Modern Ecological and Palaeoecological Contributions Towards

Postglacial Colonization Conference: Mind the Gap 2006

An ever-closing gap? Modern ecological and palaeoecological contributions towards understanding the Irish post-glacial insect fauna EILEEN REILLY* Botany Department, School of Natural Sciences, University of Dublin, Trinity College, Dublin 2 Over the last 25 years, since the 1983 conference, a range of new evidence has emerged on the changing nature of the post-glacial insect fauna of Ireland. Modern entomological research, especially new surveys and re-evaluation of existing collections, is continually improving our understanding of the Irish fauna (e.g. Alexander 2002, Speight 2004). However, findings from palaeoentomological analyses, particularly sub-fossil beetle remains, from various locations throughout Ireland have been particularly illuminating in filling gaps in our knowledge (e.g. Reilly 2005, Whitehouse 2006). This paper will attempt to evaluate these findings in order to address the following questions: " Do we know more now than we did in 1983 about the Irish native insect fauna?

" Do we know more about colonization, dispersal mechanisms and extinctions of this fauna?

" What gaps still exist in our knowledge and how do we fill them? Background At the 1983 conference, two contributors, Martin Speight and T.K. McCarthy, discussed the general state of knowledge at that time regarding the Irish postglacial insect fauna (Speight 1986, McCarthy 1986). The following key points were identified: The present day Irish fauna of approximately 14 000 species represented gradual accretion over 10 000 years and was highly mobile for the most part. It was generally depauperate in nature compared to Great Britain or north-western Europe and many groups were poorly represented, particularly among terrestrial insects. There are 'Lusitanian' elements in the Irish fauna i.e. insects that are absent from Britain today and have predominantly south-western European distributions. This might hint at south-westerly origins for the Irish fauna. However, there is also a distinct Arctic/Boreo-Alpine element to the Irish fauna. Both writers cautioned against an attempt to generalize as to the origin of the whole Irish fauna on the basis of a small number of insects - whether they were glacial relicts that survived in refugia or very early colonizers was still open to debate. Lack of known endemics, they believed, cautioned against the glacial refugia idea. Mobility, wind, accidental transport appeared to discount the need for a land-bridge. Speight highlighted the lack of particular groups of insects, most notably woodland species associated with oak, pine and 'old woodland' in general. He highlighted the need at that time for increased palaeoentomological research to understand if this lack of species represented an accurate picture of the Irish native woodland fauna and, if not, the timing and causal factors of the loss of these species (Speight 1986).

Both writers emphasized the need to examine the ecological requirements of insects, climate change and habitat diversity through time. Did insects arrive and become extinct quickly because their habitat requirements were not met, or disappear gradually or hang on in isolated pockets of certain relict habitats? What if anything could they tell us about their colonization/dispersal mechanisms? All this needed to be addressed by increased research.

* Email address: eireilly@tcdie

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Modern entomological research Since that time, modern entomological work has continued apace and has made a substantial contribution to our understanding of the origins of the Irish insect fauna. This has mostly been in the form of new surveys of different habitat types and revisions/reviews of particular families/genera of insects. Surveys Surveys have added numerous 'new' species to the Irish List and this paper can merely touch on a few of them. The beautiful damselfly Coenagrion lunulatum Char. was discovered during surveys of Co Fermanagh/Tyrone habitats (Nelson et al. 1989). At the time it was unknown in Britain and rare in western Europe. In Finland it is most common north of the Arctic Circle (Hamalainen 1984 cited in Nelson et al. 1989). It occurs in the sparse vegetation around high altitude lakes (85-310m) and is probably an example of a very early, mobile colonizer, overlooked until this time. Four species of Diptera were added to the Irish List during a survey of Blackditch Wood, The Murrough, Co Wicklow and Glendine, Co Waterford (Speight et al. 1990). Antichaeta brevipennis (Zet.) was discovered in thickly vegetated parts of wet woodland in the Murrough. It is a rare insect across Europe and on the British Red List for insects (Shirt 1987). Leucophenga maculata (Du.), discovered in Glendine, is an ancient deciduous forest insect, its larvae feeding in saproxylic fungi (Alexander 2002). It is known from various parts of Britain, from Scandinavia to Iberia and throughout Eurasia. At All Saints Bog, Speight surveyed the birch woodland and found a very important saproxylic fauna consisting of click and longhorn beetles (including Leptura quadrifasciata Linn. - only five records this century) and various tipulid flies (Speight 1990). The ground fauna was similarly indicative of ancient woodland, including the arachnid Araneus umbraticus Cl. and earthworm Eisenia eiseni Lev. The fauna as a whole appeared to indicate prolonged stable ground conditions. Speight noted that other beetles and hoverflies present as well as the rare grasshopper, Stethophyma grossum (L.), indicate that All Saints Bog is supporting a more complete, rounded fauna than equivalent sites in Ireland and can be considered a 'relict Irish biotope', in this case, birch woodland (Speight 1990).

A survey of turloughs in north Clare and south-east Galway by Good and Bulter (2001) added five new Staphilinid beetles to the Irish record and these beetles are now considered 'indicator species' of turloughs. Eight new beetle species were added to the record from a wide-ranging survey of east Clare, Galway and Offaly sites by coleopterists in 2003 (Regan and Anderson 2004) most notably many rare species associated with wetland habitats but also the longhorn beetle, Grammoptera ustulata (Sch.). It is classified as rare in Britain, and is an indicator of ancient woodland, mainly recorded from the midlands and southern England (Alexander 2002) Its find in old estate woodland in Charleville, Co Offaly is therefore of national importance. Revisions/Reviews Of the many insect groups reviewed or revised in the last 25 years the following is a mere sample to highlight the changes in our knowledge since 1983. O'Connor, Liston and Speight (1997) reviewed the sawflies (Hymenoptera: Symphyta) resulting in nineteen 'new' additions to the Irish list. The fauna is rich in 'secondary woodland' species (larvae host trees like birch, hawthorn and willow) but poor in oak and pine associates (excluding new introductions associated with spruce and larch). Two species with a Euro siberian distribution and not known from Britain include Cimbex conatus Sch., associated with alder, and Arge metallica (Klug), associated with birch.

Carabid beetles have been subject to detailed reassessment and revision (e.g. Speight et al. 1983, Anderson et al. 1997, Anderson et al. 2000). Two species, Bembidion argenteolum Ahr. and Agonum Iugens (Duff.), are not currently recorded in Britain but are both found in Scandinavia (www.habitas.org.uk/groundbeetles, Anderson 1985). They are

This content downloaded from 143.239.131.220 on Tue, 11 Sep 2018 10:35:16 UTC All use subject to https://about.jstor.org/terms Postglacial Colonization Conference: Mind the Gap 2006 characteristic of bare or sparsely vegetated ground and may have been more widespread in the early postglacial. Anderson et al. (2000) note that other boreal species are more widespread here than in Britain. They also note that the weakest categories of ground beetle species in Ireland are the Southern-temperate and Mediterranean-Atlantic groups - 62 such species occur in Britain but there are no Irish records (www.habitat.org.uk/groundbeetles). Morris, in his 1993 review of Irish weevils (Coleoptera: Curculionidae), confirmed 246 Irish species, an increase of 21 per cent from Johnston and Halbert's 1902 list of 203 (Morris 1993). Some possible 'Lusitanian' species include Otiorhynchus auropunctatus Gyll., which has only been recorded once from Scotland, otherwise is only known from southern France and Spain. Also, Barypeithes curvimanus du Val, a very rare Irish weevil is not recorded in Britain and only known from France. However, Morris noted that the taxonomy of some genera remained unworked at this time so it may be that unrecognized examples of these species exist elsewhere in collections and in nature (Morris 1993).

Summary This all too brief summary of modern entomological work highlights some important issues. Lots of species added to the Irish record in the last 25 years are described as 'new to Ireland'. In fact, this could be broken down into those that were 'overlooked' and those that are 'recent arrivals' the former being the majority because they weren't identified before or were not captured before. This is an important point because it means there is huge potential for many more species to be added to the record as a result of such work. The new total for the insect fauna of Ireland may be upwards of 16 000 species (O'Connor and Ashe 2000). Many of these findings have important biogeographical implications. The examples above give us some more evidence of the boreal elements within the Irish fauna but also others with a more southerly distribution today. Alexander describes the Irish saproxylic fauna as a 'relict hybrid of Atlantic temperate and Boreal fauna' (Alexander 2002). This appears to be true of other elements of the Irish fauna indicating multiple potential routes for the arrival of insects and, possibly, multiple phases of arrival, making the 'land-bridge' theory less and less likely. Ecological gaps still appear to exist with certain habitat types missing, reduced or modified. For example, Speight, in his review of Irish elaterid beetles notes a bias in the fauna towards species whose larvae feed in soils or on plant roots in open ground situations (Speight 1989a). Absent are species whose larvae feed in rotten wood of conifers and only five whose larvae feed on rotten wood of deciduous trees. The least well represented group are those who feed on the larvae of other saproxylics - less than ten per cent of the European elaterid fauna who prey on other saproxylics occur in Ireland. As many of these species are associated with oak, particularly long-lived oak, it is perhaps not surprising that they are absent (Speight 1989a). Over-mature, moribund and dead trees are critical to the larvae of many of the saproxylics these beetles feed on, which are themselves excessively rare or absent. These ecological gaps can perhaps only be addressed by palaeontomological research although there is certainly still scope for survey of particular rare relict habitat types and microhabitats, as the surveys of All Saints Bog and Charleville Wood show. So, has palaeoentomology helped to plug this gap and aid our understanding of colonization/dispersal mechanisms and extinctions? Palaeoecological research Palaeontomology involves the analysis of preserved insect fragments from waterlogged deposits such as lakes, bogs, woodland hollows and waterlogged archaeological deposits. It is a long established branch of palaeocology and environmental archaeology in Great Britain, primarily using Coleoptera (beetle) fragments (Reilly 2003). Patterns have

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emerged about relict landscapes from insect evidence including fen, woodland, sandy heaths, and early grasslands (e.g. Dinnin and Sadler 1999). This has resulted in increased knowledge about the origins of the British fauna including early colonizers and the many extinctions that have occurred during the last 10 000 years. A recently published review places the current extent of our knowledge of the Irish woodland beetle fauna in the context of British findings to date (Whitehouse 2006). This brief review will focus on woodland beetle fauna found during the course of palaeoenvironmental and archaeological research and includes some previously unpublished finding from the author's PhD research in the Killarney Woods, Co Kerry.

This suite of insects has many interesting features in common:

" all have fairly specific habitat requirements (stenotopic), e all are missing from the current Irish list of beetles (Anderson et al. 1997), There are also diverging aspects of their ecology and biogeography:

" some are currently missing from Great Britain and across north-western Europe, " where still extant in Britain and Europe, some have more northerly or more southerly distributions today, e other are currently widely distributed in Great Britain and north-western Europe.

Therefore, they provides many interesting clues both to the origins of the Irish fauna, possible colonization methods and its diversity through time. They also point to the fact that there were very particular factors affecting Irish woodlands thoughout the Holocene that perhaps resulted in the depauparate nature of this particular element of the Irish insect fauna today compared to Britain and north-western Europe (Reilly 2005, Whitehouse 2006). Few deposits from the very early postglacial period have been examined for insect remains. However, intertidal peats from Strangford Lough, Co Down, dated to 7320 - 6600 cal BC yielded Hylastes ater F. and Hylastes angustatus (Hbst.) (Rogers 2004 cited in Whitehouse 2006). These beetles are stenotopic saproxylics i.e. they only occur in dead wood, stumps and branches of pine. Hylastes was thought to be a recent introduction to Great Britain in plantation woodland but clearly these pine-dependents occurred here and in Britain at a very early stage and became extirpated in the following millennia (Whitehouse 2006). Other extirpated Early to Mid-Holocene species from pine layers in Sluggan Bog (6300 - 5500 cal BC) include Bothideres contractus F., Rhyncolus sculpturatus Walt and Rhyncolus elongatus Gyll. the latter two also found in similar deposits in Ballymacombs More, Co Antrim (Early-Mid Holocene in date) (Whitehouse 2006). Today, these beetles are confined to southern France, southern Germany, Poland and parts of southern Scandinavia. All are characteristic of old pine forest, the latter two being flightless, and were also recovered by Whitehouse in peat deposits in Thorne and Hatfield Moors, Humberhead Levels, Northern England (see Whitehouse 2006 for fuller discussion). Prostomis mandibularis (Fab.) was recovered in an oak plank trackway, in Derryville Bog, Co Tipperary, dated to 1606?9 cal BC (Reilly 2005). Apart from an outpost on islands off the south coast of Sweden, t i s mainly confined to woods in southern France, Italy, the Balkans and parts of Portugal. It is a stenotopic, flightless, saproxylic being found mainly in 'red rot' of fallen logs, branches and dead stumps of oak, pine and beech (Buckland 1979). It was found in mid-Holocene deposits in Thorne and Hatfield Moors and the Somerset Levels in Britain (Buckland 1979, Girling 1979, 1984; Whitehouse 1997, 2000, 2004, 2006), giving some indication of its former distribution.

A small number of species found in palaeoenironmental contexts are today rare, vulnerable or endangered in Britain but probably extinct in Ireland. Colydium elongatum (Fab.) was recovered from woodland peat, dated to c2000 cal BC, in Camillan Wood, Killarney, Co Kerry (Reilly 2008). This cylindrical, flightless saproxylic is predatory on other saproxylic

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beetle larvae and generally found in various deciduous tree species and on conifers on the Continent (Alexander 2002). It is classified as RDB3 (rare) in Britain. Teredus cylindricus 01. is another predatory, flightless saproxylic found in 'red rot' wood especially oak, often found in association with Lasius brunneus (Lat.) and anobiid beetles. In Britain, it is mainly confined to Sherwood and Windsor Forests and is classified RBD1 (endangered) (Alexander 2002). It was discovered in trackway wood associated with P. mandibularis in Derryville Bog, Co Tipperary, dated to1606?9 BC (Reilly 2005). Corticeus unicolor Pill. & Mitt. a flightless saproxylic, found in freshly dead birch, beech and oak wood, is also predatory on other saproxylic beetle larvae. It was recovered from woodland peat in Derrycunihy Wood, Killarney (c500 cal BC) (Reilly 2008). In Britain today, it is confined to two districts in the north Midlands and is classified as RDB3 (rare) (Alexander 2002). Lastly, there is a small group of beetle species that have been recovered in Irish palaeo-deposits that are not particularly rare in Britain today but have not been recorded previously in Ireland. Xyloborus dispar (Fab.) was recovered from a hazel hurdle in Derryville, Co Tipperary, dated to c200 BC and Xyloterus signatus (Fab.) was found in alder wood in a large trackway, also from Derryville Bog, dated to c600 BC (Reilly 2005). Acrantus vittatus (Fab.) and Scolytus mali (Bech.) were recovered from a house floor layers in Back Lane, Dublin, dated to the late 12th / early 13th century AD (Reilly 2003). Rhyncolus ater (Linn.), a cossinine weevil, was recovered from numerous archaeological and palaeoenvironmental deposits throughout Ireland, including the Neolithic-dated trackway of Corlea 9, Co Longford (Reilly 1996), various trackway and wood-peat deposits in Derryville Bog, Co Tipperary (Caseldine et al. 2001, Reilly 2005), two early-medieval trackways in Lemanaghan Bog, Co Offaly and 13th century floor layers from Back Lane in Dublin (Reilly 2003). In all cases it was found in association with oak, while in Britain today it is largely confined to old pine forest areas of Scotland (Alexander 2002). However, it has been found in fossil form in oak boles from Hatfield Moors, Humberhead Levels (Whitehouse 2004). In addition, recent research by the writer in Derrycunihy and Camillan Woods in Kerry has recovered other wood-dependent beetles including Xyloterus signatus, Scoyltus intricatus Ratz. and Hylesinus oleiperda (Fab.) in deposits dating from c800 cal BC to the late 17th century AD. Most of these beetles (except R. ater) are Scolytid bark beetles and in Britain range from fairly widespread to relatively restricted in their occurrence (Alexander 2002).

Summary

Of the 16 saproxylic beetles discussed above, five are primarily pine-dependents, all of which are extirpated from Ireland and four of which are from the earliest Holocene deposits examined so far in Ireland (Whitehouse 2006). Eleven are primarily deciduous woodland detrivores and primary wood-borers, two of which are definitely extirpated (B. contractus, P. mandibularis), three of which are probably extirpated (T. cylindricus, C. unicolor, C. elongatum) but what of the other six - mostly Scolytid bark beetles? The extirpated and rarest taxa are pine-dependents, 'red rot' deciduous wood dependents or predators on primary saproxylics, the very ecological groups suggested by Speight that would benefit most from palaoentomological research (Speight 1986, 1989a). Most are flightless and are termed Urwaldtiere or 'wildwood' inhabitants by palaeo entomologists in Great Britain (e.g. Buckland 1979). The presence of these ecological groups would suggest that many flightless beetles reached Ireland from early in the Holocene and were dispersed through easy availability of suitable habitats. The cumulative palaeoentomological evidence would appear to argue against the notion that forest insect species missing from the Irish record but whose presence might be expected on habitat association terms did not reach Ireland during the post-glacial. Loss of habitat is clearly the most important factor in their current absence from the Irish record. This loss was part of a complex of factors, including climate change, which would have had a knock-on effect on natural fire regimes in the Early to Mid-Holocene that kept forests open and supplied with dead wood (Whitehouse 2006). The loss of natural fire regimes from the mid-Holocene

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onward undoubtedly had a negative impact on native pinewood regeneration and, consequently, pine-dependent insect fauna (Whitehouse 2000, 2006). The loss of quality habitats and microhabitats through clearance of the forest floor, removal of dead wood and selective felling of standing dead trees would all have contributed to the loss of other specialist saproxylics (Buckland 1979, Speight 1989b, O'Sullivan and Kelly 2006). The group of Scolytid beetles (not necessarily rare in Great Britain but missing from Ireland), would appear to have succumbed to some of the above factors but were also probably casualties of the nature of forest clearance in Ireland, which appeared to have been more complete during the seventeenth to nineteenth centuries than elsewhere in Britain or Europe (Hall 1995, Cole and Mitchell 2003). A combination of expanding population in the pre-Famine years, the grubbing up of hedgerow trees, and the removal of scrub would have affected the somewhat more mobile Scolytids by increased fragmentation of their habitats and removing 'corridors' or links between patches of woodland (cf. Dennis 1997). Replacement of native trees with other wood species in the nineteenth century coupled with woodland management practices that discouraged dead wood habitats also meant that as primary saproxylics disappeared, the prey-predator chain broke down and consequently other species dependent on them disappeared (Watts 1984, Speight 1989a, O'Sullivan and Kelly 2006). However, the brief review above of modern entomological work suggests there is the possibility that some rare saproxylics are still present in relic woodland patches, as the finding of Grammoptera ustulata in Charleville Wood, Co Offaly demonstrated (Regan and Anderson 2004). Questions Answered? Undoubtedly, the state of our knowledge is much improved due to the greatly increased level of modern entomological and palaeoentomological research. Indeed, the points raised above at the 1983 meeting by Speight (1986) and McCarthy (1986) have proven to be an excellent road map in terms of where gaps are being filled and what still needs to be addressed.

The actual mechanisms of colonization are still a matter of speculation but it is clear that flightless and flighted, highly-specialized saproxylic beetles, to name just one group, made it to Ireland in numbers. In the absence of land-bridges and clearly proven 'refugia' in southern Ireland, Whitehouse (2006) suggests wind, water and, possibly, human transport were the main modes of colonization and dispersal, also suggested in 1983 by Speight (1986) and McCarthy (1986). Certainly winged insects could have been transported on the wind and this continues to be the way many small islands are colonized by insects around the world (e.g. Simberloff 1978). Human transport of insect is clearly proven from palaeoentomological research in Viking Age Iceland and Greenland (Buckland et al. 1983, Sadler 1990). However, wood rafting, in particular, is the most likely mode of transport for the wingless, specialist saproxylics discussed above. Whitehouse (2006) points out that early Holocene forests across Europe would have contained abundant dead wood and European rivers would have been full of large driftwood trees. She suggests this could explain the 'synchronous arrival of many invertebrates and plants' to both Britain and Ireland (Whitehouse 2006). This would also fit with the model of tree migration into Ireland for certain tree species suggested by Mitchell (this volume).

It is clear that more Early Holocene sites need to be examined as well as more long sequences through suitable deposits to get a clearer picture of the changing nature of the Irish native insect fauna throughout the Holocene. In particular, a greater geographical spread of sites would be valuable to examine the direction of colonization pathways.

In addition, it is clear that modern entomological surveys of relict habitats and sites of high conservation value will continue to add to our understanding of the Irish insect fauna. This requires sensitive and sustainable methodologies to ensure we don't negatively affect the very fauna we are trying to understand.

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Acknowledgements I would like to acknowledge the following people who over the years have helped me through useful discussions, aid with identifications and general encouragement: Dr Keith Alexander, Professor Paul Buckland, Dr. David Smith, Dr Martin Speight and Dr. Nicki Whitehouse. I would also like to acknowledge all my archaeological colleagues from whose sites most of the palaeoentomological data is derived. In reference to the compiling of this paper, I would particularly like to acknowledge the help of my PhD Supervisor, Dr Fraser Mitchell and my fellow palaeoecology PhD student in the department, Bettina Stefanini. Thank you to Professor Peter Woodman for inviting me to give the paper at the conference and to the editor, Julia Davenport, for her patience during the editorial process. I would also like to acknowledge the helpful comments of an anonymous reviewer. References

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