). Species thus become further isolated and confined to may have been a key factor in the extirpation of at least small biogeographic islands. some species evident in the fossil record. It would seem that disturbance, destruction, and frag- mentation of habitat, coupled with poor mobility are the 3.7. Mobility prime factors in the disappearance of many of these species.

Saproxylic species are characteristically found in only a 3.8. The decline of pine and heath proportion of potentially suitable trees, which are often only a small fraction of the total tree population of most There appears to be another contributory factor to these forests (McLean and Speight, 1993). The mobility of insect extirpations. Many species found in the Humberhead ARTICLE IN PRESS N.J. Whitehouse / Quaternary Science Reviews 25 (2006) 1755–1789 1769

Levels and Irish bog sites are pinicolous. Whitehouse Host extirpation is likely to be one of the most important (1997b) suggested that the decline of pine in the reasons for these species’ contraction in range, although British landscape might have had a significant role in the the timing of their disappearance may be more complex, decline and extirpation of at least some saproxylics. During and possibly more recent than it might appear. the Holocene, pine became increasingly restricted to marginal areas (McNally and Doyle, 1984). In Britain, 3.9. The importance of fire habitats by ca 3000 cal BC (ca 5000 cal BP) pine appears to have undergone a sudden and widespread decline Buckland and Dinnin (1993) first pointed out how the (Bennett, 1984, 1995; Gear, 1989; Bridge et al., 1990; disappearance of fire habitats might have contributed to Gear and Huntley, 1991). The most widespread view is the demise of some saproxylic elements of the British that the decline was caused by a shift to cooler and fauna. Substantial numbers of the pine and boreal insect wetter climatic conditions, by promoting the expansion of fauna are fire favoured (Hunter, 1977; Wikars and blanket peat to the detriment of pine (Bennett, 1984; Birks, Schimmel, 2001). Forest fires increase insect diversity by 1989; Gear and Huntley, 1991) and increasing wetness on creating suitable habitat for the development of strictly raised bogs (Smith and Goddard, 1991; Boswijk, 2003). pyrophilous species; secondly, they kill trees, which allows Bradshaw (1993) also suggests that the decline in popula- species which are dependant upon dense piles of sun- tions may have been connected to decreased occurrence of exposed wood to develop. Fire also alters the age-structure wildfire. and tree-species composition of an area (Wikars, 1992; Dendrochronological evidence suggests that in some Wikars and Schimmel, 2001). Muona and Rutanen (1994) areas, particularly in mire woodlands where pine was an indicate that fire specialists form a distinct community, important component, it declined in a protracted fashion developing almost exclusively in burned areas, usually only (Lageard et al., 1999). Boswijk (1998, 2003) noted that her 1–5 years after a fire. chronologies indicated its presence on the mire at Thorne Reviews by Whitehouse (2000) and Whitehouse and Moors long after the palynological pine decline (cf Smith, Eversham (2002) argued that at least four extirpated 1985, 2002). One explanation for this anomaly is that species appear to have an affinity to fire: *Zimioma pollen productivity from pine growing in wetland environ- grossum.;*Mycetina cruciata;*;*R. ments may be severely reduced (Bennett, 1995). Moreover, elongatus. These were associated with charred wood on there is good fossil beetle evidence to suggest that pine Thorne and Hatfield Moors. Two other species that are remained important in some areas of Britain into the part of the British fauna today (Pterosticus angustatus historic period (Whitehouse, 1997b, 2004). In Ireland, the (Notable B) and Caenoscelis subdeplanata), but which have fossil beetle record is rather too fragmentary to be very been regarded as introductions until their recovery in fossil informative, although the presence of the weevil R. ater contexts, are also considered to be attracted to fire. from 13th century deposits in Dublin suggests vestiges of Whitehouse (2000) argued that the decline in fire habitats pinewood or heath in the area (Reilly, 2003). Nicholls may have had a role in the extirpation and contraction in (2001) makes a convincing case based on documentary, range of some former and current invertebrates in the zoological and pollen evidence that native pinewood, British Isles. Moreover, that this raises interesting ques- particularly in small pockets in the south, survived in tions regarding the former role and frequency of natural Ireland into the post-Medieval period, but disappeared in fires within British forests, particularly within temperate prehistory elsewhere (Watts, 1985; Pilcher and Hall, 2001). coniferous forests (see Whitehouse, 2000 and Whitehouse Speight (1985) highlights the faunal effects of the disap- and Eversham, 2002 for a fuller discussion). It is worth pearance of pine from the Irish landscape, where only a noting, however, that fire-adapted species are highly mobile fraction of saproxylics characteristic of this habitat is and adapted to travelling distances in search of forest fires. found on the present Irish invertebrate list. This suggests that fires would have been highly localised Out of the 40 listed extinct species in Britain, 9 nine are and relatively uncommon, at least on an annual to decadal pinicolous (Table 1). These include *Buprestis rustica, basis, requiring mobility adaptations (Keith Alexander, *Cryptolestes corticinus,*Stagetus borealis ( ¼ pilula pers. comm., 2005). Aube),*Pissodes gyllenhali,(Whitehouse, 1993, 1997a) and *T. coerulea. The fact that four species of the pine- 3.10. Forest openness loving genus Rhyncolus have disappeared from the British Isles within the last 3000–4000 years suggests that their A recent review by Whitehouse and Smith (2004) has demise may be related to the decline of their preferred host highlighted another possible contributory factor in the (Whitehouse, 1997b), although a climatic factor may also decline in many forest invertebrates—the decline of open have been important, as discussed below. In Ireland, 4 forest ecosystems. A debate concerning the structure of pinicolous out of 15 extinct species have been recovered early Holocene forests (cf Vera, 2000; Bradshaw et al., (]H. ater, ]Hylastes angustatus, ]R. ater, mR. elongatus, 2003; Kirby, 2004) led to a review of published fossil insect Table 2), although future palaeoecological investigations data on the nature of early Holocene forest insect faunas may yet reveal much higher numbers. (Whitehouse and Smith, 2004). The authors concluded that ARTICLE IN PRESS 1770 N.J. Whitehouse / Quaternary Science Reviews 25 (2006) 1755–1789 within the limits of the published record some early mid heated by the sun. Lindroth (1945) regarded it as a relict of Holocene forest was at least partly open in character or a post-glacial warm period in Sweden. The beetle has also included open areas, whilst species usually associated with been found in several sites dating to the last interglacial the dung of grazing animals, although in small numbers, together with other southern ground beetles (Coope, 1990; were persistent. They stress, however, that the role of Keen et al., 1999) which would suggest that it is grazing animals in creating these openings is far from clear thermophilous taxon. Osborne (1969, 1976, 1982) also on the present evidence and requires new research. Ponel et attempted to examine the distribution of beetles linked to al. (2001) have drawn attention to the simultaneous habitats which are less subject to human disturbance to mixture of insects from forest, forest margins and open extrapolate climatic information. He concluded that there environment found in early Holocene samples from was some evidence to suggest that between 3000 and southern France (Lac Long Inferieur), which indicate that 2000 cal BC (5000–4000 cal BP) summer temperatures were the landscape consisted of forested areas broken by higher than the present day, declining to present levels grassland areas—could these represent areas of ‘‘pasture during the Iron Age and remaining more or less constant woodland’’? until the ‘‘Little Ice Age’’ (Osborne, 1982; Girling, 1984b). Whitehouse and Smith (2004) highlight how many forest It is possible that some locally extirpated species invertebrates require a mixed mosaic rather than full survived into the Medieval period, disappearing only with canopy forest—many species appear to require sunny, the onset of cooling of the ‘‘Little Ice Age’’ (Grove, 1988) open habitats with large amounts of dead wood (Palm, although the effects of this climatic deterioration have been 1959; Alexander, 1999; Ranius and Jansson, 2000). hard to identify and evaluate (see Buckland, 1975; Buck- Of particular interest is a recent discussion that several land et al., 1983; Girling, 1984a; Osborne, 1997; Wagner, pine and birch-dependent beetle species connected to 1997). Buckland and Wagner (2001) suggest that this open forests have become extinct in southern Sweden episode may well have had an impact upon insect faunas, (Lindbladh et al., 2003). The fossil beetle record indicates but draw attention to the fact that very few deposits such faunal changes are not new and suggests that we may spanning this period have been investigated in which a need to consider the possibility that the open structure of clear climatic signal is evident. Several beetles have been many of these Holocene forests may have been equally put forward as possible casualties of the Little Ice Age—the crucial to some of these species’ survival (Whitehouse and non-British whirligig beetle *Gyrinus colymbus (Girling, Smith, 2004). 1984a), a species which lives in standing and flowing water (Koch, 1989) and the extirpated flat bark beetle *Airaphilus 3.11. The role of climate change in extirpations elongatus (Osborne, 1974), which lives in swampy meadows and other wet places with plenty of wet vegetation (Koch, Despite the success in examining Lateglacial climatic 1989). Dinnin (1991) has suggested that the Little Ice Age change (cf Atkinson et al., 1989; Coope and Lemdahl, may have been a contributory factor in the decline of the 1995; Coope et al., 1998) Holocene climatic change has RDB 3 Greater Silver water beetle (Dytiscidae), Hydrous been less easy to infer from the coleopteran record. ( ¼ Hydrophilus of Kloet and Hincks, 1977) piceus, Terrestrial floral evidence indicates a cool start to the reducing its habitat range, while subsequent re-colonisation Holocene (Birks, 1989), but the beetle evidence suggests was prevented by the destruction and fragmentation of that this period may have been the warmest (Osborne, suitable habitats. This species has a very limited, southern 1997). Osborne (1974) inferred temperatures of 16–17 1C distribution in the British Isles and is no longer found as (about the same as today) from a fauna recovered from Lea far north as the Humberhead Levels. Its fossil record, Marston, in the English Midlands dated to the first 500 however, would suggest that it was considerably more years of the Holocene, whilst Coope (1998) has provided widespread in the past (cf Dinnin, 1991). Moreover, estimates ranging between 15 and 24 1C by about 9000 BP nineteenth century records from Derbyshire and Hudders- in southern England, about 2–31 warmer than today. field (Balfour-Browne, 1958), testify to the presence of As we have seen above, from the mid-Holocene post-Little Ice Age populations in more northerly latitudes, onwards, the records indicate that human impact is of suggesting that pollution and drainage are more likely to considerable importance, and that this may swamp and explain this species’ status. mask low magnitude climatic events. Some researchers It is perhaps useful to look more carefully at some of have suggested that there is evidence for a warmer climate these extirpated species’ modern distributions. Figs. 4–8 during the Bronze Age (Osborne, 1972a, 1988; Girling, provide an indication of the present distribution of species, 1984b) on the basis of the present distribution of some as well as fossil localities. Information concerning many of extirpated species. Girling argued that the recovery of the these species is generally rather incomplete (see Table in the non-British ground beetle (Carabidae) *Oodes gracilis Vil. online version of this article), and the lack of records from in Bronze Age deposits above the Sweet Track, Somerset, an area does not necessarily imply a species’ absence. In England (Girling, 1979) indicated a warmer climate. This many cases, the data are dependant upon records from species has a distinctly southerly distribution (see Coope, the last century, and are positively correlated with those 1990) and lives in standing eutrophic waters strongly areas most studied by entomologists (e.g. Fennoscandia, ARTICLE IN PRESS N.J. Whitehouse / Quaternary Science Reviews 25 (2006) 1755–1789 1771

Fig. 4. Modern distribution of (a) Isorhipis melasoides, (b) Tenebrioides fuscus, (c) Pcynomerus tenebrans, (d) Rhyncolus punctulatus. Stippled areas indicate areas where records are rare. ARTICLE IN PRESS 1772 N.J. Whitehouse / Quaternary Science Reviews 25 (2006) 1755–1789

Fig. 5. Modern distribution of (a) Buprestis rustica, (b) Pissodes gyllenhali, (c) Cryptolestes corticinus, (d) Prostomis mandibularis. Stippled areas indicate areas where records are rare. ARTICLE IN PRESS N.J. Whitehouse / Quaternary Science Reviews 25 (2006) 1755–1789 1773

Fig. 6. Modern distribution of (a) Bothrideres contractus, (b) Mycetina cruciata, (c) Stagetus borealis, (d) Rhyncolus sculpturatus Walt. Stippled areas indicate areas where records are rare. ARTICLE IN PRESS 1774 N.J. Whitehouse / Quaternary Science Reviews 25 (2006) 1755–1789

Fig. 7. Modern distribution of (a) Rhyncolus strangulatus, (b) Rhyncolus elongatus. Stippled areas indicate areas where records are rare.

Germany, France and to a lesser extent ). In many Thorne and Hatfield Moors and today found as far north cases, the biogeographical patterns of many saproxylic as northern Fennoscandia (Figs. 5a–c). Other species such invertebrates are not well-known. as mProstomis mandibularis, mB. contractus,*Mycetina Species on the edge of their range are more susceptible to cruciata,*Stagetus borealis and mR. sculpturatus tend to be edaphic and anthropogenic disturbance, as well as climatic distributed within central and southern Europe (Figs. 5d, change (Warren and Key, 1991). The distributions of the 6a–d), extending only into southerly localities of Fennos- Urwaldtiere beetles could imply a more continental climate, candia. Osborne (1972) highlighted the occurrence of and climatic deterioration may account for their subse- several thermophilous beetles in early Holocene deposits quent disappearance. Examining the distribution maps of from Worldsend, Shropshire. *Porthmidius austriacus and the locally extirpated species, several beetles appear to have *R. strangulatus (Fig. 7a) are no longer found in Britain or thermophilous requirements. This is best illustrated for *I. Fennoscandia, having moved southwards, whilst mR. melasoides (Fig. 4a). Horion (1953) suggests that it is elongatus (Fig. 7b) is found no further north than the restricted to areas away from the western seaboard and south of Sweden. Mediterranean, and that the species has a discontinuous *T. coerulea, which was recovered from Thorne Moors east–west distribution. It is absent from Scandinavia. Its from two localities (Roper, 1996; Whitehouse, 1997a), distribution suggests that it is relatively thermophilous, belongs to a family which is predominantly thermophilic although its absence from areas of central seems and shows a current southern European distribution to indicate that old established forest is a necessary (Whitehouse et al., 1997)(Fig. 8). Its occurrence on the component of this species’ ecology (Buckland, 1979). A island of Go¨ska Sa¨ndon, off the eastern coast of Sweden, in range of species from Thorne and Hatfield Moors (White- old forest, however, suggests a wider tolerance of house, 1998, 2004) all have distinctly southerly distribu- temperatures. In Fennoscandia, it is considered a relic of tions, south of the 17 1C July isotherm, including the warmer post-glacial climatic period (Juha Siitonen pers. *Tenebrioides fuscus, *Pycnomerus tenebrans,*Rhyncolus comm., 1997). Moreover, this species was recovered in punctulatus (Figs. 4b–d). This is in contrast to the pine- the same deposit as the bark-beetle Scolytus ratzeburgi, loving *Buprestis rustica *Pissodes gyllenhali and *Crypto- which is today confined to the uplands of Scotland and lestes corticinus all of which have also been found at areas of Fennoscandia (Whitehouse et al., 1997)(Fig. 8). ARTICLE IN PRESS N.J. Whitehouse / Quaternary Science Reviews 25 (2006) 1755–1789 1775

? Scolytus ratzeburgi ?

?

? THORNE ? ? ? ? ? ? ? ? ? ? ? 17C July ? ? isotherm ? Temnochila coerulea ? ?

? ? ?

Canaries W.Africa 0 1000 km W.Asia

0 800 mls

Fig. 8. Modern distribution of Temnochila coerulea and Scolytus ratzeburgi. Re-drawn with permission after an original by P.I. Buckland.r P.I. Buckland.

Coope (1998) recovered this species from early and later more continental and drier climate during the Holocene or Lateglacial Interstadial deposits from Holywell Coombe, are they merely a reflection of the differing histories of suggesting that it can tolerate a range of both warmer and woodland and forest in the west and Baltic regions cooler climatic conditions. Its fossil records from the earlier (Whitehouse et al., 1997)? Holocene are from sites no further north than Thorne The behaviour of some of the members of the weevil Moors, which suggest that it used to live much further genus Rhyncolus is informative. Members of this genus live south in the past than it does today, although the general in moist, rotten wood of both standing and fallen trees paucity of investigations in Scotland and northern England (Palm, 1959). They invade wood at a rather late decaying may explain this apparent contradiction. Whitehouse et al. stage and live in it for several generations. All European (1997) have argued that this species’ present distribution members of the genus with the exception of *R. reflexus may be a reflection of availability of habitat (birch forests, have been recovered from British and Irish fossil deposits, mostly now confined to upland areas) as well as a including *R. strangulatus (Osborne, 1972). The fact that preference for warmer, drier conditions. Considering the four species of the same genus have disappeared from the two species together (Fig. 8), do the distributions reflect a British Isles within the last 3000–4000 years has been ARTICLE IN PRESS 1776 N.J. Whitehouse / Quaternary Science Reviews 25 (2006) 1755–1789 attributed, at least in part, to the demise of its preferred the British fauna during the retreat of pine forest towards the host, pine (Whitehouse, 1997b). Today, only one member north; evidence from the fossil record and elsewhere on this of this genus still lives in the British Isles, R. ater, mostly in point is insufficient’’. The research presented above would the Scottish Highlands (Alexander, 2002), although Carr seem to suggest that we may have indeed lost several (1916) collected it in Sherwood Forest suggesting that it warmth-adapted pine-loving beetles. had survived in England into the 20th century. The species In the fossil record, the weevil Dryophthorus corticalis clearly used to be more widespread and is one of the (RDB 1) is closely associated with the genus Rhyncolus. commonest pine-loving species found as a fossil in forest The former species was added in 1925 to the British assemblages, ranging from the south of England (e.g. invertebrate list by Donisthorpe (1939), who discovered it Somerset, Girling, 1980), through to south and north in Windsor Forest in damp, tough wood inside old oak. Yorkshire (Buckland, 1979; Hall and Kenward, 1990; Today it is exclusively associated with oak. It is an Roper, 1996; Whitehouse, 1997a, 1998). In Ireland, it is not endangered species in Britain, endemic to Windsor Forest listed amongst the native species (Anderson et al., 1997), and Great Park, its endemicity is determined by the lack of but has been recovered from Sluggan Bog, Derryville suitable habitats elsewhere and presumably by problems of Bog—from an oak rot hole—(Caseldine et al., 2001) and mobility between patches of forest. Its fossil record Medieval Dublin (Reilly, 2003). Given its common indicates that it used to extend at least as far north as occurrence in a wide variety of deposits of different ages, Cumbria (Hughes et al., 2000), suggesting that conditions its current status both in Britain and elsewhere in Europe, were sufficiently good for it to extend its range. Its where it is not regarded as particularly rare or endangered, distribution in Europe is restricted to central Europe and nor a characteristic inhabitant of undisturbed forest, and it does not extend northwards into Fennoscandia, suggest- the fact that it can be recovered unassociated with other ing it is relatively thermophilous. Urwaldrelikt taxa, it is most likely that this species’ As a fossil, the species has been found frequently disappearance from both Irish and British forests is associated with the genus Rhyncolus, often in pine rot probably related to the decline of its host—pine—(cf holes (e.g. Koponen and Nuorteva, 1973; Lekander et al., Speight, 1985) rather than any strong requirement for 1975; Boswijk and Whitehouse, 2002). Palm (1959) Urwald forest. describes it as attacking both oak and pine, where the But why has this species survived in the British fauna wood is still hard and firm. It is possible that with the over the other members of the genus? There is some decline of pine habitats in Britain the species has been suggestion that R. ater may tolerate a broader range of constrained to live in oak, although pine habitats are still temperatures than the other members of the genus. available in southern England in areas of heath. If this is Zetterstedt (1828) includes the species in his Fauna the case, why is it not found in the Caledonian pine forest? Lapponica (fauna from Lapland) but excludes the other The fossil is frequently found associated with R. ater which members, whilst Lindroth (1948) suggests that it is spread is able to live in these northern forests, but apparently this throughout Fennoscandia, apart from the northernmost habitat is not suitable for D. corticalis. This ‘‘splitting’’ of peninsulas, and extends across the whole of Europe apart former close associates suggests that whilst R. ater was able from the southern most parts. Other members of the genus to inhabit the northern forests, its associate was not. This appear to be more temperature sensitive, as indicated by could be interpreted in terms of thermal requirements by D. their distribution maps. Species such as R. ater,*R. corticalis, as implied by its modern distribution. If oak is its punctulatus and *R. sculpturatus are not wholly restricted preferred host, then its fossil presence in pine (like the to conifers and also live in deciduous trees in more presence of R. ater in oak) could indicate that it was southern localities (Palm, 1951, 1959). R. ater is found previously not as ‘‘fussy’’ about its host because it was not exclusively on pine in the British Isles today but its at its thermal limits. However, today it is far more host- presence in oak rot holes in both English (e.g. Boswijk and specific and has contracted southwards. This could imply a Whitehouse, 2002) and Irish fossil contexts (e.g. Caseldine contemporary climate warmer than today, with subsequent et al., 2001) indicates that it was rather less fussy about its minor climatic changes causing species to become increas- host during the mid-Holocene—perhaps because it was not ingly confined to pine, in the case of R. ater, or oak in the at the limits of its range. At some stage, however, it seems case of D. corticalis. to have become more host specific, forcing it to become If there had been a change from a more continental to a largely restricted to the Caledonian pinewoods. The failure more oceanic climate, this would have resulted in less of other species to survive in the cooler northern pine successful over-winter hibernation of some beetles. Ander- forest, however, suggests that they were not as cold- son et al. (2000) have pointed out that many saproxylics are tolerant as R. ater and/or they were adversely affected by distinctly thermophilic in that they require settled, warm habitat destruction and fragmentation. Indeed, Hammond conditions for feeding and dispersal and are particularly at (1974, p. 327) drew attention to the fact that the restriction risk in a variable, windy climate. It is easy to see how a of pine forest to northern Britain produced a reduction in change from more settled, drier conditions to cooler more pinicolous species and commented that, ‘‘It is possible that oceanic conditions is likely to have adversely affected some such species, if at all warmth-adapted, were lost from such taxa. Many boreal saproxylics are thermophilous ARTICLE IN PRESS N.J. Whitehouse / Quaternary Science Reviews 25 (2006) 1755–1789 1777

(Anderson et al., 2000), a point elegantly highlighted by coming from beetles representing a variety of threatened Ponel et al. (2003) in their discussion of Eemian faunas habitats, including those associated with rapidly disappear- (130,000–110,000 BP) from La Grade Pile (France), where ing wetlands and meadowland. they found that coniferous-associated taxa were present in However, certain groups appear to have suffered greater warm or temperate mixed forest, but absent from cold declines over others—namely those associated with boreal conifer forests, where cold-resistant taxa are dominant. forest and mire woodland habitats. In a recent review, The loss of niche and micro-habitats through forest Smith and Whitehouse (2005) suggest that not all forest clearance, however, could have had similar effects to those types appear to have contained such a high proportion of described above. Saproxylics in the interior of large logs or locally extinct and rare species. They suggest that mire cavities in trees are buffered from fluctuations in moisture woodlands, in particular, may have suffered great losses content and temperature; the presence of such micro- partly because they are largely atypical environments climates within forests is key to the survival of many today, at least in the British Isles. The highest number of species (Kaila et al., 1997). The removal of trees and wood extirpations is evident amongst the pinicolous species. through human activities would have had the effect of Interestingly, there are more extirpations in this category subjecting many of these saproxylics to increased tempera- compared with numbers of rare and endangered species, ture fluctuations. This buffered habitat is particularly suggesting that these extinct species may have been subject important at the northern edges of a species’ range, where to a greater magnitude of limiting factors compared with climate may be an important limiting factor to the survival their more robust counterparts—niche or perhaps climate of a population (Warren and Key, 1991). The removal of related. With the history of pine in Ireland and Britain the insulation provided by the forest would have had an during the Holocene (cf Bennett, 1984, 1995), it is not adverse effect upon a population, without any actual surprising this has had a detrimental effect on the change in macroclimate. Many species may thus have had associated insect fauna. However, the numbers of extirpa- problems maintaining viable populations. tions associated with this tree suggests its associated beetles The disjunct modern distribution of some of the extirpated have suffered more than any other group of tree associates species indicates that their survival may largely be a matter of (Smith and Whitehouse, 2005). chance, surviving in woodland sites with a continuity of With the northward restriction of pine forests, at least a suitable habitats. With a history of forest clearance lasting proportion of their inhabitants appear not to have found several thousand years, it is not surprising that numbers of suitable habitats. This suggests that at least some of these specialised insects have become extinct. Although climate species are temperature sensitive, particularly if they were change, particularly a shift from drier and warmer to cooler already at the northern limits of their range. Temperature and wetter conditions may have played a part in the decline affects the length of larval development in some saproxylics and extirpation of saproxylics, it remains only one factor, and persistent cold winds can limit the dispersal of amongst several, which appears to have determined the emergent beetles (Bistro¨m and Va¨isa¨nen, 1988; Szujecki, modern distribution of many of these species. 1987). Additionally, other types of habitats seem to have been important for some species’ survival, such as the fire 3.12. Quantifying extirpations in Britain and Ireland ecosystem and the open forest structure of some of these forest types. Abundant dead wood would have been an The few studies that have examined ‘‘natural’’ succes- important attribute of these ecosystems, whilst temporal sions indicate that it is still really too early to quantify the continuity of habitat probably was a significant component number of extirpations in Britain and Ireland. To date, 40 in the development and maintenance of many of these pre-Linnaean (i.e. not including recent 18th–19th century saproxylic communities. extirpations) extirpations have been recorded amongst Compared with work carried out at some British sites, British Holocene Coleoptera (Table 1), whilst 15 have been the fossil insect faunal list from Ireland is somewhat recorded in Irish deposits, of which four have their nearest disappointing, lacking the plethora of ‘‘European’’ species living relatives in the forests of Central Europe (Table 2). (Table 2). Is this because these elements are lacking in the Assessing true numbers is compounded by problems Irish fauna or is it related to the fact that targeted sampling associated with the identification of species that cannot to recover these taxa has rarely been untaken? The few be recognised on characteristics recoverable from the fossil investigations carried out have been associated with record (i.e. not identifiable on head, thoraces or elytra). archaeological sites and structures (e.g. Reilly, 1996, There are also some difficulties related to the identification 2003). The low number of investigations of associated of specialist species for which modern comparative speci- palaeoecological deposits could easily explain the paucity mens are increasingly difficult to track down. All these of the present record. Evidence from Sluggan Bog and factors would suggest that these identified ‘‘extirpated’’ Ballymacombs More (Whitehouse, in press) suggests this fossils represent a tiny proportion of the range of species indeed may be the case and highlights that a range of which formerly lived in Ireland and Britain. Of the natural palaeoecological deposits need to be investigated to complete list, over 60% of species (25) are from saproxylic throw light upon the full range of Ireland’s ancient taxa associated with old and dead wood, with the rest woodland inhabitants. ARTICLE IN PRESS 1778 N.J. Whitehouse / Quaternary Science Reviews 25 (2006) 1755–1789

Unlike Britain, Ireland also appears to have lost a suite As Anderson et al. (2000) point out, this period of the of less specialised species, such as some of the wood boring last 500–1000 years is likely to have been when Ireland’s bark beetles, cossinine and long-horned beetles, Urwaldtiere suffered their greatest crisis and it is not sometime over the last 700–800 years. Many of these difficult to see why the Irish flora and fauna is so seriously species are not particularly rare in Britain. Two possibilities depleted. It seems likely that not only did any remaining are likely to explain these distinctions. Firstly, there is the specialised Urwaldtiere disappear—although many of these possibility that some of these species could have thermo- may have already gone during prehistory—but that a suite philous requirements. Many of the 13th century AD of far less specialised species also vanished, almost certainly records for these taxa occur during the Medieval Warm due to the final stripping of the Irish landscape. In Britain, period and their presence within deposits of this date could the establishment of Royal hunting forests and Chases be explained by a warmer climate during this time, with protected areas for hunting (Rackham, 1986) and probably their subsequent disappearance related to the onset of the saved equivalent species. The Irish faunal record provides a Little Ice Age. However, their continued presence in salutary reminder of what can happen to biodiversity when Britain suggests this is unlikely. extreme human impact takes place. Moreover, the extirpa- Irish forest history perhaps provides some answers. tion of native pine in Ireland exacerbated many of these By the Medieval period, forest had largely disappeared saproxylic losses (Speight, 1985). Scotland still provides a from the Central Plain of Ireland, but large stands of oak refugium for the Caledonian pine fauna, but this has long forest remained in the south-west and north-east since gone from Ireland, although its apparent survival in (McCracken, 1971). Documentary and cartographic evi- pockets of western Ireland into the post-Medieval period dence suggests there was still good quality timber available (Nicholls, 2001) may indicate that the loss of this fauna in Ireland into the 16th and 17th centuries AD, with may be a comparatively recent phenomenon. extensive woodlands in some regions (Nicholls, 2001). This Analysis of the modern Irish saproxylic fauna suggests would explain the occurrence of saproxylics surviving in that it is also impoverished of thermophilic (warmth- deposits from Medieval Dublin (cf Reilly, 2003). The loving) families or groups of species (Anderson et al., Tudor conquest of Ireland and the arrival of English 2000), suggesting that climatic conditions may also explain settlers, however, led to a period of intensive and the nature of the modern Irish fauna. The European species unprecedented exploitation of Irish woodlands, with little recovered in the Irish fossil record—mProstomis mandibu- or no regard for their survival, as timber was used for quick laris, mB. contractus, mR. elongatus and mR. sculptur- profit. By the 1670s, good timber was only available in atus—are all taxa seldom found north of southern inaccessible locations and most woods were exhausted Fennoscandia and it seems likely that climate change (Nicholls, 2001). Any remaining areas were decimated could at least partially explain their demise in Ireland. by the use of wood in the iron industry. The speed at This review provides an opportunity to visit two issues which woodland was exploited and the ‘‘colonial’’ nature which have been generally poorly considered within the of the stripping of the landscape must have had a wider literature, at least in recent times, but where the fossil devastating impact on its flora and fauna. By the 19th beetle record may provide important insights: the colonisa- century the drastic effects of deforestation were obvious, as tion of forest biota into Ireland and the European origins a French visitor commented in 1887, ‘‘The most striking of the British and Irish forest fauna. thing on first sight of the Irish landscape is the total absence of trees of any kind. They are seen only in private parks’’ 3.13. The colonisation of saproxylics into Ireland (Neeson, 1997, p. 142). Palynological investigations associated with the historic The modern Irish beetle fauna is significantly impover- period, however, indicate that the extent and nature of this ished of its saproxylic elements (ca 600 species versus ca woodland may have been somewhat over-estimated by 1800 species in Britain, Alexander, 2002), even when English colonisers, who wrote many of the historical compared with the British fauna, which is itself a sub-set records, not least because this allowed them to explain of the European fauna (Anderson et al., 1997, 2000). Many the severe problems they were experiencing in attempting old forest taxa are considered to have very limited powers to control Ireland (Pilcher and Hall, 2001). The pollen of dispersal and mobility (Warren and Key, 1991). None record shows that over the last 1000 years the Irish survived the cold of the last glacial and there is, at present, landscape was more diverse than documentary records no evidence that any arrived during the Lateglacial imply, with considerable fluctuations in woodland cover Interstadial (e.g. Coope, 1971; Coope et al., 1979; White- (Hall, 1995). Indeed, palynological work suggests that the house, in prearation), despite suggestions of survival in real damage was done much earlier, in the pre-Norman refugia (e.g. Praeger, 1932). Praeger (1932) considered that early Medieval period (Hall, 1995, 2000) but that the the upland heath Lusitanian flora and fauna found in parts remaining scrublands and much lighter woodland that of south-western Ireland were part of the longer established dominated the landscape during the historic period biota, possibly surviving in ‘‘nunataks’’, ice-free areas in (Hall, 1998) were considerably affected by 17th century the landscape. The Lusitanian (also known as Hiberno- clearances. Cantabrian) flora and fauna consist of species not found in ARTICLE IN PRESS N.J. Whitehouse / Quaternary Science Reviews 25 (2006) 1755–1789 1779 any other part of the British Isles, whose nearest relatives of insects (Sadler, 1990, 1999), although these were mostly are normally found in and Portugal (Pilcher and synanthropic rather than forest taxa. The last option— Hall, 2001). wood rafting on driftwood—is likely to have been, Since saproxylic species require continuous corridors of potentially, one of the more successful mechanisms for mature forest habitat it is clear that they must have arrived biotic movement. in Ireland during the early Holocene, when their tree hosts The dispersal of biota by driftwood has been discussed would have found favourable conditions. If a land-bridge by several biogeographers (e.g. Johansen and Hytteborn, existed during the early Holocene this could have 2001), including in the consideration of insect movement. facilitated dispersal of forest invertebrates. Mitchell Dendrochronological dating of drifted wood indicates the (1963) argued for a Holocene land-bridge between Wick- considerable long-range dispersal potential of driftwood low and the lleyn Peninsula, Wales, and between Louth, (Johansen and Hytteborn, 2001). Maximum floatability is the Isle of Man and Cumbria. The discovery of the snail found amongst coniferous woods, which can float 10–17 Discus rotundatus Muller, which shows a broadly synchro- months. The potential for transporting seeds and fruit nous pattern of arrival in both Britain and Ireland during within soil amongst the roots of driftwood trees is the early post-glacial led Preece et al. (1986) to also favour considerable. Transport of biota in this manner would a Holocene land-bridge. have greater protection from sea water and it is easy to see Unfortunately, this is not borne out by the geological that invertebrate ‘‘hitch-hikers’’ would have been protected evidence. Unlike Britain, which did not become separated within this litter layer and the thick, layered bark of from the European landmass until about ca 5000 cal BC (ca coniferous trees (Johansen and Hytteborn, 2001). Insect 7000 cal BP) when the North Sea flooded due to sea level species commonly found in driftwood include cossinine rise (Shennan et al., 2000), Ireland seems to have been weevils such as Mesites tardii (Morris, 2002) and members isolated for considerably longer. Much of the evidence of the genus Rhyncolus (Skidmore pers. comm., 2000), suggests it became an island sometime between 18,000 and precisely the sorts of taxa under consideration. 13,000 BP. Devoy (1995) argues that any land-bridge Early Holocene forests contained abundant dead wood, would have existed as a discontinuous area of temporary which are likely to have regularly fallen into rivers flowing islands between the north of Ireland and south-west outwards towards the Irish Sea, especially during periods Scotland ca 11,400–10,200 BP. Wigfield (1995) on the other of flooding. Source areas for biota rafting with driftwood hand, suggested that a series of land-bridges were formed trees would be from within British forests, but also from by a forebulge in front of the ice sheet, providing migrating the European mainland. Movement of taxa on driftwood is land connections between 11,000 BP and 9750 BP. Devoy likely to explain the arrival of some species, particularly (1995) considered this hypothesis lacking in reliable data. where there is evidence for synchronous arrival in Britain More recently, Lambeck (1996) proposed a land-bridge and Ireland (e.g. Preece et al., 1986) and may have included between 18,000 and 14,000 BP in the Bristol Channel area, a range of invertebrates and plants. disappearing by 13,000 BP. This data suggests that the Irish land-bridge had gone long before warming of the 3.14. European origins of the British and Irish forest fauna Holocene. Moreover, the tidal amplitude of the Celtic Sea would have meant that the exposed sea floor was Far less consideration has been directed to the issue of subject to tidal flooding—a hostile environment for any the Continental origins of the British and Irish forest flora or fauna attempting a crossing, let alone the entomofauna and mechanisms of insect dispersal towards flourishing of mature corridors of forest required by the end of the last ice age from glacial refugia. In the saproxylic taxa! The apparent absence of a land-bridge following section, a consideration of the origins of connecting Ireland to sources of forest during the early saproxylics’ hosts is given before turning to the saproxylics Holocene suggests that many specialist saproxylic species themselves. simply would not have arrived. The fact that species did Looking at the distribution maps of some extirpated arrive raises interesting biogeographic questions concern- species it would appear that two biogeographical elements ing the mechanisms behind insect colonisation during the are represented: those whose current distribution is south early Holocene. of the 17 1C July isotherm and which display a distinctly There are three other obvious sources of transport for southern distribution (e.g. Fig. 4) and those which appear these species: wind, humans and wood rafting. Wind could to have a greater affinity with Fennoscandian faunas have transported some of the winged taxa such as bark (Fig. 5a–c). Keith Alexander (2002) draws attention to the beetles, but it is unlikely to have had much influence on presence of two major components within the modern Irish highly specialised Urwaldtiere, since many are wingless and and British saproxylic fauna—an Atlantic version of the rarely move outside forest. Human activities may well have temperate broad-leaved forest fauna and a boreal forest accidentally introduced several components of the forest fauna. The Atlantic fauna is distributed throughout low- assemblage. Palaeoecological work in the North Atlantic land Britain, extending as far as northern Scotland. The islands (e.g. Iceland, Faro¨es) shows the success with which northern parts of Scotland have a boreal forest with a human activities can introduce viable breeding populations fauna which has greater similarities to Fennoscandia. ARTICLE IN PRESS 1780 N.J. Whitehouse / Quaternary Science Reviews 25 (2006) 1755–1789

Many species are common to both categories, but some ice sheet than previously anticipated. One suggestion is that species are particularly characteristic of the temperate or ice-free areas of coastal may have acted as boreal forest. In Ireland, the fauna is a hybrid between the immigration sources for the trees in the Scandes, with temperate and boreal forests of Britain. colonisation coming from the west rather than the east The fossil beetle record indicates that it is the boreal (Kullman, 2002). On the other hand, the existence of a fauna which appears to have suffered a particular decline in possible refugium to the north-east is highlighted by several species, although the Atlantic fauna has not remained other researchers (Taberlet et al., 1998; Sinclair et al., immune. It is worth noting that the closest comparable 1999). Whether originating from east or west, Sinclair et al. fossil insect records to the mire woodlands of Thorne and (1999), investigating the genetic make-up of Fennoscan- Hatfield Moors, come not from within Britain, but dian pine populations, found two different origins for the from Piilonsuo, a pine-mire in Finland (Karppinen and trees in this region. In southern Sweden, have genetic Koponen, 1973, 1974; Koponen and Nuorteva, 1973). markers similar to populations from central Europe, thus Many of the trees upon which these populations originating from refugia to the south, whilst pines from ultimately depend originated from refugia in southern northern Sweden, Norway and Finland are genetically Europe (Bennett et al., 1991; Hewitt, 1996; Willis, 1996). different, which they suggest may have originated from Recent research based on pollen evidence and more latterly populations in the north-east, via Finland (Sinclair et al., macrofossil charcoal from Upper Palaeolithic archaeolo- 1999). Taberlet et al. (1998) examining the genetic diversity gical sites (Willis et al., 2000; Willis and Niklas, 2004; Willis of a range of insects, mammals and tree species across and van Andel, 2004) suggests that there was full glacial Europe provide support for the different origins of the growth of trees in central and eastern Europe. There is Fennoscandian floral/faunal populations. They draw at- evidence for the survival of some tree populations in more tention to the existence of a ‘‘suture’’ zone in Fennoscan- northerly refugia, particularly those associated with con- dia, suggesting that this area may have been colonised by iferous forests, as far north as southern Poland (Stewart populations from refugia both in the south and east. and Lister, 2001; Willis and Niklas, 2004). Moreover, it Moreover, they highlight that few data are currently seems that coniferous trees were present continuously available for north-eastern Europe and Fennoscandia and between 35,000–20,000 BP, the period for which there is that we may be under-estimating the contribution of adequate data. Numbers declined towards the last glacial potential eastern refugia located in Europe and/or Asia maximum, but in and Moldavia milder condi- to this area of northern Europe. tions during this period may have allowed the continued In the British Isles, DNA evidence from Scots pine survival of trees in this region of Eastern Europe (Willis indicates that north-west Scottish pine is distinctly different and van Andel, 2004). Whether the trees grew in isolated from other British and continental populations (Sinclair et pockets or an otherwise open tundra landscape or open al., 1999) and may have originated from what Stewart and taiga forest is much more difficult to evaluate (Willis and Lister (2001) call a ‘‘cryptic’’ refugium. Birks (1989) van Andel, 2004). indicates that pine was present during the first thousand Thus, boreal species, including pine, seem to have years of the Holocene in this area. She raises the possibility survived substantially further north than originally antici- that it could have been growing in glacial refugia during pated—not necessarily isolated—whilst deciduous trees the last glacial, although she considers this unlikely in view may have remained isolated in southern refugia (Willis of the ice sheet extent across Scotland. Another possibility and Whittaker, 2000; Willis and Niklas, 2004). These is that seeds could have originated from across the North different refugial patterns have important implications for Sea basin and northern Europe (Birks, 1989). At this stage, the genetic make-up and re-colonisation of different groups sea levels would have been lower than today and England of plants and their associated organisms at the end of the would have been connected to the European Continent, ice age (Willis and Niklas, 2004). Genetic studies of trees effectively creating a continual landmass between Great suggest that the interplay of colonising populations from Britain and northern Europe. There has always been the these different refugia created high genetic tree diversity in implicit assumption that the origin of the flora and fauna central Europe (Petit et al., 2003). Boreal species, however, of the British Isles lies southwards, towards central Europe do not conform well to this overall pattern, which may be and the traditional southern refugia (cf Hewitt, 1996). due to their survival in more northerly refugia and in However, the existence of at least two different Fennos- diffuse populations during ice ages (Petit et al., 2003; Willis candian populations and the chance of possible refugia and van Andel, 2004), suggesting that post-glacial re- further north than previously suggested, either in Fennos- colonisation of organisms may be far more complex than candia itself (cf Kullman, 2002), or more likely originating previously thought (Willis et al., 2000). further east and expanding via Fennoscandia during the Kullman (2002) draws attention to boreal tree macro- early stages of warming (cf Taberlet et al., 1998), suggests fossils (mostly pine and birch) revealed by retreating we should also consider the possibility that Fennoscandia modern ice cover in the southern Swedish Scandes, may have been a potential seed source for some pine in indicating pockets of trees of Lateglacial age living closer north-west Scotland. It is worth pointing out that the coast to the southern and western margins of the Fennoscandian of Norway is not so distant to Scotland and a number of ARTICLE IN PRESS N.J. Whitehouse / Quaternary Science Reviews 25 (2006) 1755–1789 1781 agents, including birds, water or ice could have facilitated material and fossil specimens. One possible explanation seed transportation to Scotland. for these observations is the existence of different popula- A further pine population in south-west Scotland may tions of this species. Whitehead (1972) reports similar have been derived from Ireland (Sinclair et al., 1999; differences in eye curvature within individual populations Stewart and Lister, 2001), from a postulated refugium in in a taxonomic study of the ground beetle genus the west (Bennett, 1984), although there is no convincing Schizogenius in northern and middle America. Hammond palynological support for such a hypothesis (e.g. Mighall (1974) draws attention to the existence of morphological and Lageard, 1999). As several other researchers have ‘‘peculiarities’’ amongst British Coleoptera, particularly in pointed out, there is thus a possibility that a refugium for northerly relict areas, leading coleopterists to suggest that Scots pine existed within the British Isles (Bennett, 1984, certain British forms are distinct populations, brought 1995; Birks, 1989) and a seed source may also have been about by isolation. The question is whether this is a available from Fennoscandia and central Europe. By the reflection of isolation of these particular populations from Lateglacial Interstadial, trees started moving northwards, Thorne and Hatfield Moors during the Holocene or accompanied by the swift arrival of accompanying whether this reflects the existence of different European saproxylics during the early Holocene—particularly those populations of this species, brought about by isolation in associated with the boreal forest. different glacial refugia. Comparative work between a All this has important implications for the refugial range of modern and fossil beetles would be a valuable origins of many British and Irish saproxylics, particularly exercise to further explore these observations, accompanied those associated with pine and boreal forest. It suggests by modern DNA analysis. that this fauna could have originated from several different Evidence for evolutionary change is extremely rare from refugia, including northern and central Europe, and Quaternary insect assemblages (see Matthews, 1970; potentially, Fennoscandia and Ireland. Whilst the role, if Bo¨cher, 1986, 1997). Coope (1970, 1978, 1995, 2004) any, of these latter two areas may well need considerable suggested that insects had remained stable in their more research, it does indicate that the genetic structure of morphology and environmental requirements throughout our saproxylics, like that of their hosts, should reflect the whole of the Quaternary period, responding to climate genetic diversity resulting from different refugia. One change by undergoing distributional shifts. The effect of important consideration in this respect is the fact such movement would keep the gene pool constantly that whilst seeds may have found it relatively easy to mixed, preventing genetic isolation of populations move from refugia, transported by wind, water and other and mutations which would result in speciation and agents (Bennett, 1984), the same agents may not have ensuring that stasis is the norm for many Quaternary been as suitable for insect transportation, particularly insects. However, as Ashworth (2004) points out, stasis is for saproxylics. This suggests that the genetic structure not always corroborated by genetic evidence (e.g. Reiss and origins of saproxylic beetle species does not et al., 1999). He argued that this may possibly be true necessarily have to be the same as their hosts and indicates only for the northern temperate fauna, while the highly an important avenue for further research. Possible diverse tropics may be the place to look for evidence mechanisms of dispersal discussed above, including wood for divergence, where, as Coope (2004) highlights, rafting, could have all been possible from the postulated climatic oscillations may not have been so extreme. refugia. Moreover, this perspective may not hold true for As Stewart and Lister (2001) highlight, the existence of narrow-range endemic taxa. Recent DNA work indicates possible different isolated refugia has an implication for that some Iberian endemic water beetle species (Dytiscidae) speciation. Of particular interest is possible evidence for are of middle to early Pleistocene age, providing evidence population-level distinctions apparent amongst Holocene for in situ Pleistocene speciation (Ribera and Vogler, 2004). forest faunas. Whitehouse (1998) noted morphological Ribera and Vogler (2004) suggest that the northern differences between fossil specimens of R. elongatus from geographical bias of European fossil beetle investigations Thorne and Hatfield Moors and comparative modern may explain the lack of evidence for speciation amongst specimens, all of which had been collected in central Quaternary insect faunas, where climatic changes Europe. No fossils were compared with Fennoscandian forced large-scale movement of faunas. They argue material as this was not available at the time. Morpholo- that stasis may not be applicable to narrow-range endemics gical differences concerned the curvature of the eyes of and those associated with postulated refugia. Investiga- the fossils, which were significantly flatter compared to tions covering 110,000 years of beetle diversity at La modern specimens, even allowing for the sexual dimorph- Grande Pile, France (Ponel et al., 2003), have shown no ism which is pronounced in these species. Since all relevant evidence for evolutionary change, despite being located body parts were recovered amongst the fossil assemblages in an area of much higher numbers of endemics than in (in some cases bodies were completely intact apart the British Isles. Overall, however, too little fossil work from antennae and legs), it was clear this was not a case has been undertaken which covers the required periods of of mis-matching body parts. This finding hints at time and in the right location to adequately address this morphological differences between the modern type issue. ARTICLE IN PRESS 1782 N.J. Whitehouse / Quaternary Science Reviews 25 (2006) 1755–1789

The apparent inability of Holocene forest beetles to shaping the nature of our fauna today. This paper has move backwards and forwards across the landscape, in found that: response to climate or habitat change—unlike many non- phytophagous taxa which are commonly used for tem- It is still unclear when many of these species disappear perature reconstructions—also means that we should from the fossil record. At least a sizeable proportion consider whether the stasis hypothesis is entirely applicable probably disappears with primary clearance during the to these groups. Isolation leads to one of two outcomes: prehistoric period, although at least some taxa survive speciation or extirpation. Under changing conditions and into the Medieval period. There are hints that there are decreasing habitat during the Holocene, isolation resulted differences in the records between Britain and Ireland, in populations too small to survive, leading to local although our knowledge of the latter’s record is still very extirpation. However, it raises the question of whether limited. In particular, it appears that some taxa may speciation could occur given sufficient time, isolation and have survived in Ireland up until the Medieval period, continuity of habitat? Both Stewart and Lister (2001) and but that high levels of post-Medieval woodland clear- Willis and van Andel (2004) suggest that some northern ance caused the further eradication of other less refugial areas could have maintained relatively stable tree specialised taxa. It is possible that the use of wood in populations and that their relatively small sizes could have house structures in towns may have provided important brought about accelerated evolution. refugia for some species during this period. In contrast, The importance of ‘‘founder effects’’ in island popula- the establishment of Forests, Chases and the practises of tions (the genetic profile of the founding population) has traditional woodland management in England ensured been stressed by Sam Berry (1979, 2004). His ideas have the survival of some of the species which contracted in relevance both for isolated groups as well as those Ireland. colonising new areas. The effect of colonisation by (or A combination of factors has contributed to the isolation of) a small number of individuals will be a extirpation of taxa, including species’ requirement for population whose ability to cope with stresses is deter- continuity of quality habitat, increasing geographic mined by the genetic composition of the founding (or isolation in areas such as raised mires and other residual) group. This may be a sub-set of the genetic wetlands and the decline of pine and fire habitats. There composition of the parent population and thus differ quite is some evidence to support the idea that climate change substantially from the main population. This genetic may have played an important role in some local composition will dictate the organisms’ ability to respond extirpation. This may have been related to changes in to natural selection and can, on occasion, produce almost temperature, but perhaps also a move from a more instant differentiation, with sub-speciation occurring as continental climate, to a wetter, more oceanic one. quickly as a few decades, as in the case of the Faeroe house However, it is clear that extirpations are the result of a mouse (Berry et al., 1978; Berry, 2004). complex interplay between climate changes on the one These ideas suggest that it may be necessary to consider hand and human impact on the other. whether the lack of speciation evidence in the fossil insect Work on the Irish fossil forest faunas indicates that we record is true only for taxa which are highly mobile and need to consider the importance and contribution of able to move back and forth across the landscape—but several different modes of insect dispersal during the may be less applicable to specialised and endemic popula- early Holocene, including wood rafting, without re- tions which are prone to become isolated. Further research course to physical land-bridges. is undoubtedly required to examine these possibilities, but Fossil beetles associated with ancient forests have a could prove to be a valuable field of enquiry. potentially important role to play in the glacial refugial debate and in discussions of isolation and speciation. 4. Conclusions More research is required in this area.

Fossil beetle work spanning several decades of research Future research needs to take several different directions. from archaeological and palaeoecological sites suggests Firstly, we need to understand the wider context of these that Britain and Ireland have lost a large portion of their faunas and examine similar fossil faunas from across original old-growth forest Coleoptera, particularly the Europe. Is what is happening in Britain and Ireland unique saproxylic species, coinciding with the loss of primeval or are we seeing only part of a wider complex European and semi-natural forests and woodlands. Research over the picture? A series of comparative sites across Europe would last decade or more indicates that we now have some provide a much-needed context to these studies. Our insight into the decline of these species and some of the understanding of the early Holocene is relatively poor, drivers behind their extirpation as well as their wider despite its potential to provide valuable answers, particu- significance. The fossil insect work also provides a long- larly concerning the arrival and development of the forest term perspective on the effects of a range of environmental, fauna. The later Holocene is also poorly investigated and climatic and human factors on our insect fauna and no doubt holds the key to some of the events identified in highlights the important part played by time and history in the fossil record. This is partially because of a paucity of ARTICLE IN PRESS N.J. Whitehouse / Quaternary Science Reviews 25 (2006) 1755–1789 1783 suitable peat deposits which relate to this period. There are, Sippola; Pete Skidmore; David Smith; Eileen Reilly; Mark however, still many deposits—such as in-filled woodland Robinson; Frans Vera; Pat Wagner. Libby Mulqueeny, hollows, in-filled palaeochannels and shallow lakes—which Queen’s University Belfast, is thanked for cartographic could all be suitable for investigation and which could assistance. Finally, I would like to thank Dr. Gill Plunkett provide valuable insights into this period. Moreover, how and Mr. Philip Barratt, Queen’s University Belfast for their does the biotic history of the present interglacial differ comments on the manuscript, Prof. Mark Robinson, from that of previous interglacials? Investigations of University of Oxford and Dr. Scott Elias, Royal Holloway previous interglacials have been mostly concerned with College, University of London for their helpful reviews. climate reconstruction and broad environmental recon- This paper is dedicated to the memory of Stephen struction rather than biotic development, although the Warburton who tirelessly campaigned to save Thorne and relatively poor chronologic resolution of suitable deposits Hatfield Moors but sadly died in January 2004. may make such an exercise challenging. Finally, the comparison of fossil data to modern distribution patterns Appendix A. Supplementary data and DNA profiling would be immensely valuable in assessing the impact of refugia on population structure The online version of this article contains additional and development. supplementary data. Please search for doi:10.1016/j. quascirev.2006.01.010. Acknowledgments

The author would like to thank Scott Elias for the opportunity of submitting this review paper and his References patience during completion of the manuscript. Palaeoen- tomological analysis in the Humberhead Levels was Alexander, K., 1999. Should deadwood be left in sun or shade? British conducted as part of a PhD funded by the Hossein Farmy Wildlife 10, 342. Alexander, K.N.A., 2002. The Invertebrates of Living and Decaying Fund (University of Sheffield), under the supervision of Timber in Britain and Ireland. English Nature Research Reports 467. Professors Paul Buckland and Kevin Edwards. They are English Nature, Peterborough. warmly thanked for their help and guidance in the project Anderson, R., Nash, R., O’Connor, J.P., 1997. Irish Coleoptera; a revised and their continued support. The following organisations and annotated list (Special Entomological Supplement). Belfast, the provided funding and/or assistance: Hossein Farmy Irish Naturalists’ Journal. Anderson, R., Simms, M., Nelson, B., 2000. A review of lowland wood Foundation (University of Sheffield), NERC Radiocarbon pasture and parkland in Northern Ireland. Unpublished Report to Steering Committee, who granted 18 new radiocarbon Environment and Heritage Service, Northern Ireland. dates for the Humberhead Levels as part of these studies Andrieu-Ponel, V., Ponel, P., 1999. Human impact on Mediterranean (Allocation 659/0896); Kelt UK; European Science Foun- wetland Coleoptera: an historical perspective at Tourves (Var, dation (ESF) (Network on Fossil Insects), the Royal France). Biodiversity and Conservation 8, 391–407. Andrieu-Ponel, V., Ponel, P., Bruneton, H., Leveau, P., de Beaulieu, J.-L., Society. English Nature and Levingtons allowed access 2000. Palaeoenvironments and cultural landscapes of the last 2000 onto the milling areas of Thorne and Hatfield Moors. years reconstructed from pollen and Coleopteran records in the Lower Robin Goforth of Hayfield Lodge Farm, Rossington, Rhoˆne valley, southern France. The Holocene 10, 341–355. allowed access to the Rossington site. Peter Osborne Ashworth, A.C., 1973. The climatic significance of a late Quaternary insect provided the unpublished data from Misterton Carr. fauna from Rodbaston Hall, Staffordshire, England. Entomologia Scandinavica 4, 191–205. Bullrush peat, Ballymacombs More, Co. Antrim are Ashworth, A.C., 2001. Perspectives on Quaternary beetles and climate thanked for access onto the site. A considerably shorter change. In: Gerhard, L.C., Harrison, W.E., Hanson, B.M. (Eds.), version of this paper was presented at the XXIIth World Geological Perspectives of Global Climate Change. American Asso- Entomological Congress, Brisbane, Australia in August ciation of Petroleum Geologists’ Studies in Geology 47, 153–168. 2004; the Royal Society, British Academy and Queen’s Ashworth, A.C., 2004. Quaternary Coleoptera of the United States and Canada. Developments in Quaternary Science 1, 505–517. University Belfast are thanked for their support towards Atkinson, T.C., Briffa, K.R., Coope, G.R., 1989. Seasonal temperatures travel to this conference. in Britain during the past 22,000 years, reconstructed using beetle Access to insect reference collections was provided by remains. Nature 325, 587–592. Doncaster Museum; Manchester Museum; The British Atlegrim, O., Sjo¨berg, K., 1995. Effects of clear-cutting and selective Museum (Natural History); Oxford University Museum; felling in Swedish boreal coniferous forest: response of invertebrate taxa eaten by birds. Entomologica Fennica 6, 79–90. the Royal Entomological Society, London, allowed access Baker, R.G., Bettis, E.A., Schwert, D.P., Horton, D.G., Chumbley, C.A., to specialist literature. The author would also like to thank Gonzales, L.A., Reagan, M.K., 1996. Holocene palaeoenvironments of the following people for their insights: Keith Alexander; northeast Iowa. Ecological Monographs 66, 203–234. Roy Anderson; Martin Bell; Gretel Boswijk; Paul Buck- Bakker, E.S., Olff, H., Vandenberghe, C., Maeyer De, K., Smit, R., land; Russell Coope; Mark Dinnin; Brian Evesham; Sheila Gleichman, J.M., Vera, F.W.M., 2004. Ecological anachronisms in the recruitment of temperate light-demanding tree species in wooded Hicks; James Hogan; Colin Johnson; Helen Kirk; Darren pastures. Journal of Applied Ecology 41, 571–582. Mann; Petri Martikainen; Fraser Mitchell; Richard Preece; Balfour-Browne, F., 1958. British Water Beetles, vol. 3. Ray Society, Karen Rodgers; Peter Roworth; Jon Sadler; Anna-Liisa London. ARTICLE IN PRESS 1784 N.J. Whitehouse / Quaternary Science Reviews 25 (2006) 1755–1789

Bennett, K.D., 1984. The post-glacial history of Pinus sylvestris in the Bradshaw, R., Hannon, G., 1992. Climatic change, human influence and British Isles. Quaternary Science Reviews 3, 133–155. disturbance regime in the control of vegetation dynamics within Fiby Bennett, K.D., 1988. A provisional map of forest types for the British Isles Forest, Sweden. Journal of Ecology 80, 625–632. 5000 years ago. Journal of Quaternary Science 4, 141–144. Bradshaw, R.H.W., Hannon, G.E., 2004. The Holocene structure of Bennett, K.D., 1995. Post-glacial dynamics of pine (Pinus sylvestris) and North-West European temperate forest induced from palaeoecological pinewoods in Scotland. In: Aldhous, J.R. (Ed.), Our Pinewood data. In: Honnay, O., Verheyen, K., Bossuyt, B., Hermy, M. (Eds.), Heritage (Proceedings of a Conference at Culloden Academy, Forest Biodiversity: Lessons from History for Conservation. CABI Inverness). Forestry Commission, Inverness, pp. 22–39. Publishing, Wallingford, Oxfordshire, pp. 11–26. Bennett, K.D., Tzedakis, P.C., Willis, K.J., 1991. Quaternary refugia of Bradshaw, R.H.W., Hannon, G.E., Lister, A., 2003. A long-term north European trees. Journal of Biogeography 18, 103–115. perspective on ungulate-vegetation interactions. Forest Ecology and Berry, R.J., 1979. Genetic factors in animal population dynamics. In: Management 181, 267–280. Anderson, R.M., Turner, B.D., Taylor, L.R. (Eds.), Population Brayshay, B.A., Dinnin, M., 1999. Integrated palaeoecological evidence Dynamics (Twentieth Symposium of the British Ecological Society). for biodiversity at the floodplain-forest margin. Journal of Biogeo- Blackwell Scientific, Oxford, pp. 53–80. graphy 26, 115–131. Berry, R.J., 2004. Island differentiation muddied by island biogeogra- Bridge, M.C., Haggart, B.A., Lowe, J.J., 1990. The history and phers. Environmental Archaeology 9 (2), 117–121. palaeoclimatic significance of sub-fossil remains of Pinus sylvestris in Berry, R.J., Jakobson, M.E., Peters, J., 1978. The house mouse of the blanket peats from Scotland. Journal of Ecology 78, 77–99. Faroe Islands: a study of microdifferentiation. Journal of Zoology 185, Broecker, W.S., Kennett, J.P., Flower, B.P., Teller, J.T., Trumbore, S., 73–92. Bonani, G., Wolfli, W., 1989. Routing of meltwater from the Bhiry, N., Filion, L., 1996. Mid-Holocene Hemlock decline in Eastern Laurentide ice sheet during the Younger Dryas cold episode. Nature North America linked with phytophagous insect activity. Quaternary 341, 318–321. Research 45, 312–320. Buckland, P.C., 1975. Synanthropy and the deathwatch: a discussion. Birks, H.J.B., 1989. Holocene isochrone maps and patterns of Naturalist 100, 37–42. tree-spreading in the British Isles. Journal of Biogeography 16, Buckland, P.C., 1979. Thorne Moors: A Palaeoentomological Study of a 503–540. Bronze Age Site. University of Birmingham, Birmingham. Birks, H.J.B., 2005. Mind the gap: how open were European primeval Buckland, P.C., Coope, G.R., 1991. A Bibliography and Literature forests? Trends in Ecology and Evolution 20 (4), 154–156. Review of Quaternary Entomology. J.R. Collis Publications, Uni- Bistro¨m, O., Va¨isa¨nen, R., 1988. Ancient forest invertebrates of the versity of Sheffield. Pyha¨n-Ha¨kki National park in Central Finland. Acta Zoologica Buckland, P.C., Dinnin, M.H., 1993. Holocene woodlands, the fossil Fennica 185, 1–69. insect evidence. In: Kirby, K.J., Drake, C.M. (Eds.), Dead Wood Blair, K.G., 1935. Beetle remains from a block of peat on the coast of East Matters: The Ecology and Conservation of Saproxylic Invertebrates in Anglia. Proceedings of the Royal Entomological Society of London Britain. English Nature, Peterborough, pp. 6–20. 10, 19–20. Buckland, P.C., Dolby, M.J., 1973. Mesolithic and later material from Blumer, M.A., 2002. Grazing ecology and forest history. Environmental Misterton Carr, Nottinghamshire—an interim report. Transactions of History 7, 687–689. the Thoroton Society of Nottingham 17, 5–33. Bond, G., Showers, W., Chesby, M., Lotti, R., Almasi, P., deMenocal, P., Buckland, P.C., Kenward, H.K., 1973. Thorne Moor: a palaeo-ecological Priore, P., Cullen, H., Hajdas, I., Bonani, G., 1997. A pervasive study of a Bronze Age site. Nature 241, 405–407. millennial-scale cycle in north Atlantic Holocene and glacial climates. Buckland, P.C., Sadler, J., 1985. The nature of late Flandrian alluviation Science 278, 1257–1266. in the Humberhead Levels. East Midland Geographer 8, 239–251. Bond, G., Kromer, B., Beer, J., Muschler, R., Evans, M.N., Showers, W., Buckland, P.C., Wagner, P.E., 2001. Is there an insect signal for the Hoffman, S., Lotti-Bond, R., Hajdas, I., Bonani, G., 2001. Persistent ‘‘Little Ice Age’’? Climatic Change 48, 137–149. solar influence on north Atlantic climate during the Holocene. Science Buckland, P.C., Sveinbjarnardottir, G., Savory, D., McGovern, T.H., 294, 2130–2136. Skidmore, P., Andreasen, C., 1983. Norseman at Nipaitsoq, Green- Bo¨cher, J., 1986. Boreal insects in northernmost Greenland: palaeoento- land; a palaeoecological study. Norwegian Archaeological Review 16, mological evidence from the Kap Kobemhavn Formation (Plio- 186–189. Pleistocene) Peary Land. Fauna Norvegica B 36, 37–43. Buckland, P.I., Yuan Zhuo, D., Buckland, P.C., 1997. Towards an expert Bo¨cher, J., 1997. History of the Greenland insect fauna with emphasis on system in palaeoentomology. Quaternary Proceedings 5, 67–78. living and fossil beetles. Quaternary Proceedings 5, 35–48. Carr, J.W., 1916. The Invertebrate Fauna of Nottinghamshire. Notting- Boswijk, G., 1998. A dendrochronological study of oak and pine from the ham Naturalists’ Society, Nottingham. Humberhead levels, Eastern England. Unpublished Ph.D. Thesis, Caseldine, C., Gearey, B., Hutton, J., Reilly, E., Stujs, I., Casparie, W., 2001. University of Sheffield. From the wet to the dry: palaeoecological studies at Derryville, Co. Boswijk, G., 2003. The buried forest of Thorne Moors. Thorne and Tipperary, Ireland. In: Raftery, B., Hickey, J. (Eds.), Recent Develop- Hatfield Moors Papers 6, 52–65. ments in Wetland Research (UCD/WARP Occasional Paper 14). Dept. Boswijk, G., Whitehouse, N.J., 2002. Pinus and Prostomis: a dendrochro- of Archaeology, University College Dublin, Dublin, pp. 99–115. nological and palaeoentomological study of a mid-Holocene woodland Chapman, H.P., Gearey, B.R., 2003. Archaeological predictive modelling in eastern England. The Holocene 12, 585–596. in raised mires—concerns and approaches for their interpretation and Boswijk, G., Whitehouse, N.J., Smith, B.M., Buckland, P.C., 2001. management. Journal of Wetland Archaeology 2, 77–88. Thorne Moors. In: Bateman, M.D., Buckland, P.C., Frederick, C.D., Clark, P.U., Marshall, S.J., Clarke, G.K.C., Hostetler, S.W., Licciardi, Whitehouse, N.J. (Eds.), The Quaternary of East Yorkshire and North J.M., Teller, J.T., 2001. Freshwater forcing of abrupt climate change Lincolnshire (Field Guide). Quaternary Research Association, Lon- during the last glaciation. Science 293, 283–287. don, pp. 169–178. Coope, G.R., 1959. A late Pleistocene insect fauna from Chelford, Bradbury, J.P., Colman, S.M., Reynolds, R.L., 2004. The history of recent Cheshire. Proceedings of the Royal Society B 151, 70–86. limnological changes and human impact on Upper Klamath Lake, Coope, G.R., 1970. Interpretations of Quaternary insect fossils. Annual Oregon. Journal of Palaeolimnology 21, 151–165. Reviews of Entomology 15, 97–120. Bradshaw, R., 1993. Forest response to Holocene climatic change: Coope, G.R., 1971. Insecta. In: Colhoun, E.A., Mitchell, G.F. (Eds.), equilibrium or non-equilibrium. In: Chambers, F.M. (Ed.), Climate Interglacial Marine Formation and Lateglacial Freshwater Formation Change and Human Impact on the Landscape. Chapman & Hall, in Shortalstown, Co. Wexford. Proceedings of the Royal Irish London, pp. 57–65. Academy 71B, 234–238. ARTICLE IN PRESS N.J. Whitehouse / Quaternary Science Reviews 25 (2006) 1755–1789 1785

Coope, G.R., 1978. Constancy of insect species versus inconstancy of Edwards, K.J., 1996. A Mesolithic of the Western and Northern Isles of Quaternary environments. In: Mound, L.A., Waldoff, N. (Eds.), Scotland? Evidence from pollen and charcoal. In: Pollard, T., Diversity of Insect Faunas. Symposia of the Royal Entomological Morrison, A. (Eds.), The Early Prehistory of Scotland. Edinburgh Society of London 9. Blackwell, London, pp. 176–187. University Press, Edinburgh, pp. 23–38. Coope, G.R., 1979. The Carabidae of the glacial refuge in the British Isles Edwards, K.J., 1999. Palynology and people: observations on the British and their contribution to the post-glacial colonisation of Scandinavia record. Quaternary Proceedings 7, 531–544. and North Atlantic Islands. In: Erwin, T.L., Ball, G.E., Whitehead, Elias, S.A., 1994. Quaternary Insects and their Environments. Smithso- D.R. (Eds.), Carabid Beetles: Their Evolution, Natural History and nian Institution Press, Washington. Classification. Proceedings of the First International Symposium of Elton, C.S., 1966. The Pattern of Animal Communities. Methuen, Carabidology Smithsonian Institution, Washington, DC. Junk Pub- London. lishers, The Hague, pp. 407–424. Eriksson, O., Cousins, S.A.O., Bruun, H.H., 2002. Land-use history and Coope, G.R., 1990. The invasion of Northern Europe during the fragmentation of traditionally managed grasslands in Scandinavia. Pleistocene by Mediterranean species of Coleoptera. In: di Castri, F., Journal of Vegetation Science 13, 743–748. Hansen, A.J., Debussche, M. (Eds.), Biological Invasions in Europe Gear, A.J., 1989. Holocene vegetation history and the palaeoecology of and the Mediterranean Basin. Kluwer, Dordrecht, pp. 203–215. pinus sylvestris in northern Scotland. Unpublished PhD thesis, Coope, G.R., 1995. Insect faunas in ice age environments: why so little University of Durham. extinction? In: Lawton, J.H., May, R.M. (Eds.), Extinction Rates. Gear, A.J., Huntley, B., 1991. Rapid changes in the range limits of Scots Oxford University Press, Oxford, pp. 55–74. pine 4000 years ago. Science 251, 544–547. Coope, G.R., 1998. Insects. In: Preece, R.C., Bridgland, D.R. (Eds.), Late Girling, M.A., 1976. Fossil Coleoptera from the Somerset Levels: the Quaternary Environmental Change in North-West Europe: Excava- Abbot’s Way. Somerset Levels Papers 2, 28–33. tions at Holywell Coombe, South-East England. Chapman & Hall, Girling, M.A., 1979. Fossil insects from the Sweet Track. Somerset Levels London, pp. 213–233. Papers 5, 84–93. Coope, G.R., 2004. Several million years of stability among insect species Girling, M.A., 1980. The fossil insect assemblage from the Baker site. because of, or in spite of, Ice Age climatic instability? Philosophical Somerset Levels Papers 6, 36–42. Transactions of the Royal Society of London B 359, 209–214. Girling, M.A., 1982. Fossil insect faunas from forest sites. In: Bell, M., Coope, G.R., Lemdahl, G., 1995. Regional differences in the late-glacial Limbrey, S. (Eds.), Archaeological Aspects of Woodland Ecology climate of northern Europe based on Coleopteran analysis. Journal of (British Archaeological Reports International Series 146). British Quaternary Science 10, 391–395. Archaeological Reports, Oxford, pp. 129–146. Coope, G.R., Shotton, F.W., Strachan, I., 1961. A late Pleistocene flora Girling, M.A., 1984a. Investigations of a second insect assemblage from and fauna from Upton Warren, Worcestershire. Philosophical the Sweet Track. Somerset Levels Papers 10, 79–91. Transactions of the Royal Society B 244, 379–421. Girling, M., 1984b. A Little Ice Age extinction of a water beetle from Coope, G.R., Dickson, J.H., McCutcheon, J.A., Mitchell, G.F., 1979. The Britain. Boreas 13, 1–4. late-glacial and early postglacial deposit at Drumurcher, Co. Girling, M.A., 1985. An ‘old forest’ beetle fauna from a Neolithic and Monaghan. Proceedings of the Royal Irish Academy B 79, 63–85. Bronze Age peat deposit at Stileway. Somerset Levels Papers 11, Coope, G.R., Lemdahl, G., Lowe, J.J., Walkling, A., 1998. Temperature 80–85. gradients in northern Europe during the last glacial-Holocene Girling, M., 1989. Mesolithic and later landscapes interpreted from the transition (14-914 C kyr BP) interpreted from coleopteran assemblages. insect assemblages of West Heath Spa, Hampstead. In: Collins, D., Journal of Quaternary Science 13, 419–433. Lorimer, D. (Eds.), Excavations at the Mesolithic site on West Heath, Danks, H.V., Foottit, R.G., 1989. Insects of the boreal zone of Canada. Hampstead 1976–1981 (British Archaeological Reports 217). British Canadian Entomologist 121, 626–674. Archaeological Reports, York, pp. 72–89. Dennis, P., 1997. Impact of forest and woodland structure on insect Greig, J., 1982. Past and present lime woods in Europe. In: Bell, M., abundance and diversity. In: Watt, A.D., Stork, N.E., Hunter, M.D. Limbrey, S. (Eds.), Archaeological Aspects of Woodland Ecology (Eds.), Forests and Insects. Chapman & Hall, London, pp. 321–340. (British Archaeological Reports International Series 146). British Devoy, R.J.N., 1995. Deglaciation, earth-crustal behaviour and sea-level Archaeological Reports, Oxford, pp. 23–55. changes in the determination of insularity: a perspective from Ireland. Grove, J.M., 1988. The Little Ice Age. Methuen, London. In: Preece, R.C. (Ed.), Island Britain: A Quaternary Perspective. Haila, Y., Kaila, L., Koponen, P., Martikainen, P., Niemela¨, P., Puntitila, Geological Society. Special Publication 96, London, pp. 181–208. P., Siitonen, J., 1995. Effects of different forest management histories Dinnin, M.H., 1991. The sub-fossil occurrence of the greater silver on the invertebrate fauna. In: Angelstam, P., Mikusinki, G., Travina, water—beetle Hydrophilus piceus (L.) (Col.: Hydrophilidae) at Shirley S. (Eds.), Research in Eastern Europe to Solve Nature Conservation Pool, South Yorkshire. Naturalist 116, 57–59. Problems in the Nordic Countries. Swedish University of Agricultural Dinnin, M., 1997. Holocene beetle assemblages from the lower Trent Sciences, Uppsala, pp. 29–30. floodplain at Bole Ings, Nottinghamshire, UK. Quaternary Proceed- Hall, V.A., 1995. Woodland depletion in Ireland over the last millennium. ings 5, 83–104. In: Plicher, J.R., Mac an tSaoir, S.S. (Eds.), Wood, Trees and Forests Dinnin, M.H., Lillie, M., 1995. The palaeoenvironmental survey of in Ireoland. Royal Irish Academy, Dublin, pp. 23–33. Southern Holderness and evidence for sea-level change. In: Van de Hall, V.A., 1998. Recent landscape change and landscape restoration in Noort, R., Ellis, S. (Eds.), Wetland Heritage of Holderness: an Northern Ireland: a tephra-dated pollen study. Review of Palaeobo- Archaeological Survey. Humber Wetlands Project, University of Hull, tany and Palynology 103, 59–68. Hull, pp. 87–120. Hall, V.A., 2000. The documentary and pollen analytical records of the Dinnin, M.H., Sadler, J.P., 1999. 10,000 years of change: the Holocene vegetational history of the Irish landscape AD200–1650. Peritia 14, entomofauna of the British Isles. Quaternary Proceedings 7, 545–562. 342–371. Dinnin, M.H., with contributions by Ellis, S., Weir, D., 1997. The Hall, A., Kenward, H., 1980. An interpretation of biological remains palaeoenvironmental survey of West, Thorne and Hatfield Moors. In: from Highgate, Beverley. Journal of Archaeological Science 7, Van de Noort, R., Ellis, S. (Eds.), Wetland Heritage of the Humber- 33–51. head Levels. University of Hull, Hull, pp. 157–189. Hall, A., Kenward, H., 1990. Environmental Evidence from the Colonia. Donisthorpe, H. St. J.K., 1939. The Coleoptera of Windsor Forest. The Archaeology of York 14/6. Council for British Archaeology, Published privately, London. London. Duffy, E.A.J., 1968. The status of Cerambyx L. (Col., Cerambycidae) in Hall, A.R., Kenward, H.K., McCormish, J.M., 2003. Pattern in thinly Britain. Entomologist’s Gazette 19, 164–166. distributed plant and invertebrate macrofossils revealed by extensive ARTICLE IN PRESS 1786 N.J. Whitehouse / Quaternary Science Reviews 25 (2006) 1755–1789

analysis of occupation deposits at Low Fisher Gate, Doncaster, UK. Kenward, H.K., Allison, E.P., 1994. Rural origins of the urban insect Environmental Archaeology 8 (2), 129–144. fauna. In: Hall, A., Kenward, H.K. (Eds.), Urban–Rural Connexions: Hammond, P.M., 1974. Changes in the British Coleopterous fauna. In: Perspectives from Environmental Archaeology (Symposia of the Hawksworth, D.L. (Ed.), The Changing Flora and Fauna of Britain Association for Environmental Archaeology). Oxbow, Oxford, pp. (The Systematics Association Special Volume No. 6). Academic Press, 55–77. London, pp. 323–369. Kenward, H.K., Hall, A.R., 1995. Biological Evidence from 16–22 Harding, P.T., Plant, R.A., 1978. A second record of Cerambyx cerdo L. Coppergate, York (Archaeology of York: The Environment 14/7). (Coleoptera: Cerambycidae) from sub-fossil remains in Britain. York Archaeological Trust, York. Entomologist’s Gazette 29, 150–152. Kirby, K.J., 2003. What Might a British Forest-Landscape Driven By Harding, P.T., Rose, F., 1986. Pasture-Woodlands in Lowland Britain. Large Herbivores Look Like? (Research Report 530). English Nature, Natural Environment Research Council Institute of Terrestrial Peterborough. Ecology, Peterborough. Kirby, K.J., 2004. A model of a natural wooded landscape in Britain as Hayfield, C., Greig, G., 1989. Excavation and salvage work on a moated influenced by large herbivore activity. Forestry 77 (5), 405–420. site at Cowick, South Humberside, 1976. The Yorkshire Archae- Klinger, L.F., Elias, S.A., Behan-Pelletier, V.M., Williams, N.E., 1990. ological Journal 61, 41–69. The bog climax hypothesis: fossil and stratigraphic evidence Helio¨vaara, K., Va¨isa¨nen, R., 1984. Effects of modern forestry on north- in peat sections from south-east Alaska, USA. Holoarctic Ecology 13, western European Forest invertebrates: a synthesis. Acta Forestalia 72–80. Fennica 189, 1–32. Kloet, G.S., Hincks, W.D., 1977. A check list of British Insects (2nd Hewitt, G.M., 1996. Some genetic consequences of ice ages, and their role edition, revised by R.D. Pope). Royal Entomological Society, London. in divergence and speciation. Biological Journal of the Linnaean Koch, K., 1989. Die Ka¨fer Mitteleuropas. O¨kologie, 1 and 2. Goecke and Society 58, 247–276. Evers, Krefeld. Homan, T., 1991. A yearning toward wildness; environmental quotations Koponen, M., Nuorteva, M., 1973. Uber su¨bfossile Waldinsekten aus from the writings of Henry David Thoreau. Peachtree Publishers, dem Moor Piilonsuo in Sudfinnland. Acta Entomologica Fennica 29, Atlanta. 1–84. Horion, A., 1953. Faunistik der Mitteleuropa¨ischen, Ka¨fer 3. Malaco- Kullman, L., 2002. Boreal tree taxa in the central Scandes during the Late- dermata, Sternoxia (Elateridae—Throscidae). G. Frey, Munich. Glacial: implications for Late-Quaternary forest history. Journal of Horion, A., 1960. Faunistik der Mitteleuropa¨ischen Ka¨fer, 7. Clavicornia, Biogeography 29, 1117–1124. Sphaeritidae—Phalacridae. Uberlingen-Bodensee. Lageard, J.G.A., Chambers, F.M., Thomas, P.A., 1999. Climatic Hughes, P.D.M., Kenward, H.K., Hall, A.R., Large, F.D., 2000. A high- significance of the marginalisation of Scots pine (Pinus sylvestris L.) resolution record of mire development and climatic change spanning ca 2500 BC at White Moss, south Cheshire, UK. The Holocene 9 (3), the Late-glacial-Holocene boundary at Church Moss, Davenham 321–331. (Cheshire, England). Journal of Quaternary Science 15 (7), 697–724. Lambeck, K., 1996. Glaciation and sea-level change for Ireland and the Hunter, F.A., 1977. Ecology of pinewood beetles. In: Bunce, R.G.H., Irish Sea since Late Devensian/Midlandian time. Journal of the Jeffers, J.N.R. (Eds.), Native Pinewoods of Scotland. Natural Geological Society, London 153, 853–872. Environment Research Council Institute of Terrestrial Ecology, Lambrick, G.H., Robinson, M.A., 1979. Iron Age and Roman Riverside Peterborough, pp. 42–55. Settlement at Farmoor, Oxfordshire (CBA Report 32). Council for Huntley, B., 1993. Rapid early Holocene migration and high abundance of British Archaeology, London. hazel (Corylus avellana L.): alternative hypotheses. In: Chambers, Lekander, B., Koponen, M., Nuorteva, M., 1975. Eremotes larva (Col., F.M. (Ed.), Climate Change and Human Impact on the Landscape. Curculionidae) found as subfossil. Annales Entomologici Fennici 4, Chapman & Hall, London, pp. 205–215. 121–123. Hyman, P.S., 1992. A review of the scarce and threatened Coleoptera of Lindbladh, M., Niklasson, M., Nilsson, S.G., 2003. Long-time record of Great Britain; Part 1. (Revised and updated by M.S. Parsons). UK fire and open canopy in a high biodiversity forest in southeast Sweden. Joint Nature Conservation Committee, Peterborough. Biological Conservation 114, 231–243. Hyman, P.S., 1994. A review of the scarce and threatened Coleoptera of Lindroth, C.H., 1945. Die Fennoskandischen Carabidae I–III. Goteborgs Great Britain. Part 2. (Revised and updated by M.S. Parsons). UK K. Vetensk. o VitterhSamh, Goteborg. Joint Nature Conservation Committee, Peterborough. Lindroth, C.H., 1948. Interglacial insect remains from Sweden. Sveriges Johansen, S., Hytteborn, H., 2001. A contribution to the discussion of Geologiska Underso¨kning C 42, 1–28. biota dispersal with drift ice and driftwood in the North Atlantic. Long, A.J., Innes, J.B., Kirby, J.R., Lloyd, J.M., Rutherford, M.M., Journal of Biogeography 28, 105–115. Shennan, I., Tooley, M.J., 1998. Holocene sea-level change and coastal Kaila, L., Martikainen, P., Puttila, P., Yakovlev, E., 1994. Saproxylic evolution in the Humber estuary: an assessment of rapid coastal beetles (Coleoptera) on dead birch trunks decayed by different change. The Holocene 8 (2), 229–247. polypore species. Annales Zoologici Fennici 31, 97–107. Lowe, J.J., Walker, M.J.C., Scott, E.M., Harkness, D.D., Bryant, C.L., Kaila, L., Martikainen, P., Puttila, P., 1997. Dead trees left in clear-cuts Davies, S.M., 2004. A coherent high-precision radiocarbon chronology benefit saproxylic Coleoptera adapted to natural disturbances in of the Late-glacial sequence at Sluggan Bog, Co. Antrim, Northern boreal forest. Biodiversity and Conservation 6, 1–18. Ireland. Journal of Quaternary Science 19 (2), 147–158. Karppinen, E., Koponen, M., 1973. The subfossil oribatid fauna of Lucht, W.H., 1987. Die Ka¨fer Mitteleuropas. Katalog. Goeke and Evers, Piilonsuo, a bog in southern Finland. Annales Entomologici Fennici Krefeld. 39, 22–32. MacArthur, R.H., Wilson, E.O., 1967. The Theory of Island Biogeo- Karppinen, E., Koponen, M., 1974. Further observations on subfossil graphy. Princeton University Press, New Jersey. remains of oribatid (Acar., Oribatei) and insects in Piilonsuo, a bog in Marra, M., 2003. Last interglacial beetle fauna from New Zealand. southern Finland. Annales Entomologici Fennici 40, 172–175. Quaternary Research 59, 122–131. Keen, D.H., Bateman, M.D., Coope, G.R., Field, M.H., Langford, H.E., Martikainen, P., Siitonen, J., Kaila, L., Punttila, P., 1996. Intensity of Merry, H.E., Mighall, T., 1999. Sedimentology, palaeoecology and forest management and bark beetles in non-epidemic conditions: a geochronology of last Interglacial deposits from Deeping St. James, comparison between Finnish and Russian Karelia. Journal of Applied Lincolnshire, England. Journal of Quaternary Science 14 (5), 411–436. Entomology 120, 257–264. Kelly, M., Osborne, P.J., 1965. Two Faunas and floras from the alluvium Martin, P.S., 1984. Prehistoric overkill: the global model. In: Martin, P.S., at Shustoke, Warwickshire. Proceedings of the Linnaean Society of Klein, R.G. (Eds.), Quaternary Extinctions: A Prehistoric Revolution. London 176, 37–65. University of Arizona Press, Tucson, pp. 354–403. ARTICLE IN PRESS N.J. Whitehouse / Quaternary Science Reviews 25 (2006) 1755–1789 1787

Matthews Jr, J.V., 1970. Two new species of Micropeplus from the Osborne, P.J., 1974. An insect assemblage from Lea Marston, Warwick- of western Alaska with remarks on the evolution of shire and its bearing on the contemporary climate and ecology. Coleoptera: Staphylinidae. Canadian Journal of Zoology 48, 779–788. Quaternary Research 4, 471–486. McCabe, A.M., Coope, G.R., Gennard, D.E., Doughty, P., 1987. Osborne, P.J., 1976. Evidence from the insects of climatic variation during Freshwater organic deposits and stratified deposits between Early the Flandrian period: a preliminary note. World Archaeology 8, and Late Midlandian (Devensian) till sheets, at Aghnadarragh, County 150–158. Antrim, Northern Ireland. Journal of Quaternary Science 2, 11–33. Osborne, P.J., 1978. Insect evidence for the effect of man on the lowland McCracken, E., 1971. The Irish Woods since Tudor times. David and landscape. In: Limbrey, S., Evans, J.G. (Eds.), The Effect of Man on Charles, Newton Abbott. the Landscape: the Lowland Zone. Council for British Archaeology McErlean, T., McConkey, M., Forsythe, W., 2002. Strangford Lough; An Research Report 21. Council for British Archaeology, London, Archaeological Survey of the Maritime Cultural Landscape. Environ- pp. 32–34. ment and Heritage Service, Belfast. Osborne, P.J., 1980. The late Devensian-Flandrian transition depicted by McGlone, M., 1983. Polynesian deforestation of New Zealand: a serial insect faunas from West Bromwich, Staffordshire, England. preliminary synthesis. Archaeology in Oceania 18, 1–10. Boreas 9, 139–147. McGlone, M., Wilmshurst, J.M., 1999. Dating initial Maori environ- Osborne, P.J., 1982. Some British later prehistoric insect faunas and mental impact in New Zealand. Quaternary International 59, 5–16. their climatic implications. In: Harding, A. (Ed.), Climatic Change McLean, I.F.G., Speight, M.C.D., 1993. Saproxylic invertebrates—the in Later Prehistory. Edinburgh University Press, Edinburgh, European context. In: Kirby, K.J., Drake, C.M. (Eds.), Dead Wood pp. 68–74. Matters: The Ecology and Conservation of Saproxylic Invertebrates in Osborne, P.J., 1988. A Late Bronze Age insect fauna from the River Avon, Britain. English Nature, Peterborough, pp. 21–32. Warwickshire, England: its implications for the terrestrial and fluvial McNally, A., Doyle, G.J., 1984. A study of sub-fossil pine layers in a environment and for climate. Journal of Archaeological Science 15, raised bog complex in the Irish Midlands—I. Palaeowoodland extent 715–727. and dynamics. Proceedings of the Royal Irish Academy 84b, 57–70. Osborne, P.J., 1995. An insect fauna of Roman date from Stourport, Mighall, T.M., Lageard, J.G.A., 1999. The prehistoric environment. In: Worcestershire, UK and its environmental implications. Circaea (The O’Brien, W. (Ed.), Sacred Ground; Megalithic Tombs in Coastal Journal of the Association for Environmental Archaeology) 12 (2), Southwest Ireland. National University Ireland, Galway, pp. 41–59. 183–189. Mitchell, G.F., 1963. Moranic ridges on the floor of the Irish sea. Irish Osborne, P.J., 1997. Insects, man and climate in the British Holocene. Geography 4, 335–344. Quaternary Proceedings 5, 193–198. Mitchell, F.J.G., 2005. How open were European primeval forests? Palm, T., 1951. Die Holz und Rindenka¨fer der nordschwedischen Hypothesis testing using palaeoecological data. Journal of Ecology 93, Laubbaume. Meddelelser Statens Sko¨gsfors.-instituten, 40(2). 168–177. Palm, T., 1959. Die Holz und Rindenka¨fer der Sud- und Mittelschwe- Moore, P.D., 2005. Down to the woods yesterday. Nature 433, 588–589. dischen Laubba¨ume. (Opuscula Entomologica Supplementum 16). Morris, M.G., 2002. True Weevils (Part I); Coleoptera: Curculionidae. Entomologiska Sallskapet, Lund. Royal Entomological Society and the Field Studies Council, London. Parker, A.G., Robinson, A.G., 2003. Palaeoenvironmental investigations Muona, J., Rutanen, I., 1994. The short-term impact of fire on the beetle on the middle Thames at Dorney, UK. In: Howard, A.J., Macklin, M., fauna in boreal coniferous forest. Annales Botanici Fennici 31, Passmore, D. (Eds.), Alluvial Archaeology in Europe. Swets and 109–121. Zeitlinger, Lisse, pp. 43–60. Needham, S., 1985. Neolithic and Bronze Age settlement on the buried Parker Pearson, M., 1997. The environmental overview. In: Parker floodplain of Runnymede. Oxford Journal of Archaeology 4, 125–137. Pearson, M., Sydes, R.E. (Eds.), Iron Age Enclosures and Prehistoric Needham, S., 1992. Holocene alluviation and interstratified settlement Landscape of Sutton Common, South Yorkshire. Proceedings of the evidence in the Thames valley at Runneymede Bridge. In: Needham, Prehistoric Society 63, 250–253. S., Macklin, M.G. (Eds.), Archaeology Under Alluvium. Oxbow Patterson, W.A., Backman, A.E., 1988. Fire and disease history of forests. Books, Oxford, pp. 249–260. In: Huntley, B., Webb, T. (Eds.), Vegetation History. Kluwer Needham, S.P., 2000. The Passage of the Thames; Holocene Environment Academic, New York, pp. 603–632. and Settlement at Runnymede. (Runnymede Bridge Research Excava- Patterson, W.A., Edwards, K.J., Maguire, D.J., 1987. Macroscopic tions, Volume 1). British Museum Press, London. charcoal as a fossil indicator of fire. Quaternary Science Reviews 6, Neeson, E., 1997. Woodland in History and Culture. In: Foster, J.W. 3–23. (Ed.), Nature in Ireland. Lilliput Press, Dublin, pp. 133–156. Pearson, R.G., 1966. The origins of the British insect fauna. The Nicholls, K., 2001. Woodland cover in pre-modern Ireland. In: Duffy, Entomologist 99, 69–71. P.J., Edwards, D., FitzPatrick, E. (Eds.), Gaelic Ireland ca 1250- Pearson, G.W., Pilcher, J.R., Baillie, M.G.L., Corbett, D.M., Qua, F., c.1650; Land, Lordship and Settlement. Four Courts Press, Dublin, 1986. High precision 14C measurement of Irish oaks to show the pp. 181–206. natural 14 C variation from AD 1840–5210 BC. Radiocarbon 28 (2B), Niemela¨, J., 1997. Invertebrates and boreal forest management. Con- 911–934. servation Biology 11 (3), 601–610. Penny, L.F., Coope, G.R., Catt, J.A., 1969. Age and Insect Fauna of the O’Connor, J.P., 1979. Blaps lethifera Marsham (Coleoptera: Tenebrioni- Dimlington Silts, East Yorkshire. Nature 224, 65–67. dae), a beetle new to Ireland from Viking Dublin. Entomologist’s Peterken, G.F., 1996. Natural Woodland: Ecology and Conservation in Gazette 30, 295–297. Northern Temperate Regions. Cambridge University Press, Cam- Opdam, P., Wascher, D., 2004. Climate change meets habitat fragmenta- bridge. tion: linking landscape and biogeographical scale levels in research and Petit, R., Aguinagalde, I., de Beaulieu, J-L., Bittkau, C., Brewer, S., conservation. Biological Conservation 117, 285–297. Cheddadi, R., Ennos, R., Finschi, S., Grivet, D., Lascoux, M., Osborne, P.J., 1965. The effect of forest clearance on the distribution of Mohanty, A., Mu¨ller-Starck, G., Demesure-Musch, B., Palme, A., the British insect fauna. Proceedings of the XII International Congress Martin, J.P., Rendell, Vendramin, G., 2003. Glacial refugia: hotspots of Entomology, London, 1964, pp. 556–557. but not melting pots of genetic diversity. Science 300, 1565. Osborne, P.J., 1969. An insect fauna of Late Bronze Age date from Pilcher, J., Hall, V., 2001. Flora Hibernica; the Wild Flowers, Plants and Wilsford, Wiltshire. Journal of Ecology 38, 555–566. Trees of Ireland. The Collins Press, Cork. Osborne, P.J., 1972. Insect faunas of Late Devensian and Flandrian age Pilcher, J.R., Baillie, M.G.L., Brown, D.M., McCormac, F.G., MacSwee- from Church Stretton, Shropshire. Philosophical Transactions of the ney, P.B., McLawrence, A.S., 1995. Dendrochronology of sub-fossil Royal Society of London B 263, 327–367. pine in the north of Ireland. Journal of Ecology 83, 665–671. ARTICLE IN PRESS 1788 N.J. Whitehouse / Quaternary Science Reviews 25 (2006) 1755–1789

Plunkett, G.M., Hall, V.A., Whitehouse, N.J., 2003. Palaeoecological Rogers, K., 2004. An environmental reconstruction of Strangford Lough Investigations at Toome By-Pass: Final Report to the Environment in the early Holocene; a fossil insect perpsective. Unpublished and Heritage Service: Natural Heritage. Unpublished Report, Queen’s undergraduate Dissertation, Queen’s University Belfast. University Belfast. Roper, T., 1993. A ‘‘waste’’ of resources: the origins, growth and decline Ponel, P., Andrieu-Ponel, A., Parchoux, F., Juhasz, I., de Beaulieu, J.-L., of Thorne Moors; a palaeoentomological study. Unpublished MSc 2001. Late-glacial and Holocene high-altitude environmental changes dissertation, University of sheffield. in Valle´e des Merveilles (Alpes—Maritimes, France): insect evidence. Roper, T., 1996. Fossil insect evidence for the development of raised mire Journal of Quaternary Science 16, 795–812. at Thorne Moors, near Doncaster. Biodiversity and Conservation 5, Ponel, P., Orgeas, J., Samways, M.J., Andrieu-Ponel, V., de Beaulieu, J.- 503–521. L., Reille, M., Roche, P., Tatoni, T., 2003. 110,000 years of Roper, T., Whitehouse, N.J., 1997. Beetle remains. In: Parker-Pearson, Quaternary beetle diversity change. Biodiversity and Conservation M., Sydes, R.E. (Eds.), Iron Age Enclosures and Prehistoric Land- 12, 2077–2089. scape of Sutton Common, South Yorkshire. Proceedings of the Praeger, R.L., 1932. Recent views bearing on the problem of the Irish Prehistoric Society 63, 238–245. flora and fauna. Proceedings of the Royal Irish Academy B 41, Rundgren, M., Ingo´lfsson, O., 1999. Plant survival in Iceland during 125–145. periods of glaciation? Journal of Biogeography 26, 387–396. Preece, R.C., Coxon, P., Robinson, J.E., 1986. New biostratigraphic Sadler, J.P., 1990. Beetles, boats and biogeography: insect invaders of the evidence of Post-glacial colonisation of Ireland and for Mesolithic North Atlantic Islands. Acta Archeologica 61, 199–212. forest disturbance. Journal of Biogeography 13, 487–509. Sadler, J.P., 1999. Biodiversity on oceanic islands: a palaeoecological Rackham, O., 1986. The History of the Countryside. Dent and Sons, assessment. Journal of Biogeography 26, 75–87. London. Schwert, D.P., 1996. Effect of Euro-American settlement on an insect Rackham, O., 2003. Ancient Woodland. Its History, Vegetation and Use fauna: paleontological analysis of the recent chitin record of Beetles in England (New Edition). Castlepoint Press, Dalbeattie. (Coleoptera) from Northeastern Iowa. Ecology and Population Ranius, T., Jansson, N., 2000. The influence on forest regrowth, original Biology 89, 53–63. canopy cover and tree size on saproxylic beetles associated with old Shennan, I., Lambeck, K., Flather, R., Horton, B., McArthur, J., Innes, oaks. Biological Conservation 95, 85–94. J., Llyod, J., Rutherford, M., Wigfield, R., 2000. Modelling western Rasmussen, P., 1990. Pollarding of trees in the Neolithic: often North Sea palaeogeographies and tidal changes during the Holocene. presumed—difficult to prove. In: Robinson, D.E. (Ed.), Experimenta- In: Shernnan, I., Andrews, J. (Eds.), Holocene Land-Ocean Interaction tion and Reconstruction in Environmental Archaeology. Symposia of and Environmental Change around the North Sea (Geological Society the Association of Environmental Archaeologists 9. Oxbow Books, Special Publications 166). Geological Society, London, pp. 299–319. Oxford, pp. 139–148. Shirt, D.B., 1987. British Red Data Books: 2. Insects. Nature Conservancy Reilly, E., 1996. The insect fauna (Coleoptera) from the Neolithic Council, Peterborough. trackways Corlea 9 and 10: the environmental implications. In: Siitonen, J., Martikainen, P., 1994. Occurrence of rare and threatened Raftery, B. (Ed.), Trackway Excavations in the Mountdillon Bogs, Co. insects living on decaying Populus tremula: a comparison between Longford 1985–91, vol. 3. Irish Archaeological Wetland Unit Finnish and Russian Karelia. Scandinavian Journal of Forestry Transactions: Crannog Publications, Dublin, pp. 403–410. Research 9, 185–191. Reilly, E., 2003. The contribution of insect remains to an understanding of Simmons, I.G., 1996. The Environmental Impact of Later Mesolithic the environment of Viking-age and medieval Dublin. In: Duffy, Cultures. Edinburgh University Press, Edinburgh. E. (Ed.), Medieval Dublin IV. Proceedings of the Friends Simmons, I.G., 2001. An Environmental History of Great Britain, From of Medieval Dublin Symposium 2002. Four Courts Press, Dublin, 10,000 Years Ago to the Present. Edinburgh University Press, pp. 40–62. Edinburgh. Reimer, P.J., Hughen, K., Guilderson, T.P., McCormac, G., Baillie, M., Sinclair, W.T., Morman, J.D., Ennos, R.A., 1999. The postglacial history Bard, E., Barratt, P., Beck, J.W., Buck, C.E., Damon, P.E., Freidrich, of Scots pine (Pinus sylvestris L.) in western Europe: evidence from M., Kromer, B., Bronk Ramsey, C., Reimer, R., Remmele, S., mitochondrial DNA variation. Molecular Ecology 8, 83–88. Southon, J.R., Stuiver, M., van der Plight, J., 2002. Preliminary report Smith, A.G., 1958. Post-glacial deposits in south Yorkshire and north of the first workshop of the INTCAL04 radiocarbon calibration/ Lincolnshire. New Phytologist 57, 19–49. comparison working group. Radiocarbon 44 (3), 653–661. Smith, A.G., 1970. The influence of Mesolithic and Neolithic man on Reiss, R., Ashworth, A.C., Schwert, D.P., 1999. Molecular genetic British vegetation: a discussion. In: Walker, D., West, R.G. (Eds.), evidence for the post-Pleistocene divergence of populations of the Studies in the Vegetation History of the British Isles. University Press, artic-alpine ground beetle Amara Alpina (Paykull) (Coleoptera: Cambridge, pp. 81–96. Carabidae). Journal of Biogeography 26, 785–794. Smith, A.G., 1981. Palynology of a Mesolithic–Neolithic site in county Reitter, E., 1911. Fauna Germanica. Die Ka¨fer des Deutchen Reiches, Antrim, N. Ireland. Proceedings of the IVth International Palynology Stuttgart. Conference (1976–1977), Lucknow, vol. 3, pp. 248–257. Ribera, I., Vogler, A.P., 2004. Speciation of Iberian diving beetles in Smith, A.G., Goddard, I.C., 1991. A 12,500 yr record of vegetational Pleistocene refugia (Coleoptera, Dytiscidae). Molecular Ecology 13, history of Sluggan Bog, Co. Antrim, N. Ireland. New Phytologist 118, 179–193. 167–187. Riley, D.N., 1980. Early landscapes from the air. Studies of crop marks in Smith, B.M., 1985. A palaeoecological study of raised mires in the South Yorkshire and North Nottinghamshire. Department of Archae- Humberhead Levels. Unpublished Ph.D. Thesis, University of Wales. ology and Prehistory, University of Sheffield, Sheffield. Smith, B.M., 2002. A palaeoecological study of raised mires in the Robinson, M.A., 1991. The Neolithic and late Bronze age insect Humberhead Levels. Thorne and Hatfield Moors Conservation Forum assemblages. In: Needham, S. (Ed.), Excavation and Salvage at (Monographs), and Archeopress, Doncaster and Oxford. Runnymede Bridge, 1978: The Late Bronze Age Waterfront Site. Smith, D., Whitehouse, N.J., 2005. Not seeing the woods for the trees; a British Museum, London, pp. 277–325. palaeoentomological perspective on woodland decline. In: Brinkley, Robinson, M.A., 2000. Middle Mesolithic to late Bronze age insect M., Smith, W., Smith, D.N. (Eds.), ‘‘The Fertile Ground’’; Papers in assemblages and an early Neolithic assemblage of waterlogged Honour of Susan Limbrey. Oxbow Books, Oxford. macroscopic plant remains. In: Needham, S.P. (Ed.), The Passage of Smith, D.N., 2001. Disappearance of elmid ‘‘riffle beetles’’ from lowland the Thames: Holocene Environment and Settlement at Runnymede. river systems—the impact of alluviation. In: O’Connor, T., Nicholson, Runnymede Bridge Research Excavations: Volume 1. The British R. (Eds.), People as Agents of Environmental Change. Oxbow, Museums Press, London, pp. 146–167. Oxford, pp. 75–80. ARTICLE IN PRESS N.J. Whitehouse / Quaternary Science Reviews 25 (2006) 1755–1789 1789

Smith, D.N., Howard, A.J., 2004. Identifying changing fluvial conditions Whitehouse, N.J., 1997a. Silent witnesses: an ‘‘Urwald’’ fossil insect in low gradient alluvial archaeological landscapes: can Coleoptera assemblage from Thorne Moors. Thorne and Hatfield Moors Papers 4, provide insights into changing discharge rates and floodplain evolu- 19–54. tion? Journal of Archaeological Science 31, 109–120. Whitehouse, N.J., 1997b. Insect faunas associated with Pinus sylvestris L. Speight, M.C.D., 1985. The extinction of indigenous Pinus sylvestris in from the mid-Holocene of the Humberhead Levels, Yorkshire, UK. Ireland: relevant faunal data. Irish Naturalist Journal 21, 449–453. Quaternary Proceedings 5, 293–303. Speight, M.C.D., 1989. Saproxylic invertebrates and their conservation. Whitehouse, N.J., 1998. The evolution of the Holocene Wetland landscape Nature and Environment Series No. 42. Council of Europe, Strasbourg. of the Humberhead levels from a fossil insect perspective. Unpublished Spence, J.R., Langor, D.W., Hammond, H.E.J., Pohl, G.R., 1997. Beetle Ph.D. Thesis, University of Sheffield. abundance and diversity in a boreal mixed-wood forest. In: Watt, Whitehouse, N.J., 2000. Forest fires and insects: palaeoentomological A.D., Stork, N., Hunter, M.D. (Eds.), Forest and Insects. Chapman & research from a sub-fossil burnt forest. Palaeogeography, Palaeocli- Hall, London, pp. 287–301. matology, Palaeoecology 164, 231–246. Stace, C., 1991. New Flora of the British Isles. Cambridge University Whitehouse, N.J., 2004. Mire ontogeny, environmental and climate Press, Cambridge. change inferred from fossil beetle successions from Hatfield Moors, Stewart, J.R., Lister, A.M., 2001. Cryptic northern refugia and the origins eastern England. The Holocene 14, 79–93. of the modern biota. Trends in Ecology and Evolution 16, 608–613. Whitehouse, N.J., in press. The study of fossil insect remains in Sutherland, W.J., 2002. Conservation biology: openness in management. environmental and archaeological investigations: an Irish perspective. Nature 418, 834–835. In: Murphy, E., Whitehouse, N.J. (Eds.), Environmental Archaeology Svenning, J.-C., 2002. A review of natural vegetation openness in north- in Ireland. Oxbow Books, Oxford. western Europe. Biological Conservation 104, 133–148. Whitehouse, N.J., Eversham, B.C., 2002. A fossil specimen of Pterostichus Szujecki, A., 1987. Ecology of Forest Insects. Dr. W. Junk Publishers, angustatus (Duftschmid) (Carabidae): implications for the importance Warszawa. of pine and fire habitats. The Coleopterist 11, 107–113. Taberlet, P., Fumagalli, L., Wust-Saucy, A.-G., Cossons, J.-F., 1998. Whitehouse, N.J., Smith, D.N., 2004. ‘‘Islands’’ in Holocene forests: Comparative phylogeography and postglacial colonisation routes in implications for forest openness, landscape clearance and ‘‘culture- Europe. Molecular Ecology 7, 453–464. steppe’’ species. Environmental Archaeology 9 (2), 199–208. Tipping, R., 1994. The form and fate of Scotland’s woodlands. Whitehouse, N.J., Boswijk, G., Buckland, P.C., 1997. Peatlands, past, Proceedings of the Society of Antiquaries of Scotland 124, 1–54. present and future; some comments from the fossil record. In: Parkyn, Turner, J., 1962. The Tilia decline: an anthropogenic interpretation. New L., Stoneman, R., Ingram, H.A.P. (Eds.), Conserving Peatlands. CAB Phytologist 61, 328–341. International, Wallingford, pp. 54–64. Va¨isainen, R., Bistro¨m, O., Helio¨vaara, H., 1993. Sub-cortical Coleoptera Wigfield, R.T.R., 1995. A model of sea-levels in the Irish and Celtic Seas in dead pines and spruces: is primeval species composition maintained during the end-Pleistocene to Holocene transition. In: Preece, R.C. in managed forests? Biodiversity and Conservation 2, 95–113. (Ed.), Island Britain; A Quaternary Perspective. Geological Society, Van de Noort, R., 2004. The Humber Wetlands; the Archaeology of a Special Publication 96, London, pp. 209–242. Dynamic Landscape. Windgather Press, Cheshire. Wikars, L.O., 1992. Skogsbra¨nder och insekter (Forest fires and insects). Vera, F.W.M., 2000. Grazing Ecology and Forest History. CABI Entomologisk Tidskrift 113, 1–11. Publishing, Oxforshire. Wikars, L.-O., Schimmel, J., 2001. Immediate effects of fire-severity on Wagner, P., 1997. Human impact or cooling climate? The ‘‘Little Ice Age’’ soil invertebrates in cut and uncut pine forests. Forest Ecology and and the beetle fauna of the British Isles. Quaternary Proceedings 5, Management 141, 189–200. 269–276. Williams, C.T., 1985. Mesolithic Exploitation Patterns in the Central Walker, M.J.C., Coope, G.R., Lowe, J.J., 1993. The Devensian (Weichse- Pennines: A Palynological Study of Soyland Moor (British Archae- lian) Lateglacial palaeoenvironmental record from Gransmoor, East ological Reports 139). British Archaeological Reports, Oxford. Yorkshire, England. Quaternary Science Reviews 12, 659–680. Willis, K.J., 1996. Where did all the flowers go? The fate of temperate Warren, M.S., Key, R.S., 1991. Woodlands: past, present and potential European flora during glacial periods. Endeavour 20, 110–114. for insects. In: Collins, N.M., Thomas, J.A. (Eds.), The Conservation Willis, K.J., van Andel, T.H., 2004. Trees or no trees? The environments of Insects and their Habitats. 15th Symposium of the Royal of central and Eastern Europe during the last glaciation. Quaternary Entomological Society of London, 14–15 September 1989. Academic Science Reviews 23, 2369–2387. Press, London, pp. 155–211. Willis, K.J., Niklas, K.J., 2004. The role of Quaternary environmental Watts, W.A., 1985. Quaternary vegetation cycles. In: Edwards, K.J., change in plant macroevolution: the exception or the rule? Warren, W.P. (Eds.), The Quaternary History of Ireland. Academic Philosophical Transactions of the Royal Society of London B 359, Press, London, pp. 155–185. 159–172. Whitehead, D.R., 1972. Classification, phylogeny, and zoogeography of Willis, K.J., Whittaker, R.J., 2000. The refugial debate. Science 287, Schizogenius Putzeys (Coleoptera: Carabiidae: Scartini). Quaestiones 1406–1407. Entomologicae 8, 131–348. Willis, K.J., Rudner, E., Su¨megi, P., 2000. The full-glacial forests of Whitehead, P.F., 1989. Changing environments and Coleoptera faunas central and southeastern Europe. Quaternary Research 53, 203–213. from Aston Mill, Worcestershire, England. Entomologist’s Monthly Wormwell, P., 1977. Woodland insect populations changes on the Isle of Magazine 125, 187–197. Rhum in relation to forest history and woodland restoration. Scottish Whitehouse, N.J., 1993. A mid—Holocene forested site from Thorne Forestry 31, 13–36. Moors: the fossil insect evidence. Unpublished M.Sc. Thesis, Zetterstedt, J.W., 1828. Fauna Insectorum Lapponica. Pars 1. Libraria University of Sheffield. Schulziana, Hammone.