Turloughs – Ireland’s unique wetland habitat

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Review

Turloughs – Ireland’s unique wetland habitat ´ . O Connora,b,d, E. Regana,b,e, C.E. Coxone, M. Sheehy Skeffingtona,*, J. Morana,b,c, A N.E. Scottf, M. Gormallyb a

Department of Botany, NUI, Galway, Galway, Ireland Applied Ecology Unit, Centre for Environmental Science, NUI, Galway, Ireland c Teagasc, Athenry, Co. Galway, Ireland d National Parks and Wildlife Service, 7 Ely Place, Dublin 2, Ireland e School of Natural Sciences, Trinity College Dublin, Dublin 2, Ireland f ‘Seagal’, Ballinacourty, Clarinbridge, Co. Galway, Ireland b

A R T I C L E I N F O

A B S T R A C T

Article history:

Turloughs are karst wetland ecosystems that are virtually unique to Ireland. Flooding annu-

Received 23 February 2006

ally in autumn through springs and fissures in the underlying limestone and draining in the

Received in revised form

springtime, often through the same fissures or swallow-holes, they have been described as

30 June 2006

‘temporal ecotones’. Over 300 have been documented. They are priority habitats in the EU

Accepted 30 June 2006

Habitats Directive and support a variety of wet grassland and fen type vegetation. Though

Available online 27 September 2006

the vegetation has been recorded and mapped for over 80 turloughs, records for invertebrates are more sporadic. Characteristic species include some aquatic species-often benefit-

Keywords:

ing from the absence of fish-, and many wetland terrestrial species, including carabid beetles

Biodiversity

that are rare on a European scale. Due to their shallow nature and the full vegetation cover of

Conservation

the basin, turloughs can host internationally significant numbers of visiting winter wildfowl,

Karst

particularly whooper swans. The variety of plant and invertebrate communities between

Turloughs

turloughs is primarily due to different hydrogeomorphological characteristics, but also

Wetlands

depends on the range of grazing practices on turloughs. Since these often vary within a tur-

Birds

lough basin, this helps maintain within-turlough biodiversity. The main threat to turloughs

Hydrology

in the past was drainage, but pollution by nutrients is also now potentially detrimental. How-

Invertebrates

ever, a more recent and important threat may be the cessation of farming within turloughs.

Land use

As potentially threatened wetlands of European importance, turloughs require a full inven-

Review

tory of their biodiversity and the factors affecting it. The collation here of all literature con-

Vegetation

cerning turloughs will provide a basis for an integrated approach to future research on turloughs that is essential for a full understanding of these complex ecosystems. Ó 2006 Elsevier Ltd. All rights reserved.

Contents 1.

Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 266 1.1. Background . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 266

* Corresponding author: Tel.: +352 91492682. E-mail address: [email protected] (M. Sheehy Skeffington). 0006-3207/$ - see front matter Ó 2006 Elsevier Ltd. All rights reserved. doi:10.1016/j.biocon.2006.06.019

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2.

3.

4.

5.

6.

7.

8.

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1.2. What are turloughs? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1.3. Turlough distribution. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hydrology and geomorphology of turloughs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2.1. Swallow holes and estavelles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2.2. Depth and duration of flooding. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2.3. Mechanisms for turlough flooding . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2.4. Origin and geomorphology of turloughs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2.5. Calcium carbonate deposition. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Vegetation of turloughs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3.1. Introduction. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3.2. Plant species of turloughs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3.3. Vegetation studies – phytosociology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3.4. Vegetation studies – turlough mapping . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3.5. Conservation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Invertebrates – aquatic fauna . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4.1. Introduction. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4.2. Turlough communities . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4.3. Species adaptation to the turlough environment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4.4. Conservation – rare species. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Invertebrates – terrestrial fauna . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5.1. Introduction. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5.2. Species occurring in turloughs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5.3. Environmental factors affecting terrestrial invertebrates . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5.4. Conservation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Vertebrates . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6.1. Fish and amphibians . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6.2. Mammals. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6.2.1. Conservation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6.3. Birds. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6.3.1. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6.3.2. Frequent visitors to turloughs. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6.3.3. Site ecology and evaluation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6.3.4. Conservation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Turlough management and conservation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.1. Background . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.2. Drainage . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.3. Nutrient/trophic status . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.4. Grazing . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Conclusion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Appendix 1. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

‘‘Horse and boat races are held on the same ground, but at different seasons’’—Dutton (1824) re Turloughmore, Tuam, Co. Galway (in MacGowran, 1985).

1.

Introduction

1.1.

Background

Turloughs are virtually unique to Ireland and as such are listed as priority habitats in Annex I of the EU Habitats Direc-

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tive (EEC, 1992). They are wetlands, a type of habitat that is increasingly rare throughout Europe due to land drainage and intensification of land-use (Merne, 1980; Stevenson and Frazier, 1999; Moser, 2000; Thompson and Finlayson, 2001). Turloughs occur in the karst limestone region of Ireland and although unique, their conservation is linked with other European limestone-related Annex I habitats. Thus an understanding of this Irish ecosystem will benefit the conservation of other EU habitats, especially in Slovenia, which has extensive karst areas with poljes that closely resemble turloughs. Yet what is known about turloughs is

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disparate and focused on specific topics rather than on turloughs as integrated ecosystems. Within the Water Framework Directive (EC, 2000), Ireland must review likely threats to turloughs as part of water catchment systems and this requires an understanding of turlough ecology. Thus it is important to draw together existing knowledge to evaluate gaps and areas for future research, in order to develop policies for turlough conservation. This paper reviews all the main published literature concerning turloughs as well as several unpublished theses and reports and summarises the salient management and conservation issues. Nomenclature follows Stace (1997) for vascular plants and Smith (2004) for mosses. The authorities for invertebrates are included in the text and follow Moorkens and Speight (2001) for molluscs; Ashe et al. (1998) for aquatic insects; Anderson et al. (2000) for carabid and Lott (2006) for staphylinid beetles; Rozkosˇny´ (1987) for dipterans.

1.2.

What are turloughs?

The essence of turloughs is in their hydrology. They can be defined as depressions in karst areas, seasonally inundated mostly by groundwater and supporting vegetation and/or soils characteristic of wetlands (Working Group on Groundwater, 2004). They usually flood through underground passages and springs in autumn, when rainfall exceeds evapotranspiration, form a lake for several months in winter (Plate 1) and empty underground through swallow holes in springtime (Coxon, 1987a). There may also be sporadic rises at other times in response to high rainfall. Some turlough basins retain standing water in channels, pools or small lakes when flooding subsides. Although they harbour an aquatic fauna, they are not true lakes, since most drain in the summer, revealing fen or grassland vegetation (Plate 2) which is frequently grazed by livestock. Reynolds et al. (1998) describe them as ecotones, since they are transitional between aquatic and terrestrial systems, the transition being temporal as much as spatial. The name ‘turlough’ is usually interpreted from the Irish, tur meaning ‘dry’ and lough meaning ‘lake’, but could also be interpreted differently (Royal Irish Academy, 1998). The current spelling in Irish is turlach. Joyce (1869) refers to -lach as a suffix and interprets turlach instead as meaning ‘a dried-up place’. The genitive form: turlaigh, with -lach as a suffix, occurs in Irish place names. On the Irish-speaking island of Inis Mo´r in the Aran Islands, there is an area called Ceathru´ an Turlaigh (Turlough Quarter). Parish names such as Killaturly (Mayo) and Ballinturley (Roscommon) are similar examples (all documented turloughs are listed by county in Appendix 1). It makes sense that turloughs were originally distinguished for their ability to dry up, not for their status as lakes, as it would be the grazing which was valued. Indeed, MacGowran (1985) says turloughs are as much ‘vanishing fields’ as ‘vanishing lakes’.

1.3.

Turlough distribution

Within the very extensive Carboniferous limestones of Ireland, turloughs occur mostly in areas of thin glacial drift and are restricted to areas of grey calcarenite that has a greater

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degree of karstification than other limestones due to its purity and well-developed bedding (Coxon, 1987a). Other seasonal lakes in comparable karstic regions are not considered to be identical to turloughs, due to their differing hydrology, seasonality, size or geomorphology (Coxon, 1986). These include the Slovenian poljes, the Breckland Meres on chalk in England and lakes/depressions in the North American karst regions (Reynolds, 1998; Goodwillie and Reynolds, 2003). The poljes appear to be the most similar to turloughs, but are much bigger, covering valleys several km long and although they flood seasonally, snow melt–water adds to the springtime water-level rise. However, on a recent visit to Slovenia, two of the authors visited much smaller wetlands that appeared to be more like turloughs. Reynolds et al. (1998) refer to the lacs myste´rieux in Eastern Canada and a seasonal water body in karstic Catalonia, but affirm that only in Wales is there a hydrological feature similar to Irish turloughs at Pant-y-llyn (see Campbell et al., 1992; Blackstock et al., 1993). Early topographical and statistical surveys referring to turloughs in Ireland are reviewed by MacGowran (1985). Coxon (1986); Coxon (1987b), selecting for turloughs >10 ha, identified 90, of which 30 had been lost through drainage schemes from the 19th century to the present. Other hydrologically active sites (sometimes <10 ha), have since been identified (Fogg and Kelly, 1995; Southern Water Global, 1998; Reynolds et al., 1998; Goodwillie and Associates, 2000). These sites, along with those from Reynolds (1985a) and some noted by the authors, are in a database in the Environmental Change Institute, NUI, Galway. Another database of >300 (based on these, GSI and other records) is held by the Geological Survey of Ireland (GSI), Dublin. This is a baseline list and not all have been formally verified. These are plotted in Fig. 1 in relation to limestone bedrock which shows how turloughs occur either on, or immediately adjacent to, areas of pure bedded limestone (see Coxon, 1987a). The greatest density of turloughs is in the western third of Ireland. Here rainfall more frequently exceeds evapotranspiration and the degree of exposure of the limestone (due to shallower glacial drift) is more extensive, facilitating karstification. The biggest turlough in Ireland, appropriately named Turloughmore (turlach mo´r = big turlough), used to be in the Clare River catchment in Co. Galway, near Tuam, appearing on Petty’s 1685 map of Galway. Stretching some 400 ha in winter, it was part of a complex of ca. 770 ha before the first drainage of 1847 (D’Arcy, 1983). Rahasane Co. Galway, (ca. 260 ha) is now the largest (Goodwillie, 1992).

2.

Hydrology and geomorphology of turloughs

2.1.

Swallow holes and estavelles

Turloughs are karst features and their flooding is the surface manifestation of ground water that is linked characteristically via swallow holes to subterranean karst geomorphological features. When the same hole acts both as a spring through which the turlough fills in the autumn and a swallow hole through which it empties in the springtime, it is termed an estavelle. Swallow holes were identified by Coxon (1986) as present in 80 out of the 90 sites studied. They are mostly lo-

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Fig. 1 – Distribution of turloughs in Ireland in relation to limestone bedrock, based on the GSI database (see Appendix 1) (courtesy of the Geological Survey of Ireland – GSI).

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Plate 1 – Ballinacourty turlough, January, 2005.

cated around the periphery of the basin, where bedrock is at or close to the ground surface and at the edge of the unconsolidated deposits such as peat and marl which occupy the turlough floor, often to a thickness of several metres. In a small distinctive group, the swallow holes occur in the centre of the turlough or are scattered over the floor as depressions or collapses in the deposits (Coxon, 1986, 1987b) (see Plate 2). Occasionally the turlough acts as a sink for a stream or river coming from outside the basin (Coxon, 1986; Goodwillie, 1992; Goodwillie and Reynolds, 2003). However, even sites with a major surface water input also contain groundwater inputs, e.g. Rahasane is fed not only by the Dunkellin River but also by a series of large springs.

2.2.

Depth and duration of flooding

Several hydrological factors contribute to defining turlough characteristics. These include hydroperiod (length of flooding time), time of year when flooding occurs and recedes, flood frequency (how many floodings a year) and flooding depth (Southern Water Global, 1998). In addition, severe or catastrophic flooding may occur in some turlough systems when winter rainfall exceeds a given maximum, such as occurred in the Gort lowlands, SE Galway in the 1990s (Southern Water Global, 1998). The depth of flooding in the turloughs studied by Coxon (1987b) varied from 0.5 m to 6 m, with median 1.5 m. These depths were deduced from the upper limit of the epiphytic or epilithic moss Cinclidotus fontinaloides. Most turloughs contain an upper zone where rock surfaces are dominated by

Cinclidotus and a lower one where it is replaced by the moss Fontinalis antipyretica or sometimes by a tufaceous crust. Where Cinclidotus occurs to the base of the turlough, (35% of the 60 sites) it is considered to indicate a shorter flood duration, the turlough perhaps also filling and emptying more frequently (Coxon, 1986, 1987b). Most of the latter are shallower turloughs, flooding less than 1 m, but the small distinctive group mentioned in 2.1 above, are deeper with an apparently shorter duration of flooding and have compound or undulating floors rather than the typical turlough flat floor. They contain unsorted glacial drift, sand, silt or clay, rather than peat or marl (Coxon, 1987b). In the Gort lowlands which have been studied in some detail, there is a variety of lake–turlough systems: some empty each summer, others only fully empty in dry summers and others always retain some water (Coxon and Drew, 1998). This is linked with the mode of filling and emptying: some fill via surface waters and show large water level fluctuations (such as Hawkhill, with an annual range of 15 m, and Blackrock/Peterswell, which can change its water level by up to 9 m in 48 h) while others, filling via springs or estavelles usually have a much narrower range in water level (3–5 m) (Johnston and Peach, 1998).

2.3.

Mechanisms for turlough flooding

Detailed studies of turlough hydrology and the mechanisms of flooding are rare. In south-east Mayo, turlough water levels and flood timing were found to coincide reasonably well with the water table as determined from a borehole over a 2 year

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Plate 2 – Ballinacourty turlough, May, 2005.

period (Coxon, 1986; Coxon and Drew, 1986). Turloughs were not randomly distributed through the aquifer, but were preferentially located along lines of concentrated flow within it. These higher permeability zones were identified by troughs in water table maps and by high water velocities (of the order of 100 m h1) determined by tracing experiments (Coxon, 1986; Coxon and Drew, 1986). The exact nature of the zones is uncertain: they may correspond to actual cave conduits or to solutionally widened fissure networks. The hydrology of the Gort lowlands is rather different. Whereas other turloughs in most of the Irish limestone lowlands are fed only by limestone-derived water, the karst aquifer in the Gort lowland is fed by several sinking rivers originating on Devonian sandstones on the Slieve Aughty uplands. Thus the area contains major karst conduits associated with these chemically aggressive water inputs. An intensive study of turloughs in the region over 2 years (the Gort Flooding Study: Southern Water Global, 1998) showed that some of the turloughs are fed predominantly by these conduits and their flooding has been modelled successfully using a pipe network model (Southern Water Global, 1998; Johnston and Peach, 1998). Apart from the major conduits in each region, there is also a degree of distributed flow within the limestones and a very significant component of shallow flow in the epikarst (shallow karst at the surface of a limestone rock system), which provides the main water input to some turloughs (Coxon and Drew, 1998; Drew, 2003). While the majority of turloughs occur on the western Irish limestone lowlands, a few are found on the Burren plateau. Williams (1964) made a distinction between upland turloughs

(e.g. Lough Aleenaun) which flood at any time of year following a few days of heavy rain, due to rapid runoff into a basin with a semi-permeable floor where flood water is temporarily held up, and lowland turloughs where the flooding is more regularly seasonal. Drew (1990) noted that the depressions at L. Aleenaun and Carran, are underlain by low permeability cherty limestones, glacial drift and lacustrine marl, and that flooding is due to temporary ponding of spring water entering along the flanks. Thus the mechanism for filling and emptying of the turloughs is complex, reflecting the nature of the karst aquifers feeding them. To some extent they can be viewed as water-table phenomena, filling as the water table rises and the zone of saturation intersects the ground surface and emptying as the water table falls. Alternatively they can be viewed as pressure release points along an underground pipe network, filling when the discharge through the conduit and the pressure increases and water is forced up to the ground surface at a particular location, and emptying when the pressure falls again. The role of shallow epikarstic water inputs is also likely to be significant in many instances.

2.4.

Origin and geomorphology of turloughs

Lack of detailed knowledge of turlough basin structure precludes definitive conclusions on turlough origins, but probably most if not all are in bedrock hollows and not merely in hollows in glacial till (as suggested by Williams, 1964; Coxon, 1986). The question arises as to how these bedrock hollows were formed. There are two main theories as to how they

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form; that they are glacial in origin (e.g. Williams, 1964, 1970; Southern Water Global, 1998), or that they are pre-glacial, having developed as a result of tertiary karstic processes (Coxon, 1986; Coxon and Coxon, 1997). As evidence of a pre-glacial tertiary karst landscape continues to be found (Mitchell, 1980, 1985; Drew and Daly, 1993; Coxon and Coxon, 1997; Southern Water Global, 1998; Karst Working Group, 2000; Simms, 2003), it is thought likely that at least some turloughs developed as karst depressions in the tertiary and were possibly subsequently modified by repeated glacial scouring and/or till deposition. Indeed, some turloughs may have formed by karstic processes in tertiary times, others by glacial processes in the more recent quaternary period (Coxon and Coxon, 1997). The post-glacial history of turloughs also lacks data. However, it is believed that seasonal emptying has not existed for all turloughs throughout their existence as flooded areas. At least 51% of the 90 sites surveyed by Coxon (1986, 1987b) contains pure calcareous marl or lacustrine chalk, indicating that they were permanent lakes at some time in the past. Marl was found in most turloughs surveyed in the Gort lowlands, sometimes forming an impermeable seal on the base of the turlough, broken occasionally by a swallow hole or estavelle (Southern Water Global, 1998). The period of marl deposition was dated at one site to 10,000–9000 years B.P. (Coxon and Coxon, 1994). The reason for the change from a permanent water body to seasonal flooding is unclear. It does not appear to reflect climate change or a change in hydrological base level, but may have involved a combination of factors, such as silting up of the depression, or increasing karstification in postglacial times, resulting in the development of swallow holes and a zone of higher permeability within the aquifer associated with the turloughs. The presence of mushroom stones (limestone rocks eroded below the top surface by standing water, forming overhanging lips of rock) is evidence that a water body has remained for long spells at one particular level marked by the ´ Riaga´in, 1972; Goodwillie and Reynolds, lip of the rock (O 2003; Dunne and Feehan, 2004). They have been found around the periphery of several turloughs (e.g. Caranavoodaun and Coole, SE Galway), but require more research.

2.5.

Calcium carbonate deposition

In addition to the lacustrine marls referred to above, carbonate deposition also occurs in turloughs at the present day: one of their distinctive characteristics is the whitish coating of calcite crystals seen on vegetation after emptying in the springtime. Annual calcite deposition rates estimated from the chemical data for four turloughs in south-east Mayo were of the order of 500–700 g m2 year1, comparable to reported deposition rates for permanent lakes (Coxon, 1994). A study of their water chemistry over two seasons suggested that carbonate deposition is due predominantly to supersaturation caused by the loss of carbon dioxide to the atmosphere from water entering the turloughs. Calcite precipitation appears to occur throughout the winter rather than being restricted to the springtime before emptying. Thus the deposition was attributed to physico-chemical processes, with biological influences being only minor in causing supersaturation, although further studies are required to investigate the role

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of phytoplankton in triggering calcite precipitation (Coxon, 1994). In the absence of detailed nutrient water chemistry for turloughs, they are generally assigned a trophic status in relation to the plant communities they support (Goodwillie, 1992, 2001; Working Group on Groundwater, 2004). However, it is also considered that extreme calcareous waters could immobilise soluble phosphate to some extent and that such turloughs tend to be oligotrophic (Goodwillie, 2001; Southern Water Global, 1998). The trophic status of turloughs is complex, however and cannot be assessed on groundwater properties alone (see Section 7.3).

3.

Vegetation of turloughs

3.1.

Introduction

Turloughs are grass- or sedge-dominated basins which sometimes have a marsh or occasionally a permanent pond in the centre. They are notable for the absence of trees or shrubs (Praeger, 1932) which are controlled not only by grazing but also by duration of flooding (Praeger, 1932; Goodwillie, 1992, 2003). In the exceptional flooding of 1990 around Gort, the high water level killed juniper Juniperus communis, gorse Ulex europaeus, ash Fraxinus excelsior and blackthorn Prunus spinosa around the edge of affected turloughs, though other species, such as purging buckthorn Rhamnus cathartica, revived late in the summer season (Goodwillie, 1992, 2003). The presence of the black moss Cinclidotus fontinaloides on walls and surrounding shrubs is a regular indicator of the location of a turlough with its alternating wet-and-dry regimes (Praeger, 1932; Coker, 1993). Growth experiments have suggested that Cinclidotus does not necessarily have a preference for variably wet conditions, but is excluded from a more aquatic habitat by competition with more vigorous aquatic mosses such as Fontinalis antipyretica (Allott, 1976). Though these two moss species are the only plants listed as characteristic of turloughs in the Habitats Directive manual (Roma˜o, 1999), many Drepanocladus species recorded in turloughs and fens by Ivimey-Cook and Proctor (1966) are nationally rare and require further work (vide Hill et al., 1992; Holyoak, 2003). Calliergon trifarium, a relict montane species, is recorded only in two sites in Ireland, one a turlough (Perry, 1983; Hill et al., 1992) The extensive bryophyte lists of Ivimey-Cook and Proctor (1966), contain very few liverworts which suggests that these are almost absent from turloughs. Lichens, generally epiphytic or epilithic in turloughs, have not been documented specifically for turloughs.

3.2.

Plant species of turloughs

The flora of turloughs includes rare wetland species such as the fen violet Viola persicifolia and the shrubby cinquefoil Potentilla fruticosa, common on some upper margins (Webb and Scannell, 1983; Pullin, 1986) and the annual northern yellowcress Rorippa islandica, found on bare muds of late-drying turloughs (Scannell, 1973; MacGowran, 1979; Goodwillie, 1995). The only record to date of Callitriche palustris for Ireland is from Coole Lough (Scannell and Jebb, 2000; Goodwillie, 2003). Plants of some species can exhibit different growth forms to those found on normal ‘dry’ soil, e.g. Ranunculus repens,

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Potentilla reptans and Mentha aquatica (White, 1984). The turlough form of R. repens shows morphological variation in leaf dissection (White, 1984; Lynn, 1998; Lynn and Waldren, 2001, 2003) and root aerenchyma (White, 1984) as well as physiological differences in response to experimental flooding (Lynn and Waldren, 2002, 2003). Most spectacularly, when submerged in the very deep Hawkhill turlough, it has been noted to develop 4 m long etiolated shoots floating up through the water, but disappear when the water subsides (B. MacGowran, pers. commun.). Aquatic plants including charophytes can be found in turloughs with summer ponds or pools (Goodwillie, 1992; Stewart and Church, 1992; Roden, 2001). Those with basin floors that dry out late are characterised by open mud with ruderal species, such as the annual knotgrasses (Persicaria spp.) and water purslane Lythrum portula. If the mud remains wet, it favours the rare annual mudwort Limosella aquatica (Curtis et al., 1985; Goodwillie, 1992). Such weed communities may also be a function of heavy grazing and trampling (Goodwillie, 1992; Nı´ Bhriain et al., 2002). The basins of some shallow turloughs, on drying in spring are strewn with carpets of drying filamentous algae, referred to as ‘algal paper’ (Scannell, 1972; Allott, 1976; Reynolds, 1983).

3.3.

Vegetation studies – phytosociology

Several vegetation studies refer to single or localised turlough sites in SE Galway and Clare (Louman, 1984; O’Gorman, 1984; Vink and van Kruysbergen, 1987; Nı´ Bhriain, 1999; van Ravensberg and van der Wijngaart, 2000) or a group of turloughs as part of a catchment study (Lockhart, 1984; Lockhart, 1985; Goodwillie et al., 1992). However, Ivimey-Cook and Proctor (1966) and O’Connell et al. (1984) describe the phytosociology of the vegetation of turloughs in detail, as part of larger vegetation studies of the Burren and Irish wetlands, respectively. In the last two cases, the main plant communities described are also found on lake-shores, fens or marshes, demonstrating that broadly speaking, turlough vegetation is not unique. Turlough plant communities grade into those surrounding the basin, such as woodland, limestone heath or pasture, all of which may be flooded for many days in any year (Goodwillie, 1992, 2003). Most turlough wetland communities can be classified into two main phytosociological classes, the Scheuzerio-Caricetea fuscae, and the Plantaginetea majoris, describing small-sedge, and disturbed habitat communities respectively. A third class, the Littorelletea, describes lakeshore communities found on the margins of more permanent water bodies within turloughs (Ivimey-Cook and Proctor, 1966; O’Connell et al., 1984). Although these authors ascribe the differences in occurrence of the plant communities in part to inundation frequency and substrate saturation, the two communities can vary in dominance within the same flood-zone in a turlough, separated only by a wall or fence (Nı´ Bhriain et al., 2002), suggesting that some of the vegetation differences are due to variations in land-use within the same turlough. The alliance Caricion canescenti-nigrae (or the related Caricion davallianae) and the Agropyro-Rumicion crispi within the above-mentioned classes respectively describe the majority of plant communities of turloughs (Ivimey-Cook and Proctor, 1966; O’Connell et al., 1984). The

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Caricion davallianae and the Lolio-Potentillion are the phytosociologically related alliances listed in the Habitats Directive Manual (Roma˜o, 1999) for turloughs. Although so far only described for the NE Clare–SE Galway region (Ivimey-Cook and Proctor, 1966; O’Connell et al., 1984) there appear to be discrete plant sub-communities of the broader grassland/fen vegetation which occur only in turloughs. Specifically the Burren Carex demissa nodum (the Carex panicea–Carex flava agg. community of O’Connell et al. (1984)) and perhaps the Potentilla anserina–Drepanocladus lycopodioides nodum of Ivimey-Cook and Proctor (1966) are almost exclusively described from in turlough basins.

3.4.

Vegetation studies – turlough mapping

Goodwillie’s survey of turlough vegetation (1992) is the most comprehensive to date, describing and mapping 32 separate vegetation units in Coxon’s (1987b) 60 undrained turloughs. However releve´s were rarely taken and these units can only be tentatively related to the phytosociological communities described above. Later, as part of the Gort lowlands flooding study (Southern Water Global, 1998), Goodwillie et al. (1997) include smaller turloughs (and other wetlands) within SE Galway and NE Clare. A total of 36 vegetation types influenced by flooding are described. The detailed vegetation maps in both studies highlight the variety in turlough plant communities; each turlough supporting a particular combination of about a dozen communities. Goodwillie (2003) summarises these into 24 principal communities. Vegetation patterns are related to flooding. Praeger (1932) was among the first to describe the zonation of vegetation, remarking on the gradual change from dry soil species on the edge to wetland species in the centre. The plant communities encircle the basin in roughly concentric zones, according to different submergence spells (Ivimey-Cook and Proctor, 1966; O’Connell et al., 1984; Goodwillie, 1992; Goodwillie et al., 1997). The vegetation of the uppermost zone most closely resembles the adjacent grassland pasture, as it is rarely flooded for long (Goodwillie, 1992). Flooding undoubtedly controls vegetation zonation, but it is now clear that duration of flooding (hydroperiod), rather than depth, most affects vegetation. However, both period of inundation and ‘release’ date can vary considerably from year to year (Goodwillie, 2003) and some turloughs are susceptible to out-of-season flooding, even in summer (Southern Water Global, 1998). Goodwillie (2003) suggests that less typical turlough species, such as flag iris Iris pseudacorus and smooth rush Juncus effusus may not tolerate such summer inundation.

3.5.

Conservation

The Red Data Books list several turlough species: alder buckthorn Frangula alnus, fen violet Viola persicifolia, northern yellow-cress Rorippa islandica, mudwort Limosella aquatica, shrubby cinquefoil Potentilla fruticosa and the charophyte Nitella tenuissima (Curtis and McGough, 1988; Stewart and Church, 1992). Though none of these species is confined to turloughs, their presence in a turlough adds to its conservation value. Only the water starwort Callitriche palustris has so

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far only been recorded in turloughs in Ireland (Goodwillie, 2003). All are wetland species and thus are at risk in a European context through habitat loss. Though the main plant communities of turloughs occur elsewhere, the small-sedge communities have some elements that appear to be almost unique. These require more permanently wet soils than do grassland communities and so are rarer and more susceptible to drainage. The mosaic of different plant communities, often individual to one basin unit, also enriches turlough biodiversity and conservation value. Maintenance of the unique hydrology of a turlough basin is of primary importance in conserving the vegetation, but agricultural land-use is also essential for maintaining vegetation diversity (Nı´ Bhriain et al., 2002, 2003).

4.

Invertebrates – aquatic fauna

4.1.

Introduction

The factors determining the aquatic fauna of turloughs are incompletely understood, although flooding is generally accepted to have a major influence (Foster et al., 1992; Reynolds, 1982, 1996a,b, 2000). The way flooding regimes vary from year to year, causes the freshwater fauna to be unpredictable (Reynolds, 1982, 1996a, 2000). With each new flood, new species may colonise a turlough and new competitive interactions occur (Reynolds, 1982). Irregular flooding, combined with the oligotrophic nature of many sites, also results in sparse populations (Grainger, 1991; Reynolds, 1996b, 2000). Low productivity can also lead to slow growth rates in invertebrates, as found in aquatic snails in Lough Geala´in and Knockaunroe (Byrne et al., 1989). Turlough aquatic fauna is also highly spatially variable, in terms of both species diversity and abundance (Reynolds, 1997, 2000). Even adjacent sites can have very different flooding regimes and consequently, discrete aquatic assemblages (Reynolds, 1982). Sampling strategies can affect species catches; the use of box sampling methods for turlough aquatic invertebrates can be more efficient and accurate than the pond net (O Connor et al., 2004).

4.2.

Turlough communities

Despite these adversities, there is a suite of aquatic invertebrates that regularly occurs in turloughs. Many of these species are widespread and common elsewhere (Lansbury, 1965), but all are well adapted to the temporary nature of turloughs (Reynolds, 1982). Typical groups include flatworms, gastropod molluscs, cladocerans, copepods, ostracods, mayfly nymphs, and beetles (Reynolds, 1982, 1985a, 1996a, 1997, 2003), most of which are commonly found in small ponds (Reynolds, 2000). Fully aquatic species such as snails and the crustaceans Gammarus and Asellus may survive in refugia and reproduce rapidly upon flooding. The dominant turlough snails are Radix baltica and Bithynia tentaculata (Donaldson et al., 1979; Tattersfield, 1998). Mayfly and caddis-fly diversity is restricted, however, when compared to more permanent water-bodies. Byrne (1981) found that Cloeon simile normally a bivoltine mayfly species in Ireland, only succeeded in producing one generation per year in two upland turloughs studied and appeared to over-winter as diapausing eggs, a

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response attributed to the fluctuations and low productivity of the water. Reynolds (1985b) recorded 20 cladoceran species (six of which were new to County Clare) in two turloughs adjacent to Mullagh More, and 15 in a larger series of 28 turloughs studied in early spring (Reynolds et al., 2004). Although cladoceran and copepod diversity in turloughs is high overall, species richness in individual turloughs is often relatively low, with one or two species dominating many sites (Duigan, 1988, 1992; Reynolds, 1997; Reynolds and Marnell, 1999). The low species richness may represent dominance of the first colonists from the ephippial egg bank (Reynolds et al., 2004). Abundance is also low in many turloughs, such as those around Mullagh More (Reynolds, 1982). Phytoplankton, the main food of many of these crustaceans, is scarce in turloughs (Reynolds, 1982). Low phytoplankton abundance is probably a result of the synchrony between the time of highest photosynthetic activity and the dry phase of turloughs. Phytoplankton is an area of turlough ecology which requires further research. The corixid bugs (Corixidae) have a high dispersal ability and can presumably therefore recolonise turloughs on reflooding. Turloughs sampled in Co. Galway appear to have a slightly different assemblage of corixid species than nearby lakes, with Rahasane (being large) containing an intermediate assemblage (Tobin and McCarthy, 2004).

4.3.

Species adaptation to the turlough environment

Invertebrates can colonise the aquatic phase of turloughs in a number of ways: from internal and external sources and through development of in situ resting stages (Reynolds, 1982, 1996a, 2000). For turloughs that dry fully, colonisation from external sources requires that the appropriate life stages (e.g. aerial adult insects) coincide with the onset of flooding. When no colonists are available during the flood, diversity is low and populations sparse (Reynolds, 1996a). Turloughs may have small areas of still or flowing permanent waters (Reynolds, 1982), which allow species of dragonfly, mayfly, caddis-fly, etc. to lay eggs in summer. Turloughs which do not recede fully are expected to contain greater invertebrate abundance and to demonstrate greater community complexity (Duigan, 1988; Reynolds, 1997). When the floodwaters rise, individuals move from refugia within the basin (which might be just a sinkhole) to occupy the littoral zone. Aquatic invertebrates also appear able to survive dry periods in damp mud; live snails, cladocerans, Gammarus, water bugs, dragonfly and fly larvae and beetles have been observed there during summer. Aquatic organisms can also be transferred directly from permanent water bodies to turloughs via overland flow during high floods. Eighteen crustaceans dominate a list of 24 aquatic invertebrates considered to be characteristic of, though not confined to, turloughs (Reynolds, 2003). Some species of crustaceans, such as cladocerans and copepods, are particularly well adapted to colonising turloughs (Reynolds et al., 2004). They use resting stages, which in permanent water bodies allow the species to withstand food shortages or other stresses, to survive dry periods. Turlough cladocerans increase rapidly by parthenogenesis to saturate available niches and develop

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gametogenetic populations in late spring that produce ephippia to survive the dry summer and hatch upon autumn flooding (Duigan and Frey, 1987b; Duigan, 1988; Duigan and Kovach, 1991). The large chydorid Eurycercus glacialis is a winter–spring form, which may help it to evade competition or predation (Reynolds et al., 2004). Of the Odonata the turlough spreadwing, Lestes dryas is the rarest and perhaps the most specialist Irish damselfly. Like the ruddy darter dragonfly, Sympetrum sanguineum, it has a life cycle appropriate to the seasonality of the water levels of turloughs (Nelson and Thompson, 2004). At least 12 other Odonata species have been recorded on turloughs (Bond, 1997; Nelson and Thompson, 2004).

4.4.

Conservation – rare species

Turloughs can support species and assemblages of restricted distribution and high conservation value, containing diverse and characteristic communities of crustaceans (Reynolds, 1996a), aquatic beetles (Bilton, 1988; Bilton and Lott, 1991; Foster et al., 1992), and some Odonata (Nelson and Thompson, 2004). Bilton (1988) identified a characteristic water’s-edge mossdwelling aquatic/semi-aquatic beetle assemblage, found in the margins of undisturbed turloughs and unknown from Great Britain. It included three species new to Ireland, Graptodytes bilineatus, Dryops similaris (Red Data Book category three species (Shirt, 1987) and Berosus signaticollis. Other characteristic turlough beetle species include Hygrotus quinquelineatus, Agabus labiatus, Agabus nebulosus, Hygrotus impressopunctatus and Helophorus spp. (Bilton, 1988). H. quinquelineatus, A. labiatus and B. signaticollis are ‘nationally notable B’ species in Great Britain and this important assemblage is considered of high conservation value (Foster and Eyre, 1992). Bradish et al. (2002) have confirmed the occurrence of G. bilineatus and B. signaticollis in turloughs, but the assemblage has also been recorded in other Irish sites with fluctuating water levels (G.N. Foster, pers. commun.). It is likely that it is vulnerable to disturbance by anthropogenic factors (Bilton, 1988) and extremely sensitive to alterations in flooding. Rare peri-glacial relict species of Crustacea have also been recorded in turloughs, including the cladoceran Eurycercus glacialis (Duigan and Frey, 1987a,b; Reynolds, 1997, 2000; Reynolds and Marnell, 1999; Reynolds et al., 2004) and Diaptomus castor (Ali et al., 1987; Reynolds, 1997). These species are largely restricted to temporary habitats, making the presence of turloughs integral to their conservation. Typically, these characteristic turlough organisms are found with a mix of common species and species which have restricted ranges (e.g. Daphnia longispina, Eurycercus lamellatus (Reynolds, 1997; Reynolds, 2000)). The only Irish freshwater anostracan, Tanymastix stagnalis, was discovered in Rahasane turlough in 1974 (Young, 1975, 1976). This was the first record for the crustacean in Britain or Ireland, and it was presumed to have been introduced as eggs by birds or humans (Young, 1976; Grainger, 1991). The species was subsequently recorded in six temporary field ponds (Grainger, 1976, 1991). Grainger (1976) considered small, temporary ponds and not turloughs to be the natural habitat for the species. Diaptomus cyaneus is another crusta-

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cean species to be first recorded in Britain or Ireland from Irish turloughs and temporary field ponds (Ali et al., 1987; Grainger and Holmes, 1989). An absence of fish is thought to be responsible for the success of some rare invertebrate species in turloughs. Eurycercus glacialis has been recorded at 18 turlough sites, 17 of which lacked vertebrate predators (Duigan and Frey, 1987a,b; Duigan, 1988; Reynolds and Marnell, 1999; Reynolds et al., 2004). It seems the large size of both Eurycercus glacialis and Tanymastix stagnalis render them particularly vulnerable to visual predators, confining them to temporary water bodies (Duigan and Frey, 1987a,b). The turlough spreadwing, Lestes dryas may also benefit from the absence of larger predatory invertebrates and fish (Nelson and Thompson, 2004). However, over-intensive grazing of aquatic vegetation may adversely affect egg-laying and remove shelter for adult dragonflies (Nelson and Thompson, 2004).

5.

Invertebrates – terrestrial fauna

5.1.

Introduction

The terrestrial invertebrate fauna of turloughs includes several rare species and communities. Yet to date only some faunal groups have received much attention. The beetles and butterflies have been described to some extent, but the remaining terrestrial orders are poorly documented.

5.2.

Species occurring in turloughs

The first turlough records of interest for terrestrial invertebrates include three ground beetle species new to Ireland from the Coole–Garryland complex: Platynus livens, Badister meridionalis, and Badister peltatus (Speight, 1976; Speight, 1977). All three are considered rare and threatened due to the loss of wetlands throughout Great Britain (Hyman and Parsons, 1994). During surveys of wetland sites in 1987–1989 a total of nine turloughs were visited, adding five new species of beetles to the Irish list for turloughs, including Philonthus furcifer, a rare rove beetle in Ireland which does not occur in Great Britain (Lott and Foster, 1990) and recording a number of other rare species, most of which are strongly associated with wetlands (Lott and Bilton, 1991; Lott, 2003). More recently, several more rare beetles have been recorded for turloughs, confirming the importance of this habitat for these invertebrates (Anderson, 1997; Owen, 1997; Good and Butler, 2001; Nı´ Bhriain et al., 2002; Moran et al., 2003; Regan and Anderson, 2004; Regan, 2005a; Regan and Moran, 2005). Among the flies (Diptera) recorded from the Coole–Garryland turlough complex, Clusiodes caledonica and Zabrachia minutissima (Zetterstedt) are new to Ireland (Speight and Cogan, 1979; Speight et al., 1979), though these are two saproxylic species and are probably associated solely with the adjacent woodland and not the turloughs per se. In one turlough in Co. Mayo, 67 morphospecies from 34 dipteran families were found (Ryder et al., 2005). Many species of these families have larvae that are strongly associated with wetland habitats and feed on decaying vegetation (Ryder et al., 2005). The Sciomyzidae were

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identified to species level, with Pherbellia nana being the most abundant and Colobaea distincta recorded new to Ireland (Ryder et al., 2003; Ryder et al., 2005). Two hundred and forty species of butterflies and moths (Lepidoptera) were recorded from turloughs and adjacent areas as part of the Gort Flooding Study (Bond, 1997), 8% of which are known only from a few Irish localities (Bond, 1997). Winter-flooding areas (such as turloughs) that frequently have a high cover of silverweed Potentilla anserina, or are otherwise species-poor for plants were not found to harbour many lepidopteran species (Bond, 1997). However, turloughs which remained damp in summer hosted several species, notably the scarce Paraponyx stratiotata. Two other Lepidoptera Bactra furfurana and Deltote uncula occurred on many wetlands, but were noted as abundant in or around turloughs (Bond, 1997).

5.3. Environmental factors affecting terrestrial invertebrates Once the importance of turloughs for rare terrestrial Coleoptera had been established, interest developed and detailed studies were undertaken. As part of the Gort Flooding Study the ground and rove beetle communities of turloughs in the Gort area were surveyed (Good, 1997; Good and Butler, 2001). A total of 115 species of ground and rove beetles were recorded and distinct differences between the faunas on different soils detected (Good and Butler, 2001). The conservation importance of turloughs for these communities was confirmed, with 18% of the species recorded considered indicators of well-developed habitat: defined as an ecosystem that ‘is sufficiently undisturbed by human activity to retain many local or rare characteristic species’ (Good and Butler, 2001). The environmental factors influencing the communities are complex, involving, inter alia, rate and timing of draw-down and recharge of the turlough waters, soil type and surrounding over-wintering habitat (Good and Butler, 2001). Comparing roughly the same flood zone in two turloughs in SE Galway, Nı´ Bhriain et al. (2002) found two species, Blethisa multipunctata and Chlaenius nigricornis, scarce in Britain and possibly declining in Ireland, only in the turlough with peaty soils and less intensive land-use. Pelophila borealis, very rare in Britain, occurred at both turloughs on bare ground in more heavily grazed areas. Variation in carabid species composition between fields in the same turlough was also noted. Results highlight inter alia the importance of land management practices for species occurrence and the potential of terrestrial invertebrates as bioindicators (Nı´ Bhriain et al., 2002). A significant positive relationship was found between Dipteran morphospecies richness and mean vegetation height in a vegetation zone with different stocking densities on Skealoghan turlough in Co. Mayo (Ryder et al., 2005). The authors suggest that some element of rotational grazing be employed in turloughs to ensure high dipteran diversity levels. Bond (1997) found the main factors resulting in high Lepidoptera abundance and diversity in the Gort lowlands to be floral diversity (food plants), a low (to absent) grazing re´gime and the presence of adjoining areas of undisturbed habitats such as scrub, deciduous woodland and limestone pavement.

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Turlough areas that remain wet in summer also revealed a relatively richer fauna (Bond, 1997).

5.4.

Conservation

A number of beetle species found in turloughs are listed in the review of the scarce and threatened Coleoptera of Great Britain (Hyman and Parsons, 1992, 1994). Agonum lugens and Philonthus furcifer are not found in Britain, but occur in Ireland mainly on turloughs (Anderson et al., 2000; Owen, 1997). Five of the six key invertebrates associated with turloughs listed in the Interpretation Manual of European Union Habitats (Roma˜o, 1999) are ground beetles. Turloughs are, therefore, a refuge for many rare species of terrestrial invertebrates associated with wetlands, many of which are characteristic of intermittently flooded areas (Good and Butler, 2001). The terrestrial invertebrate fauna of turloughs appear to be influenced by the flooding regime (Bond, 1997; Good and Butler, 2001; Lott, 2003). There is also some evidence to suggest that land-use on turloughs can be an important factor influencing terrestrial invertebrate diversity (Bond, 1997; Nı´ Bhriain et al., 2002; Ryder et al., 2005). Therefore, drainage and change in management practices would be detrimental to these communities.

6.

Vertebrates

6.1.

Fish and amphibians

Fish are rarely present in turloughs as they are long-lived, fully aquatic organisms; where they do occur, populations are likely to be small and limited by summer water and prey availability (Reynolds, 1982). Three-spined sticklebacks Gasterosteus aculeatus and occasionally small pike Esox lucius have been recorded (Reynolds, 1982, 1997). Eels Anguilla anguilla are abundant in the Dunkellin catchment, which includes Rahasane turlough (Callaghan and McCarthy, 1993, 1996). Eels used to be common in some turlough sites, but with a national decline in numbers, this seems no longer to be the case (Goodwillie, 2001). Amphibians, however, are widespread in turloughs. Many use ephemeral water bodies for spawning and hatching in the spring, when turloughs are still wet. Frogs Rana temporaria are widespread in Ireland, but demonstrate a preference for ponds surrounded by good terrestrial cover (Marnell, 1998b, 1999). Frog-spawn and tadpoles were noted in several turloughs (Reynolds, 1982, 1997) and the frogs in and adjacent to Carran turlough were the subject of a detailed demographic study that showed turloughs are just some of many semi-ephemeral sites frogs use remarkably consistently for spawning (Gibbons, 1983; Gibbons and McCarthy, 1984, 1986). Smooth Newts Triturus vulgaris (L.) are less widespread than frogs in Ireland and appear to be generally associated with Carboniferous limestones (Marnell, 1994, 1998a, 1999). Of the newt breeding sites recorded in a national survey of the species, 5.6% were turloughs (Marnell, 1998a) Newt tadpoles metamorphose during July and August, at which stage they can leave the water, therefore newts will only breed in turloughs that retain some water throughout the summer

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and early autumn. An absence of predators is an advantage and newts prefer fish-free ponds for breeding (Marnell, 1996a,b). Newts may themselves be the top predators in turloughs during the early part of the year (F. Marnell, pers. commun.), but large invertebrates, such as beetle larvae (family Dytiscidae) and greater water-boatmen (Notonecta spp.), can prey on amphibians in turloughs. All amphibians are protected in Ireland under the Wildlife Acts 1976, 2000.

6.2.

Mammals

Though Ireland holds one of the more important remaining otter Lutra lutra populations in western Europe (Chanin, 1985; Springer, 2002), a national otter survey only recorded otters in a few turlough sites (Chapman and Chapman, 1982) and in 2000 these did not show any evidence of otters (Springer, 2002). The dominant fish prey for otters appears to be eel (O’Sullivan, 1996) and otters may use turloughs only at specific times of the year, for example when the waters are receding and eels are an easy prey (S. Springer, pers. commun.), or in spring, when frogs are an important seasonal prey at spawning time (J. Reynolds, pers. commun.). It may be that only turloughs close to the sea are important for otters, as they would be a freshwater source for washing salt out of the pelt (S. Springer, pers. commun.). Hares Lepus timidus hibernicus are often abundant on turloughs, which provide suitable habitat for them, with short swards for feeding as well as longer vegetation for cover (R. Jeffrey, pers. commun.). The non-native brown rat Rattus norvegicus is common throughout Ireland and its watercourses. It has been recorded as a predator of eggs on a turlough when bird nests are exposed by receding waters (Humphreys, 1978). Relatively little work has been published on bat association with freshwater habitats (Hutchinson et al., 1998). Although turloughs may provide open water for only part of the bat feeding season, they may be important in mild winters, when bats temporarily emerge from hibernation to feed (K. McAney, pers. commun.). Leisler’s bats Nyctalus leisleri are generally active over open habitats such as rivers, lakes and grasslands and have been recorded foraging over turloughs (O’Mahony, 1998). Turloughs could also be a foraging habitat for other species such as Daubenton’s Myotis daubentoni, whiskered Myotis mystacinus and natterer’s bats Myotis nattereri (O’Mahony, 1998), though the last two favour woodland and therefore may only be present where there is adjacent woodland or scrub such as at Coole–Garryland. The lesser horseshoe bat Rhinolophus hipposideros is thought to be most vulnerable to karst-related flooding events as its winter roosts are frequently in underground cave systems (Galway Bat Group, 1997). Excessive turlough winter flooding in the Gort area in 1994/1995 is likely to have reduced the population at Coole Cave (Galway Bat Group, 1997).

6.2.1.

Conservation

All bat species and otters are protected under the Irish Wildlife Act (1976 and 2000). The otter and the lesser horseshoe bat are also priority species under Annex II of the Habitats Directive while all bat species are covered by Annex IV of the directive. Otters and seven species of Irish bats are listed

1 3 3 ( 2 0 0 6 ) 2 6 5 –2 9 0

in the Irish Red Data Book for vertebrates (Whilde, 1993) and as protected species in the Bern Convention on the Conservation of European Wildlife and Natural Habitats. Turloughs may not be essential to their survival, but they are often part of a complex of karst, wetland and scrub woodland elements that combine to ensure their survival.

6.3.

Birds

6.3.1.

Introduction

Many turloughs have long been known to ornithologists as good sites for over-wintering wildfowl (Ruttledge, 1966, 1989; Cabot, 1999). The seasonality of the wildfowl coincides with the lake phase of turloughs, but unlike lakes, turloughs are particularly good feeding grounds for wildfowl as they are often shallow with wide, gently sloping margins and the entire basin floor is covered with vegetation (Madden and Heery, 1997). When the basin is partly full, there is plenty of food for dabblers such as wigeon Anas penelope and teal A. crecca and when it is full, dabblers still find shallow margins and the diving duck such as tufted duck Aythya fuligula and pochard A. ferina can reach the deeper areas.

6.3.2.

Frequent visitors to turloughs

The preference for turloughs of several species (mostly dabblers and grazers), such as whooper swan Cygnus cygnus, wigeon, teal, Greenland white-fronted goose Anser albifrons flavirostris Dalgety and Scott and pintail Anas acuta has been noted by Ruttledge (1989). Rahasane turlough is the most famous for wildfowl and was long rated of international importance for its numbers of 11 species of wildfowl and wader (Ruttledge, 1966; Hutchinson, 1979; Scott, 1980; Anon., 1981; Whilde, 1986; Grimmett and Jones, 1989; Sheppard, 1993). However, recent counts at Rahasane have shown a significant drop in bird numbers and only black-tailed godwit Limosa limosa are now at numbers of international importance and whooper swans, Greenland white-fronted geese, wigeon, shoveler Anas clypeata, pintail, lapwing Vanellus vanellus and golden plover Pluvialis apricaria are of national importance (Crowe, 2005). It is also the only major inland site in both Britain and Ireland for wintering dunlin Calidris alpina (Sheppard, 1993), though they have also declined (Crowe, 2005).

6.3.3.

Site ecology and evaluation

Aside from studies on the Rahasane area (Buckley, 1993; Buckley and McCarthy, 1987), little bird research has concentrated solely on the turlough ecosystem. However, as part of the South Galway Flooding Study, 28 sites were surveyed for breeding waterfowl (Madden and Heery, 1996), of which 19 were turloughs, two of which were studied in detail (Madden and Heery, 1999). The sites were classified according to their flooding levels and their response times to a wet period. Where flooding is in the middle ranges (low to moderate or moderate to high) it is most conducive to stable bird populations, both feeding and roosting. However with regular, sometimes fast changes in water level at other sites, birds move frequently between the sub-sites of the complexes. Thus, should a turlough fill to above-moderate levels, dabblers might move onto the drier turloughs. The importance of such

B I O L O G I C A L C O N S E RVAT I O N

‘dry’ turloughs as Ballylee or Cahermore is perhaps critical at times of such extreme flooding elsewhere and highlights the need to preserve a diversity of adjacent sites with respect to flooding levels (Madden and Heery, 1997). At Rahasane, disturbance of birds is notable (Buckley, 1993), but it was not considered a factor in the South Galway sites (Madden and Heery, 1996). Only turloughs that do not dry out completely are important for breeding, provided they are not uniform well-grazed swards (Madden and Heery, 1996). A relatively high number of breeding birds was noted at the Williamstown turlough complex (N Galway), where an abundance of sedge communities and the presence of peat, indicate more permanently wet conditions (Environment Consultancy, 1997). Birds which commonly breed in dry or damp grassland such as meadow pipits Anthus pratensis, lapwing and snipe Gallinago gallinago have been recorded on turloughs (Buckley and McCarthy, 1987; Buckley, 1993). The two rare birds that once nested in a few turloughs, the black-necked grebe Podiceps nigricollis Brehm and red-necked phalarope Phalaropus lobatus appear to have ceased due to drainage (Humphreys, 1978; Ruttledge, 1978).

6.3.4.

Conservation

Systematic recording of sites for bird conservation rating has only taken place in recent decades. Rahasane was the first turlough to be listed (Ruttledge, 1966; Hutchinson, 1979; Scott, 1980; Whilde, 1986), but several more appear in Ruttledge (1989) and Sheppard (1993). They are often listed as complexes, since the birds commute between adjacent turloughs that can act as near-by refuges, should the birds be disturbed (Buckley, 1993; Madden and Heery, 1997). The turloughs in the Gort region form two main groups, the Coole and the L. Coy complexes. These sites, along with the adjacent Ballinduff turlough, have each hosted numbers of international and national importance for whooper and Bewick’s swans respectively (Madden and Heery, 1997). Both swans are listed in Annex I of the EU Birds Directive (EEC, 1979). The Coole complex is a Ramsar site and it, along with Rahasane and three sites in Roscommon, are the only turloughs listed as Special Protection Areas (SPAs) under the Birds Directive (Working Group on Groundwater, 2004) (see Appendix 1). Sheppard (1993) lists some 25 turloughs or clusters such as the 17-turlough Castleplunket complex in Roscommon and the ‘southern group’ which includes the Coole–Garryland complex, all of which are at least of regional/local importance for winter birds. In a recent wetland bird survey (I-WeBS), 45– 50 of the sites are turloughs, of which over 20 are in two complexes, ranked in the top 35 sites of national importance for wildfowl and wader counts, while Rahasane and Coole–Garryland rank of international importance (Crowe, 2005). The fact that turloughs rank alongside other wetland sites that are far larger (e.g. Lough Corrib is 18,240 ha) is because they can support far higher densities, because the vegetation fully covers the turlough basin, yet no study has specifically compared the relative importance of turloughs with other wetland sites. Over 40% of the Icelandic breeding population of whooper swans over-winter in the Republic of Ireland (Cranswick et al., 2002) and of the 27 sites in Britain and Ireland rated as of

1 3 3 ( 2 0 0 6 ) 2 6 5 –2 9 0

277

international importance for whooper swans during 1995– 2000, six are turloughs or include turlough complexes (Robinson et al., 2004). The Coole and L. Coy complexes can together host at least 20% of the over-wintering population of whooper swans in Ireland (this was over 1000 birds when the threshold number of whooper swans for a site to be rated of international importance was 160) (Sheppard, 1993; Madden and Heery, 1997). Drainage, such as happened with the Clare River catchment, has resulted in the loss of turloughs that were important wintering bird sites (Stronach, 1973; D’Arcy, 1983) and is clearly undesirable for any aspect of turlough conservation (O’Gorman, 1985; Buckley and McCarthy, 1987). However, extremely high levels of flooding have also been noted as being detrimental to numbers of bird visitors (Rutledge and Ogilvie, 1979). Wildfowl numbers at Rahasane declined in the late 1970s and 1980s possibly due to excessive flooding, as a result of ‘un-regulated’ drainage upstream, causing the turlough downstream to flood to higher levels (Rutledge and Ogilvie, 1979; Whilde, 1986; Buckley and McCarthy, 1987). High water levels at Rahasane reduce food availability for whooper swans, which are known to favour turloughs for feeding and roosting (Robinson et al., 2004). Since then the main channel in Rahasane was deepened, reducing excessive floods, but also causing the turlough to drain very fast. Winter wetland bird counts there have further declined, by 63% between 1984–1986 and 1996–2000, resulting in the site being dropped from the top 20 wildfowl and wader sites in the island (Crowe, 2005). Water fluctuation clearly affects feeding wildfowl and, whereas complexes of turloughs that have different flood levels supply the variety needed for all species, it is important to monitor levels such that an excessively high level does not prevail any more than an excessively low one (Madden and Heery, 1997). As teal and to some extent tufted duck and pochard feed on seeds of sedges and pondweeds, the maintenance of these plant communities will be essential for supporting them (Madden and Heery, 1997). Sedge communities occur in turloughs that are permanently moist and so depend in turn on the turlough hydrology (Regan, 2005b).

7.

Turlough management and conservation

7.1.

Background

Although there is some indication that birds may require relatively subtle site-specific flooding regimes and vegetation, it is more difficult to determine the particular environmental factors influencing, for example, terrestrial invertebrates. It is now thought that specific turlough types exist with respect to invertebrate fauna, but the factors that determine the differences such as soil type, hydroperiod and land-use not only of the turlough, but of the surrounding area, are as yet ill-defined. It is important, therefore, to define and conserve as diverse a number of turloughs as possible, in relation to both habitat and management practices. Whereas drainage up until now has been the most significant threat to turloughs, land-use clearly influences their vegetation and soils and hence their terrestrial fauna. Specific site studies emphasise

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not only the importance of managing diversity within as well as between sites, but also the basic necessity of retaining and recognising the farming community as crucial to the conservation of the turlough habitat (Nı´ Bhriain et al., 2002, 2003). Though former cultivation ridges are still visible in some turloughs and are evidence of past crop planting, turloughs are now thought to be entirely used for grazing. It would seem that there are now three principal threats to turlough conservation: drainage, nutrient input and management (grazing) changes.

7.2.

Drainage

Detailed research on turlough ecology and hydrology has often been in relation to proposed drainage plans for a catchment (O’Gorman, 1983, 1984, 1985; Lockhart, 1984, 1985; Goodwillie et al., 1992; Environment Consultancy, 1997; Southern Water Global, 1998) or the effects of drainage on a catchment (D’Arcy, 1983). Roughly one third of turloughs over 10 hectares have been irreversibly damaged by drainage (Coxon, 1986). The impact of land drainage on groundwater resources is particularly acute in karst areas because of the unique characteristics of karstic aquifers. Arterial drainage, or drainage of river systems to dry out land within the catchment, of karst lowlands in Ireland since the mid-19th century has resulted in losses of recharge, lowering of water tables, drying up of turloughs, alteration of underground flow routes, and periodic groundwater contamination (Drew and Coxon, 1988). Though large-scale drainage has ceased, it resulted in the loss of at least 50% of flooded turlough area (Coxon, 1986; Goodwillie, 2001). Any changes in the flooding regime will have dramatic impacts, especially on the rarer biological communities of turloughs. Several turloughs dried out during the drainage of the Clare River catchment and farmers found that although this resulted in more land being available for grazing for a longer season, fertiliser was now needed and the shallower soils were susceptible to poaching (D’Arcy, 1983). Elsewhere, drainage has been seen as of benefit to the land-owners, though completely altering the ecology and species composition of the turlough basin (Tenthorey, 1994). Since karst hydrology is particularly complex, it is not likely that restoration would be feasible, particularly of the larger (and therefore more unusual) ones of the R. Clare catchment.

7.3.

Nutrient/trophic status

The evaluation of turloughs with respect to nutrient status is complex. They have been ranked according to the trophic status of their vegetation and Ellenberg Fertility Scores for the dominant plant species (Goodwillie, 1992; Working Group on Groundwater, 2004). The vegetation has been classified into mesotrophic, disturbance-indicator, grassy plant communities, or oligotrophic sedge-dominated communities (IvimeyCook and Proctor, 1966; O’Connell et al., 1984). However, the extent to which these are influenced by anthropogenic factors such as water pollution and fertilising is not yet clear. Only on Rahasane have plant nutrient analyses been carried out, showing the vegetation there to have nitrogen levels comparable to those of fertilised grassland (soil and plant P were rela-

1 3 3 ( 2 0 0 6 ) 2 6 5 –2 9 0

tively low) (Sheehy Skeffington, 1985). Elsewhere, preliminary turlough water sampling in SE Galway showed that nitrate levels were highest at those turloughs that were most strongly grazed, indicating manure effects or possible fertiliser addition (Southern Water Global, 1998). Water total phosphorus and to a lesser extent total nitrogen, were higher for turloughs in SE Galway that had more grass-dominated vegetation (Regan, 2005b). Some turloughs have a relatively long history of low-level chemical fertilising (Nı´ Bhriain et al., 2003), while others do not (Aughney and Gormally, 1999). Catchment water is thought to be the main factor determining nutrient input into turlough systems (Southern Water Global, 1998). Limited water sampling in a small ground-water fed turlough in north Clare over winter/early spring 1998/1999, revealed waters to be mesotrophic (Nı´ Bhriain, 1999). Changes in the vegetation over recent years were known to have occurred and ground water pollution from fertiliser and possibly slurry was suggested as the main cause of these changes (Nı´ Bhriain, 1999). Turlough flooding may, however, also supply essential nutrients for plant growth (MacGowran, 1985; Sheehy Skeffington, 1985). Risk assessment for turlough nutrient pollution is required under the Water Framework Directive (Working Group on Groundwater, 2004).

7.4.

Grazing

Grazing is a major ecological factor in turloughs, either as commonage or in privately owned fields. They are grazed, sometimes heavily by cattle, sheep and even horses and domestic geese (MacGowran, 1985; Goodwillie, 1992, 2003; Feehan, 1998; Aughney and Gormally, 1999; Nı´ Bhriain et al., 2003). Time and density of stocking can vary even between adjacent fields that may in turn support different plant and invertebrate communities (Nı´ Bhriain et al., 2002, 2003). Thus management prescriptions for turloughs may need to be complex and flexible to maintain existing diversity of flora and fauna. Apart from limiting shrub growth, the sward is directly affected by grazing through the removal of the more palatable species and by physical damage and manuring. When stock is put out to pasture before the vegetation has regenerated following water subsidence, the soil surface is broken up, encouraging the proliferation of ‘weedy’ species (Goodwillie, 1992). If not widespread, this is not detrimental and is an intrinsic part of turlough biodiversity. The stocking density for turloughs recommended by the National Parks and Wildlife Service is 1.5 livestock units per hectare (LU ha1). Research on past and present agricultural practices on turloughs so far suggests an upper limit of 1 LU ha1 (Aughney and Gormally, 1999; Nı´ Bhriain et al., 2002; Nı´ Bhriain et al., 2003) referring mostly to cattle. However, large variation in times and rates of stocking occurs between sites, even on one turlough (Nı´ Bhriain et al., 2003). No research is published to date on behaviour of animal type and breed, or on their effects on sward height and composition in turloughs. It has been shown above that different sward heights favour different plant and invertebrate species and thus management diversity within a turlough is important for its biodiversity. In fact, it is likely that the major threat to turlough biodiversity, given that turloughs are considered marginal land, is the abandonment of land. An

B I O L O G I C A L C O N S E RVAT I O N

absence of grazing, if extensive over a whole turlough basin is likely to be detrimental to both vegetation and invertebrate diversity. Management plans for turloughs must therefore be cognisant of national trends in farming and include incentives to retain farming systems on the turloughs, as well as incorprating the experience of farmers managing these habitats.

8.

Conclusion

Turlough importance as unique hydrological features has been recognised (Coxon, 1987a,b; Drew, 1990; Reynolds, 1996a,b; Goodwillie and Reynolds, 2003). However, though it is clear that turloughs support a number of unusual plant and invertebrate communities, knowledge on their biodiversity is far from complete. Whereas there have been several attempts to classify turloughs in relation to their flooding, geomorphology and trophic status (e.g. Coxon, 1987a,b; Goodwillie, 1992; Goodwillie and Reynolds, 2003), it is also apparent that each turlough has its own unique hydrological features that in turn have a bearing on substrate, vegetation type and/or invertebrate communities (Moran, 2005; Regan, 2005b). Turloughs are wetlands, which are perhaps the most threatened and fragile habitats in Europe, Many of the rarer turlough species are wetland species. Some of them are adapted either in life cycle or behaviour, to the unique seasonal flooding. Many of the unusual terrestrial species may be there because the wetter turloughs are refuges of lowintensity agriculture in a more intensively used landscape. However, in investigating the factors important to the distribution of these species, it cannot be assumed that turloughs supporting rare plant species are always those that support rare invertebrates or indeed that the invertebrate or bird fauna respond to the same factors. Many insects require tall vegetation for cover, birds require shallow winter flooding and some insects and birds have specific food dependence on particular plant assemblages. For aquatic invertebrates, factors such as the absence of fish as well as the oligotrophic, calcareous nature and mossy-fen plant communities of some turloughs are probably important. Many turloughs host large numbers of over-wintering birds, several species of which reach levels of international or national importance (Sheppard, 1993; Crowe, 2005). However, there has been no specific research on the importance of turloughs for such birds relative to other wetland sites. They may have to be viewed as complexes with a diversity of complementary flooding patterns (Madden and Heery, 1997) or as part of larger wetland systems, acting for example as freshwater refuges for birds feeding in adjacent coastal complexes, such as Inner Galway Bay (designated as a SPA under the Birds Directive). The shallow depth of turloughs and their extensive underwater swards of vegetation are of particular importance to birds such as whooper and Bewick’s swans as well as many dabbling and diving duck. Not all feed on grasses however, and the importance of seed-producing vegetation such as sedges has also to be recognised. As wildfowl move between sites, it is not possible to select a single representative site for conservation. This mobility may also apply to other biota and the inherent diversity of

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279

turloughs requires that a range of turloughs, even adjacent, be conserved. It is apparent that the role of the farming community is essential to the maintenance of turlough biodiversity. Sward height, time and length of grazing, stocking density, fertiliser use and even the land-use of surrounding habitat are important factors, as are the variety of management regimes that exist, even within one turlough system. These are not just intrinsic to maintaining plant biodiversity, they also define invertebrate communities and affect bird populations. The Habitats Directive requires the drawing up of management plans for each turlough designated a Special Area of Conservation (SAC). A total of 43 SACs have been designated for turloughs but as several SACs contain more than one turlough, the total number of turloughs under designation is 71 and of these, eight are designated as SPAs and one is a Nature Reserve (Stapleton et al., 2000; Working Group on Groundwater, 2004). It is important that further information concerning management on turloughs is acquired and that, in imposing any regimes, flexibility be applied, such that farmers are encouraged to maintain their use of turloughs (Nı´ Bhriain et al., 2003). The Water Framework Directive also requires risk assessments for all water bodies, yet little is known about groundwater catchments and therefore about potential sources of pollution. This is a much more difficult task than predicting changes in land-use or potential drainage plans (Working Group on Groundwater, 2004). As potentially threatened wetlands of European importance, turloughs require a full inventory of their biodiversity and the factors affecting it. Priority research areas are the documentation of the temporal and spatial variability in invertebrate communities, an investigation of the abiotic and biotic factors affecting plant and animal communities on turloughs, a documentation of current and past land-use in turloughs and the identification of the impact of external factors such as drainage and nutrient enrichment on these communities. Turloughs are hydrogeological phenomena that support complex biological systems that in turn are integrally maintained by a diversity of farming systems. While drainage continues to be a major threat to turloughs, designation of the prominent ones has given them protection. It is possibly the current economic threat to the survival of relatively small farms in the more extensively farmed regions of Ireland where turloughs occur, that may now be the single greatest threat to the conservation of turlough biological diversity, through the cessation of grazing on turloughs.

Acknowledgements A very special thanks to Julian Reynolds who gave invaluable comments on the final draft. Very many thanks also to Caoimhe Hickey, Geological Survey of Ireland, who so willingly compiled and perfected the map. Thanks are also due to the following for supplying information: Roy Anderson, Tom Bolger, Ken Bond, Philip Buckley, Paul Corbett, Don Cotton, Olivia Crowe, Donal Daly, Gordon D’Arcy, Ian Enlander, Roger Goodwillie, Erin Gibbons, Maria Gibbons, Maggie Hall, Rebecca Jeffrey, Daniel Kelly, Neil Lockhart, Kate McAney, T. Kieran McCarthy, Brian MacGowran, Colin McLeod, Brian

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Madden, Ferdia Marnell, Eleanor Mayes, Oscar Merne, Myles Nolan, Jim O’Connor, Ciara O’Mahony, Cilian Roden, Jim Ryan, Martin Speight, Sabine Springer, Genevie`ve Tenthorey,

1 3 3 ( 2 0 0 6 ) 2 6 5 –2 9 0

John Wilson, Marjolein Visser, Chris Williams, Mike Wyse Jackson. Three referees also gave useful comments for which we are grateful.

Appendix 1 List of turloughs in Fig. 1, with grid references and listed by townland name within each county. Where known, the area is given and for any that are within a SAC, the name of that SAC (complex) is listed, where possible. All turlough SPAs are in SACs and that designation is in brackets alongside the SAC name, where it applies. Data by permission of the Geological Survey of Ireland (GSI) and amended by the authors. N/A means the turlough name is not known. Townland

County

Easting

Northing

Name

1

Aughrim

Clare

135550

196770

Travaun–Skaghard Turlough Carran and Termon turloughs Ballyvelaghan Castle Lough N/A Fahee Turlough N/A Turloughmore N/A Lough Geala´in N/A N/A N/A N/A N/A Knockaunroe N/A N/A Muckinish Lough Lough Gash Ballyvaughan Turlough N/A

2

Ballyconry

Clare

128200

198560

3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32

Ballyvelaghan Carrownagoul/Leitra Creehaun Fahee South Glenquin Gorteen/Derrybeg Gortlecka 1 Gortlecka 2 Kilcorney Killeany Killeenmacoog South Knockaunroe Knockaunroe Knocknagroagh Lackareagh Glenquin Muckinish East Newmarket on Fergus Newtown Nooan Pouleenacoona Poulroe Sheshymore Turlough Turlough Turloughmore Ballyadam Carrigane Crockacapple Dunmuckrum Fardrum

Clare Clare Clare Clare Clare Clare Clare Clare Clare Clare Clare Clare Clare Clare Clare Clare Clare Clare Clare Clare Clare Clare Clare Clare Clare Cork Cork Donegal Donegal Fermanagh

127990 134520 134470 129620 131860 147480 131450 131900 122340 116550 135320 131400 130700 122800 131680 127560 139210 122300 132270 136610 137740 124740 128500 128050 134950 184000 184120 187850 186000 217700

211400 198230 198180 198580 196050 176360 194730 195000 199400 200930 201300 194180 193450 206850 196600 208750 167820 207200 185310 202210 195630 195440 205100 204600 199480 73950 74100 363150 360000 350700

33 34

Aggard Beg Ardskea More

Galway Galway

150180 144440

218050 241130

Fardrum Lough, Roosky Lough, Green Lough Aggard N/A

35 36

Ashfield Demesne Ballinastack

Galway Galway

144410 164820

197390 264820

Lough Awock Ballinastack

37 38 39 40

Ballinderreen Ballinduff Ballinillaun Ballinvoher South

Galway Galway Galway Galway

138520 146300 144450 178200

216100 208200 222600 247250

Ballinderreen Lough N/A N/A Lough Aunaculaskey

Size (ha)

SAC (SPA) name East Burren Complex

90.3

East Burren Complex

Ca. 8 37

Galway Bay Complex East Burren Complex

17.3

East Burren Complex

42.5

East Burren Complex

Moneen Mountain 21.9 9

Ballyvaughan Turlough

Poulroe Turlough Lough Aleenaun

10 10.7

East Burren Complex East Burren Complex

Turloughnagullaun N/A N/A N/A Legaltan Lough

20.4 21

East Burren Complex

Dunmuckrum Turloughs

Fardrum and Roosky Turloughs (N. Ireland designation) 24.9 83 29

Lisnageeragh Bog and Ballinastack Turlough Lough Fingall Complex Ballinduff Turlough

B I O L O G I C A L C O N S E RVAT I O N

281

1 3 3 ( 2 0 0 6 ) 2 6 5 –2 9 0

Appendix 1 – continued Townland

County

Easting

Northing

Name

41 42 43 44 45 46 47 48 49 50 51

Ballymore Ballynakillew 1 Ballynakillew 2 Ballynew Ballysheedy Balrobuck Beg 1 Balrobuck Beg 2 Balrobuck Beg 3 Balrobuck Beg 4 Baunmore Beagh

Galway Galway Galway Galway Galway Galway Galway Galway Galway Galway Galway

139960 125670 125570 131900 142960 132230 132620 133150 133400 137080 156680

213380 245970 246300 241570 201480 241570 241730 241750 241500 237500 268300

N/A N/A Balrobuck Beg N/A N/A Polleagh Lough

52 53

Beagh Beagh

Galway Galway

156800 156390

268150 269160

Curragh Lough

54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78

Beagh More Beaghroe Blindwell Blindwell Boyounagh Bredagh Caherakenny Caherbroder Cahercrin Cahercrin Caherkelly Cahermore Caherpeak West Caherpeak West Cappadarock Carheenshowagh Carraghadoo Carraghdoo 1 Carraghdoo 2 Carraghy Carrowholla Carrowneany Cashla Castlegar Castlequarter

Galway Galway Galway Galway Galway Galway Galway Galway Galway Galway Galway Galway Galway Galway Galway Galway Galway Galway Galway Galway Galway Galway Galway Galway Galway

134060 145750 134380 134640 159930 131430 134570 144200 148950 148700 146290 141710 141480 143210 152950 141500 142130 142740 143000 138150 167430 159680 143220 143640 142620

251890 255650 259150 259760 265020 244420 250850 198270 224400 223370 210290 207660 216150 215340 253450 244720 215080 214880 214950 237650 233750 270540 230050 218340 202680

78a 79 80 81 82 83 84 85

Castletaylor Castleweelder Clooncon West Cloondoyle Beg Cockstown Coldwater Common Coole Demesne

Galway Galway Galway Galway Galway Galway Galway Galway

145450 144990 163050 164300 148520 145830 143100 143110

215450 215750 263580 261200 210350 222450 240900 204260

Caranavoodaun N/A N/A N/A

86 87 88 89 90 91

Corbally South 1 Corbally South 2 Corbally South 3 Corbally South 4 Cornananta More Corrandrum

Galway Galway Galway Galway Galway Galway

141850 142250 142320 142700 176470 141270

239850 239800 239200 239300 247240 240750

Pollakilleen Meelick Pollnacloya Cloghdo

Size (ha)

SAC (SPA) name

N/A N/A N/A 3

Williamstown Turloughs Williamstown Turloughs

Beagh Bodarrig Boyounagh N/A Turlough O’Gall

21.2 50.9

Turlough Naheltia Turloughakip

Esker Tullaghnafrankagh Lough Levally Lough Turloughour Carraghadoo Turlough Derreen Moneardotia N/A N/A N/A KiItullagh N/A Newtown

46

Cahermore Turlough

48.5

Levally Lough

6 2

194 31

Lough Fingall Complex

Coole–Garryland Complex (SPA) Castletaylor complex

7 Pollnakirka N/A Coole Lough

180

Coole–Garryland Complex (SPA)

N/A (continued on next page)

282

B I O L O G I C A L C O N S E RVAT I O N

1 3 3 ( 2 0 0 6 ) 2 6 5 –2 9 0

Appendix 1 – continued Townland

County

Easting

Northing

Name

92 93 94 95 96 97 98 99 100 101 102 103 104

Crannagh Cuildooish Derrydonnell North Doughiska Drumacoo Drumminacloghaun Eenhugh Eskeromullacaun Fearagha Fearaghta Frenchpark Garraun Garryland Wood

Galway Galway Galway Galway Galway Galway Galway Galway Galway Galway Galway Galway Galway

142600 141060 143190 134980 139820 143680 142470 161250 134280 132800 141710 145600 141750

207220 214920 226270 226470 216660 201520 211850 262500 245810 246400 214920 254600 204050

105

Glenamaddy

Galway

163220

261220

Glenamaddy

106 107

Gortatleva Gortduff

Galway Galway

139650 157390

231680 269160

N/A North Gortduff Turlough

108 109 110 111 112 113 114

Kilcoona Kilcornan Kilcornan Killeenavarra Killeeneen More Killkerrin Killomoran

Galway Galway Galway Galway Galway Galway Galway

131270 144250 142390 142020 146800 163210 141550

243670 220200 219270 212900 219130 256100 206340

N/A Turlough Nacashlaun N/A

115 116 117 118 119 120 121 122

Killora Kilmacduagh Kilquain Kilternan West 1 Kilternan West 2 Kiltullagh Knockatogher Knockaunatouk

Galway Galway Galway Galway Galway Galway Galway Galway

151000 140060 145150 143400 144050 136850 158950 141140

219000 199730 217700 214510 213780 229800 226950 202320

123 124 125 126 127 128 129 130 131 132 133 134 135 136 137 138 139 140

Knockavanny Lackan 1 Lackan 2 Laghtshaughnessy Lahardaun Lehinch Liscloonmeeltoge Lisheen Lisheenageeha Lissananny Lough Curra South Lydacan Moneyteige Moyne Moyode Muckcoort Newcastle Newhall

Galway Galway Galway Galway Galway Galway Galway Galway Galway Galway Galway Galway Galway Galway Galway Galway Galway Galway

147460 147390 148000 144570 136350 163210 168880 141590 126000 137350 138330 143670 147800 124950 152500 128130 175650 147270

254060 211740 212650 199790 219120 256140 246450 199810 244300 259530 208830 207930 224700 248770 223950 242500 227200 206790

Muggaunagh N/A N/A

Size (ha)

SAC (SPA) name

12

Lough Fingall Complex

18.8 26.8 5

Lough Fingall Complex

Loughaunwee N/A Turlough Monaghan Lough Fingall N/A Garryland

25 177.5

Coole–Garryland Complex (SPA) Lough Lurgeen Bog/ Glenamaddy Turlough Williamstown Turloughs

6 N/A Kilkerrin Caherglassaun

16.6 68

Caherglassaun Turlough

Killora Turlough Turloughnacloghdoo N/A Leeragh Kiltullagh Turlough N/A Hawkhill

17

Kiltiernan Turlough

28.9 6

Coole–Garryland Complex (SPA)

Knockavanny Turlough

Ballinacourty Turlough

Ca. 20

Galway Bay Complex

N/A Polliffern lough N/A Lissananny Turlough 9 9 Millmount Turlough Rostaff Turlough N/A Turloughnacrusha N/A Carrowbaun, Newhall and Ballylee Turloughs (continued on next page)

B I O L O G I C A L C O N S E RVAT I O N

283

1 3 3 ( 2 0 0 6 ) 2 6 5 –2 9 0

Appendix 1 – continued Townland 141 Newhall

County

Easting Northing

Name

Galway

147520

206130

Ballylee River Turlough

142 Patch Galway 143 Pollynoon Galway 144 Rahasane/Carrigeen West Galway

159750 157720 147920

263130 271150 219230

N/A N/A Rahasane

145 146 147 148 149 150 151 152 153 154 155 156 157 158 159 160 161 162 163 164 165 166 167 168 169 170 171 172 173 174 175 176 177 178 179 180 181 182 183 184 185 186 187 188 189 190

Galway Galway Galway Galway Galway Galway Galway Galway Galway Galway Galway Galway Galway Galway Galway Galway Galway Galway Galway Galway Galway Galway Galway Galway Galway Kilkenny Leitrim Limerick Longford Longford Mayo Mayo Mayo Mayo Mayo Mayo Mayo Mayo Mayo Mayo Mayo Mayo Mayo Mayo Mayo Mayo

144420 141710 134560 144700 138630 133460 143320 148250 147730 150750 140920 142080 140350 141600 140330 138930 137550 150120 150370 129500 143500 149780 138020 142350 173130 231520 176000 154570 201500 202820 147690 127360 136070 122300 122730 119810 123000 123030 121750 122400 122240 120600 126810 122310 118500 126900

207280 207660 261140 218800 202210 209810 212200 207560 207710 222350 197350 197760 201650 241650 221120 213710 252130 225370 225480 244050 236900 208130 249900 240300 227230 163450 343100 152190 263160 263590 282210 262500 269810 251140 251950 261720 277450 278070 251500 279850 261630 258600 252160 252100 276660 264510

Mayo Mayo

119990 121750

253000 261290

Raheen Demesne Raheenkilkelly Rathbaun Roevehagh Roo Roo Demesne Rooaunmore Skehanagh (Kiltartan By) Srah (Kiltartan By) Templemartin Termon Termon Tirneevan Tonamace Tonroe Tooreen East Treanbaun Turloughalanger 1 Turloughalanger 2 Turloughcor Turloughmore Common Turloughnacloghdoo Turloughnaroyey Turloughrevagh Woodberry Balief Lower Formoyle Loughmore Common Commons Newpark Annagh Ardkill Ballindine Ballisnahyny Ballisnahyny Ballyargadaun Ballyglass Ballyglass Ballynalty Ballynamuddagh Ballytrasna Ballywalter Brodullagh Burnafollistran Burriscarra Caheravoostia

191 Carheens 192 Carrowkeel

Size (ha)

SAC (SPA) name Carrowbaun, Newhall and Ballylee Turloughs

267.1

Rahasane Turlough (SPA)

3 Turlough Rathbaun Dunkellin Roo West Turlough N/A Owenbristy Turlough

Turloughmartin Termon Lough Lough Mannagh N/A Tonroe Cloghballymore Lough N/A N/A N/A Turloughcor Turloughmore Common Blackrock/Peterswell Turloughnaroyey (Belclare) Turloughrevagh N/A The Loughans Loughmore Common Fortwilliam Turlough Cordara Turlough Island Lake Ardkill N/A N/A N/A Ballyglass Mountpleasant Turlough Poolbeg N/A N/A Shrule N/A N/A Caheravoostia

66.9 28

East Burren Complex

39 49 23

Termon Lough Termon Lough East Burren Complex

4

50 98.8

Peterswell Turlough

21.7

The Loughans

39 44

Fortwilliam Turlough

16

Ardkill Turlough

25.6

23.8

Kilglassan/ Cahervoostia Turlough Complex

N/A (continued on next page)

284

B I O L O G I C A L C O N S E RVAT I O N

1 3 3 ( 2 0 0 6 ) 2 6 5 –2 9 0

Appendix 1 – continued Townland

County

Easting Northing

193 194 195 196 197 198 199 200 201 202 203 204 205 206 207 208 209 210 211 212 213 214 215

Carrowkeel North Carrowmore Carrownacon Carrowreaghmony Coollisduff Coollisduff Cornacartan Cornaroya Creagh Demesne Cregduff Cregnanagh Doocastle Drumelly 1 Drumelly 2 Drummeennavaddoge Forkfield Frenchbrook North Frenchbrook South Greaghans Greenwood Illanmore 1 Illanmore 2 Kilglassan

Mayo Mayo Mayo Mayo Mayo Mayo Mayo Mayo Mayo Mayo Mayo Mayo Mayo Mayo Mayo Mayo Mayo Mayo Mayo Mayo Mayo Mayo Mayo

129810 124400 119700 122100 122760 121940 122220 120800 117750 124950 122500 158460 120500 119550 120100 107850 123500 124300 129050 144690 121850 122500 127860

268900 263780 276740 277350 259630 259870 256550 263950 265550 258000 255700 305550 256500 257200 282050 283200 259200 258800 262640 280560 257180 257200 264550

216 217 218 219 220 221 222 223 224 225 226 227 228 229 230 231 232 233 234 235 236 237 238 239 240 241 242 243 244 245

Killaturly Knockaderraghy Knockglass Knockysprickaun Larganboy East Lisduff Loughanboy Mannin Mocollagan Mocorha Moorgagagh/Cahermaculick Mountpleasant Nymphsfield Parks Parks Pollavaddy Scardaun East Skealoghan Slishmeen Thomastown Westport Demesne Ballyloghlan Ballyloughlan Ballyloughlan Donaghmore Kilmactrasha Monaltyduff Ardconra Attishane Ballinlig

Mayo Mayo Mayo Mayo Mayo Mayo Mayo Mayo Mayo Mayo Mayo Mayo Mayo Mayo Mayo Mayo Mayo Mayo Mayo Mayo Mayo Monaghan Monaghan Monaghan Monaghan Monaghan Monaghan Roscommon Roscommon Roscommon

141250 125150 119400 97450 145190 140300 124090 146250 126760 123500 126320 121400 115140 120450 121700 126320 133660 124750 122440 123380 195750 283710 283880 284130 284600 283370 286260 188020 153150 182100

298440 281320 267740 288730 282320 269400 260850 283440 252220 254200 254270 278280 256250 277900 277030 284350 269070 262900 278950 261000 286070 301100 301330 300750 307970 302620 302200 301200 273450 261380

Name Pollelamar Lough N/A N/A N/A

N/A Turloughosheenan Turloughmarlagh Turlough Faugh N/A Doocastle N/A Kiltogorra Tulroughaganny Cashel Turloughagurkall N/A Greaghans Greenwood Turlough N/A Turloughmore Kilglassaun

Killaturley Lough N/A N/A Grallaghlea Lough N/A N/A Mannin Lake Lough Nakill N/A Shrule Turlough N/A Turloughatsallain N/A N/A Pollaghard Scardaun Slishmeen Turloughaclara N/A

Size (ha)

SAC (SPA) name

30.1

Carrowkeel Turlough

38.2

Doocastle Turlough

36.5

Greaghans Turlough

49.9

Kilglassan/ Cahervoostia Turlough Complex

34

104.1

18.3 28 19.6

Srule Turlough

Skealoghan Turlough Clyard kettle Holes

N/A

N/A Attishane Turlough N/A

17.2 (continued on next page)

B I O L O G I C A L C O N S E RVAT I O N

285

1 3 3 ( 2 0 0 6 ) 2 6 5 –2 9 0

Appendix 1 – continued Townland

County

Easting Northing

Name

246 Ballinturly

Roscommon 184820

260100

N/A

247 248 249 250 251 252 253 254 255 256 257 258 259

Roscommon Roscommon Roscommon Roscommon Roscommon Roscommon Roscommon Roscommon Roscommon Roscommon Roscommon Roscommon Roscommon

185630 181700 178080 181600 192200 178420 186070 185200 189040 184420 177670 193230 183710

261150 271600 301090 276560 238800 275080 294630 301200 237750 261570 277840 238970 251620

Ballygalda Turlough N/A

260 Coolcam 261 Corbally 262 Cornageena/Pollalaher

Roscommon 157420 Roscommon 184900 Roscommon 192840

263 264 265 266 267 268 269 270 271 272 273 274 275 276 277 278 279 280 281 282 283 284 285 286 287 288 289 290 291 292 293 294 295 296

Roscommon Roscommon Roscommon Roscommon Roscommon Roscommon Roscommon Roscommon Roscommon Roscommon Roscommon Roscommon Roscommon Roscommon Roscommon Roscommon Roscommon Roscommon Roscommon Roscommon Roscommon Roscommon Roscommon Roscommon Roscommon Sligo Sligo Sligo Sligo Sligo Sligo Tipperary Tipperary Tipperary

Ballygalda Ballymacurly South Breandrum Brierfield Carrowduff and Carbally Carroweagh Carrowmore Carrownagashel Carrownure Carrowstellen Castleplunket Cloonagh Cloonlaughnan

Cuileenirwan Farranagalliagh Feacle Glennanea Inchiroe Keenagh Kilnamryall Kinclare Kinitty Knockroe Lisduff Lissergool Lissnaboll Lissonuffy Loughnaneane Mullygolan Newtown Onagh Rathnalulleagh Stonepark Tonroe Turlagh Turlagh Turlaghmore Turrock Doongelagn Leitrim South Moylough 1 Moylough 2 Quarryfield West Treanscabbagh Downamona Drum Firgrove

188800 187550 190790 190070 193460 182980 182150 180960 184810 186860 184250 164750 185080 193290 186850 180080 178150 190420 177710 185200 178890 192350 186650 191000 187430 179400 154460 153720 154250 159100 173650 185600 190300 185370

Size (ha) 130.4

Brierfield

52.9

N/A

26.3

N/A N/A Carrowkeel Turlough Castleplunket

60

Cloonloughlin

59.1

271390 280160 243470

Coolcam Lough N/A Corkip

67.1 9.2

246520 301600 243390 238170 251240 256080 288710 288790 272430 294270 255500 291580 290210 277050 264900 279450 272680 241390 273760 262350 281370 253920 303790 238600 249830 317450 313270 308210 308130 310200 312650 176050 198000 192520

Cuileenirwan Lough N/A Feacle Lough

SAC (SPA) name Ballinturley Turlough (SPA)

Four Roads Turlough (SPA) Coolcam Turlough Ballynamona Bog and Corkip Lough

15.7

Lough Funshinagh Kilnamryall Turlough N/A N/A Lisduff

54.1

Lisduff Turlough

Turlaghnamuck N/A Loughnaneane Turlough N/A N/A

31.8 12.4

Mullygollan Turlough

Rathnalulleagh Stonepark Turlough N/A N/A Turlaghmore Lough Croan Lough Nasool Turlough More Turloughbeg Turloughmore

26.4

106.9

19.5 25.7

Lough Croan (SPA)

Turloughmore (Sligo)

N/A Spring Park Wetlands Newchapel Turlough (continued on next page)

286

B I O L O G I C A L C O N S E RVAT I O N

1 3 3 ( 2 0 0 6 ) 2 6 5 –2 9 0

Appendix 1 – continued 297 298 299 300 301 302 303 304

Townland

County

Easting

Northing

Name

Gorteen Higginstown Kilgask Lismacrory Lismaline Raheen Upper Rahinane Cruckboeltane

Tipperary Tipperary Tipperary Tipperary Tipperary Tipperary Tipperary Westmeath

199400 222650 195420 196620 196180 188350 196850 245000

198050 140250 202560 198360 195700 138520 196400 262000

Ballingarry Ballynascullogue Sluggary Pool Liskeenan N/A N/A N/A Cruckboeltane

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26

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