Fresh insights into long-term changes in flora, vegetation, land use and soil erosion in the karstic environment of the Burren, western Ireland

Fresh insights into long-term changes in flora,

vegetation, land use and soil erosion in the karstic

environment of the Burren, western Ireland

Ingo Feeser* and Michael O’Connell

Palaeoenvironmental Research Unit, Department of Botany, National University of Ireland Galway, Galway, Ireland

Summary

1. The study focuses on species-rich, upland, heathy vegetation with arctic-alpine floristic affinities

and Sesleria grasslands in the karstic Burren region, western Ireland. The investigations aimed at

reconstructing the long-term development of these high conservation-value communities and the

role of farming in their formation and long-term survival.

2. The methods used included pollen analysis and 14C-dating of short monoliths and investigation

of grykes (fissures in karstic limestone) for evidence of soil erosion. Special attention was paid to

fossil, coprophilous fungal spores as indicators of local grazing. The strong local character of the

pollen records facilitated identification of inter-site differences as well as regional patterns. It is

shown that open pine woodland characterized the Cappanawalla uplands between c. 1500 BC and

500 BC. It is proposed that such woodlands, with floristic affinities to Scandinavian open pine

woodlands on calcareous soils, provided a suitable environment for the present-day, open heath

vegetation with species such as Dryas octopetala, Arctostaphylos uva-ursi, Geranium sanguineum

andEmpetrum nigrum.

3. Burning of vegetation as a management tool was important in the uplands over most of the last

two millennia. Firing seems to have ceased with the onset of more intensive grazing regimes in the

18th century.

4. Synthesis. Upland palaeoarchives, derived from shallow peaty deposits, show that the upland

Burren supported mainly plagioclimax Corylus-dominated woody vegetation and grasslands from

c. 1500 BC (beginning of present record), until possibly as late as the 17th century AD. In the

uplands of the north-western Burren, open, species-rich pinewoods with hazel dominated. The

northern-arctic elements in the present-day upland flora survived clearances, involving initially

Pinus sylvestris (c. 500 BC) and subsequently Corylus avellana (c. AD 1600). Colluvial material

retrieved from grykes supports the idea of considerable soil loss occurring as late as the first and

early 2nd millennium AD. The investigations highlight the potential of upland palaeoarchives,

consisting of short sequences, for elucidating vegetation and land-use dynamics in karstic environ-

ments such as the Burren.

Key-words: biogeography, coprophilous fungal spores, environmental change, karst, land use,

late Holocene, non-pollen palynomorphs, pollen analysis, soil erosion, vegetation dynamics

Introduction

The Burren, a mainly upland region extending over some

300 km2 in north-west County Clare, western Ireland (Fig. 1),

is internationally recognized for its karstic geology, an excep-

tional flora that includes arctic-alpine and northern elements

often growing in great abundance and in close proximity to

southern elements, and a rich archaeology that testifies to

settlement and farming extending back to the early Neolithic

(c. 4000 cal. BC; dates are given in calibrated years BC ⁄AD,

except 14C dates which are cited as BP, i.e. before AD 1950).

Research by Irish and visiting scientists has elucidated several

aspects of this region including the geology and soils (Finch

1971; Pracht et al. 2004), vegetation (Ivimey-Cook & Proctor

1966; Kirby 1981) and environmental plant physiology

(Osborne et al. 2003). More recently, the role of management

— essentially extensive farming as influenced by the EU Com-

mon Agricultural Policy (CAP) and specific instruments of

CAP such as the Rural Environment Protection Scheme*Correspondence author. E-mail: [email protected]

Journal of Ecology 2009, 97, 1083–1100 doi: 10.1111/j.1365-2745.2009.01533.x

� 2009 The Authors. Journal compilation � 2009 British Ecological Society

(REPS) — has received considerable attention (Sheehy Skeff-

ington & Gormally 2001; Dunford 2002). Much of this infor-

mation has been collated and synthesized in general

overviews (D’Arcy & Hayward 1992; O’Connell & Korff

2001; Jones 2004) and detailed accounts (e.g. Webb & Scan-

nell 1983; Osborne & Jones 2003). Despite these advances,

however, the impression remains of a ‘‘false sense of security’’

provided by the wealth of ‘‘descriptive biological detail’’

whereas ‘‘the truth is we don’t … understand the factors that

have shaped the present communities associated with the Bur-

ren’’ (Osborne & Jeffrey 2003, p. 107).

The present paper is conceived as a contribution to

improved understanding of the Burren and, specifically, the

plant communities and soils of the Burren uplands (altitude c.

300 m a.s.l.). These areas are home to some of the most dis-

tinctive plant communities and habitats, such as the Arcto-

staphylos uva-ursi—Dryas octopetala heaths (cf. Ivimey-Cook

& Proctor 1966) on the Cappanawalla–Gleninagh–Black

Head uplands in the north-western Burren and the Calluna

heaths and species-rich limestone grasslands on Gortaclare

and Slieve Rua–Mullach Mor uplands in the eastern Burren.

The particular aim of these investigations is the recon-

struction of environmental change in the areas referred to,

and, specifically, to determine if farming played a key role

in the establishment and long-term maintenance of these

species-rich communities or if, on the other hand, these com-

munities are best regarded as representing ‘‘a true climax ...

within a certain range of altitude and exposure’’, as proposed

by Ivimey-Cook & Proctor (1966, p. 251) for the Arctostaph-

ylos uva-ursi–Dryas octopetala heaths. Attention is also

focussed on possible widespread erosion and hence large-

scale loss of soil as a result of early farming. Techniques

employed include pollen analysis and AMS 14C dating and

probing of grykes, i.e. fissures in the karstic limestone, for

evidence of former soils. Particular attention is given to

identification of critical pollen taxa and also non-pollen

palynomorphs (NPP) including coprophilous fungal spores

(CFS) which are particularly useful for the reconstruction of

grazing activity (cf. van Geel et al. 2003; Blackford & Innes

2006; Davis & Shafer 2006; Feeser & O’Connell 2010).

Site descriptions

CAPPANAWALLA

The plateau-like upland of Cappanawalla (c. 300 m a.s.l.;

Fig. 1) is characterized by extensive limestone pavement with

deep grykes and patches of shallow rendzina-like soil. Shallow

Fig.1. Contoured maps showing the extent of the Burren region (eastern lowlands excluded) and details of the three study areas, Cappanawalla

(CPW), Gortaclare (GTC) and Sliabh Rua (RUA) (the squares are 10 · 10 km). Location of sampling sites, including Lios LairthınMor (profile

LLM II; Jelicic &O’Connell 1992), is indicated. Archaeological features are after Robinson (1999) and OSDiscovery Seriesmaps, with additions

andminormodifications.

1084 I. Feeser & M. O’Connell

� 2009 The Authors. Journal compilation � 2009 British Ecological Society, Journal of Ecology, 97, 1083–1100

depressions with peat deposits, mainly developed on minero-

genic, probably drift-derived soils several decimetres thick, are

a feature of the area.

Cappanawalla is one of five upland areas in the Burren of

exceptionally high floristic interest (Webb & Scannell 1983).

The heath communities are definedmainly byCalluna vulgaris,

Dryas octopetala, Arctostaphylos uva-ursi and Empetrum

nigrum (higher plant nomenclature follows Stace 1997). Erica

cinerea, Succisa pratensis, Carex panicea and C. binervis are

common; other wet heath and bog species include Narthecium

ossifragum,Trichophorum cespitosum andParnassia palustris.

Species associated with the upland limestone, i.e. pavement,

grykes and small patches of shallow soils, include Sesleria

caerulea, Thymus polytrichus, Achillea millefolium, Phyllitis

scolopendrium, Polystichum setiferum, Cystopteris fragilis and

the neophyte Mycelis muralis. Like most Burren uplands, the

Cappanawalla plateau is extensively grazed, mainly by cattle,

in winter as part of a transhumance-type practice known as

winterage (Dunford 2002).

On the slopes of Cappanawalla, the vegetation changes

with altitude, bedrock and soil characteristics, and land use.

The eastern slopes, which have little soil cover, harbour the

more floristically interesting vegetation. Dryas octopetala is

locally abundant above c. 100 m a.s.l. and Empetrum nigrum

occurs regularly above c. 200 m a.s.l. Calluna-dominated

heath with Molinia caerulea is locally common along the

mid-slopes and Filipendula ulmaria is frequent. At lower

elevation, above the intensively managed valley-floor

pastures, open scrubby vegetation with Corylus avellana,

Crataegus monogyna, Prunus spinosa, Ilex aquifolium, Rosa

pimpinellifolia is common, while in open rocky parts Solidago

virgaurea, Campanula rotundifolia, Geranium sanguineum and

Pteridium aquilinum are common.

GORTACLARE

Gortaclare, together with Slievecarran, forms the most exten-

sive upland in the eastern Burren (Fig. 1). Calluna heath and

short-sward Sesleria-rich limestone grassland characterizes the

southern part of the uplands. Heath is common here, its devel-

opment being favoured by the frequent outcropping of dark

grey chert that forms isolated nodules, ridges and

semi-continuous beds. Apart from the occasional, wind-bent

specimen of C. monogyna, the uplands are bare of tall woody

vegetation. At lower elevations, e.g. the eastern side of

Slievecarran and the Glen of Clab, Corylus scrub, with

Fraxinus excelsior, is extensive and largely dominant. The

uplands, like at Cappanawalla, are important for winterage

(Keane & Sheehy Skeffington 1995). The valley floors, in con-

trast to the uplands, have substantial, drift-derived soil cover

that supports rather intensive, pastoral-based farming.

SLIABH RUA

Sliabh Rua lies at the centre of the Knockanes–Sliabh Rua–

Mullach Mor uplands (Fig. 1) which, together with the sur-

rounding lowlands, constitute the Burren National Park. The

soils, which are very patchy, show considerable small-scale var-

iation in mineralogical composition and glacial origin, but

apparently do not include loess deposits (Moles & Moles

2002), as has been postulated (Moles et al. 1995; Jeffrey 2003).

The uplands carry mainly Sesleria-dominated, limestone

grassland on shallow soils intermingled with bare limestone

pavement often with well-defined grykes. There are extensive

patches ofCalluna-dominated heath, especially between Sliabh

Rua andMullachMor and on the side of Knockanes. Cultiva-

tion ridges, indicative of arable farming (probably potato

growing), occur on drift on the north-eastern flank ofMullach

Mor (Moles &Moles 2002).

ARCHAEOLOGICAL F IELD EVIDENCE FOR PAST HUMAN

ACTIV ITY

A rich archaeological field assemblage points to substantial

human activity from Neolithic times onwards (last c.

6000 years) in the Burren, but, surprisingly, little has been

published on the upland areas that were investigated here

(Fig. 1). Features noted on Cappanawalla include a ringfort

(diameter c. 20 m), a cairn and several stone-wall enclosures

including possible hut sites. Several low walls, including

so-called mound, tumble and slab walls (possibly of Bronze

Age, Iron Age, and late Iron Age or mediaeval period, respec-

tively), are present, superimposed on which are high, so-called

single walls that are probably nomore than a few centuries old

(dating of walls after Plunkett Dillon, 1985, and Jones, 2004).

Small U-shaped stone structures, presumably of relatively

recent origin and used for management of sheep and ⁄or goatsand possibly also for drying sods and cow pats as a fuel source

(Dunford 2001), are common. Stone walls, c. 5–10 m long and

about 20 in all, serve as windbreaks for cattle on the exposed

hilltop. The surrounding lowlands, on the other hand, are

much richer in archaeological features, indicative of a substan-

tial human presence from the Bronze Age onwards and espe-

cially during themediaeval period (Fig. 1).

Field evidence for past farming on Gortaclare consists

mainly of stone walls. Conspicuous single walls and also

mound, tumble and slab walls suggest active landmanagement

extending back to at least the Bronze Age. There is the occa-

sional stone-wall windbreak, presumably of rather recent ori-

gin (Fig. 2). Cairns, including a particularly large and

prominent cairn on Slievecarran, suggest substantial pre-his-

toric human presence (cf. also Turlough Hill where there is a

large enclosure, a very substantial cairn and c. 150 hut sites;

undated but possiblyNeolithic).

In the Knockanes–Sliabh Rua–Mullach Mor uplands, the

main archaeological features are cairns on Mullach Mor and

Knockanes (Fig. 1).

Materials and methods

FIELDWORK

Cappanawalla.A small depression, c. 5 · 4 m, on a shelf to the south

of the highest part of the plateau, was selected for sampling (Fig. 1;

Long-term environmental change in the Burren 1085


Table 1). The monolith, CPW1, was taken in August 2006 where the

deepest sediments were noted using a gouge corer (Fig. 2). Common

species at the sampling site included Calluna vulgaris (dominant),

Erica cinerea, Succisa pratensis, Carex spp., Lathyrus linifolius and

Potentilla erecta.

Gortaclare. The main sampling location, GTC7, lies towards the

southern end of the plateau (Fig. 1; Table 1) where there is a patch-

work of partially bare limestone, grassland and peat-covered chert

overlying limestone. Species of note include Dryas octopetala and

occasionally Rosa pimpinellifolia and Gentiana verna. Heathy vege-

tation with Calluna and a well-developed bryophyte layer domi-

nates where peat forms the substrate. The monolith, GTC7,

was taken in May 2006 from peat on a pronounced chert ridge,

c. 10 m long and elevated c. 50 cm above the surrounding lime-

stone grassland (Fig. 2).

On a c. 40-m wide terrace — referred to as the Gortaclare terrace

— that lies c. 5 m below the main plateau on the north-western side,

sediments were found in grykes (Fig. 1, GTC1–5). Here a c. 5-m

wide strip of jointed limestone pavement with patchy grassy cover

formed a zone, distinct from otherwise densely vegetated areas

(grassland and Calluna-dominated heath) with less jointing and

fewer grykes.

Sliabh Rua. In the Knockanes–Sliabh Rua–Mullach Mor uplands,

grykes and heath-covered surfaces were probed using a gouge corer

and spade. Potentially suitable sites were identified only in the saddle

between Sliabh Rua and Mullach Mor where Calluna on shallow

peat dominates over a relatively extensive area. Otherwise, the area

is characterized by jointed limestone pavement with patchy grass-

land and heath. The peat deposits, resting on up to several-decimetre-

thick minerogenic sediments, proved to be very shallow (maximum

Fig.2. Photographs of monoliths and sam-

pling sites. At top, monolith Cappanawalla

(CPW1) (left) and removing the monolith

(right); limestone grassland ⁄ heath domi-

nates in the foreground and the background

consists mainly of bare limestone pavement

(view to SW; 21 August 2006). At bottom,

monolithGortaclare (GTC7) (left; scale divi-

sions = 2 cm), and view of sampling area

(right) with Calluna-dominated heath and

stone-wall windbreak in the background (12

May 2006).

Table 1. Geographical location and other details of sites investigated

Location and

monolith ⁄ core number Latitude ⁄ longitudeAltitude

(m a.s.l.) Comments

Cappanawalla

(CPW1)

53�7.062¢ N, 9�11.812¢ W 296 Monolith from a Calluna-dominated heath

on shallow peat that rested on a silty-sandy soil

Gortaclare

(GTC1 and GTC5)

53�4.931¢ N, 9�2.111¢ W 285 Cores GTC1–5 from grykes on terrace below main

plateau (co-ordinates relate to the end points,

GTC1 and GTC5). Calluna was common at GTC5

53�4.945¢ N, 9�2.041¢ W

Gortaclare

(GTC7)

53�4.646¢ N, 9�2.651¢ W 272 Monolith from peat on cherty limestone

RUA1–3 53�0.884¢ N, 8�59.672¢ W 167 Cores from grykes in grassland ⁄ heath;co-ordinates given for core RUA2

RUA4 53�0.881¢ N, 8�59.690¢ W 170 Shallow peat deposit that rested on silty-sandy soil;

examined only in field



depth noted: 18 cm in core RUA4) and were not investigated in

detail. Corings of grykes (RUA1–3; gryke depths 180–200 cm)

yielded c. 70 cm of minerogenic sediment with charcoal flecks near

the base.

LABORATORY INVESTIGATIONS

Samples of 2 cm3 from 1-cm thick slices were taken from monoliths

CPW1 and GTC7 for pollen analysis. Lycopodium clavatum spores,

in tablet form, were added to facilitate calculation of pollen concen-

trations (Stockmarr 1971; tablets supplied by Department of Geol-

ogy, Lund University, Sweden). Standard treatments were given that

involved use of KOH, HCl and HF, and acetolysis (cf. Moore et al.

1991). At the end of the preparation procedure, the pellets were sieved

through a 5-lm mesh sieve in an ultrasonic water bath to remove

unwanted fine particles. The samples were mounted in glycerol and

counted using a Leica DM 4000B microscope (Leica Microsystems,

Wetzlar, Germany) at ·400magnification; phase contrast ·1000mag-

nification was used in the case of critical grains.

Counting, data calculations and plotting of pollen diagrams were

carried out using CountPol 3.2, a java program by I. Feeser. A

pollen sum (PS) of about 1000 terrestrial pollen grains, excluding

heath taxa, was generally achieved. For spectra with exceptionally

large numbers of heath taxa or exceedingly poor pollen concentra-

tion, a lower PS was accepted. Percentage representation was calcu-

lated on the basis of total terrestrial pollen (TTP, including fern

spores), but excluding heath taxa which are expressed relative to

TTP.

Pollen and spore identification follows mainly Moore et al.

(1991) and Beug (2004). As regards cereal-type pollen, the pollen

and annulus-size criteria of Beug (2004) were followed, but modi-

fied as follows. Four pollen-size categories were distinguished (max-

imum length cited), i.e. 37–39 lm, 40–44 lm, 45–49 lm and

>50 lm. Secale pollen were noted separately. The smallest size cat-

egory was subsequently included in the Poaceae (undifferentiated)

because of the high probability that at least some, and probably

several, of the grains were from non-cultivated grasses (see Inter-

pretation of data). Ericaceous pollen were distinguished using

Moore et al. (1991) and modern reference material. Non-pollen

palynomorphs (NPP), i.e. microfossils other than pollen, were

identified using published photomicrographs and descriptions (e.g.

van Geel (1978); Pals et al. (1980); Bakker & van Smeerdijk (1982);

van der Wiel (1982); van Geel et al. (1989, 2003)). Conspicuous

NPP that were regularly observed but not identified with certainty

were assigned a type number, e.g. Type BRN1 (BRN to indicate

Burren).

Material retained after initial sieving of the pollen samples

(>100 lm) was scanned under a stereo-microscope for macrofossil

remains and other matter including charcoal fragments (referred to as

macro-charcoal as distinct from micro-charcoal recorded on the pol-

len slides). Abundances were noted using a four-point scale (rare, +;

occasional, 1; frequent, 2; abundant, 3).

To obtain material suitable for AMS 14C dating, 1-cm thick slices

were taken from the monoliths, sieved and washed with distilled

water, and scanned for macrofossils. As far as possible, terrestrial

macrofossils of known origin were submitted for AMS 14C dating.

Loss-on-ignition at 550 �C (LOI) was determined using c. 2-cm3

samples taken from 1-cm thick slices. Procedures recommended by

Heiri et al. (2001) were adhered to.

In addition to the above investigations, charcoal fragments (at

most a few millimetres long, i.e. too small to be identified) were

picked from basal sediments retrieved from grykes on Gortaclare

and Sliabh Rua (cores GTC1 and RUA2 and 3) and submitted

for dating.

Results

DATA PRESENTATION

Stratigraphical descriptions of the monoliths are presented in

Table 2. Photographs showing monoliths and sampling in the

field are provided in Fig. 2. Pollen and NPP curves, LOI and

macrofossil (pollen sievings) data relating to profiles CPW1

and GTC7 are presented in Figs 3 and 4. Local pollen assem-

blage zone (PAZ) boundaries are indicated where at least one,

and usually several, pollen curves show substantial changes.

Results from constrained incremental sum of squares cluster-

ing (CONISS), carried out using ZONE (ver. 1.2; program by

S. Juggins), supported the zone boundaries as distinguished by

eye. An overview of the CFS and indeterminate NPP is pre-

sented in Table 3 and photomicrographs are presented in

Fig. 5.

Results from the 14C dating are given in Table 4. The 14C

dates were calibrated using the program OxCal 4.0 (Ramsey

2001) and the IntCal04 calibration curve (Reimer et al. 2004).

Table 2. Stratigraphical descriptions for monoliths CPW1

(Cappanawalla) andGTC7 (Gortaclare)

Depth (cm) Description

Monolith CPW1

0–6 Poorly decomposed peat with many

roots. Brownish black (7.5 YR 2 ⁄ 2)*6–22 Well-decomposed peat with fibres.

Dark reddish brown (5 YR 3 ⁄ 2)22–29 Black organic-rich clayey silt with

few fibres. Black (7.5 YR 2 ⁄ 1)29–32 Silty clay with c. 1% small stones

up to 3 mm in diameter. Brownish

black (7.5 YR 3 ⁄ 1)32–36 Silty clay. Brownish grey (5 YR 5 ⁄ 1)

with greyish brown (7.5 YR 4 ⁄ 2) toorange (7.5 YR 6 ⁄ 8) mottling

36–45 Clayey silt. Orange (7.5 YR 6 ⁄ 8)

Monolith GTC7

0–15 Dark brown, well decomposed peat

with roots (brownish black, 7.5 YR 2 ⁄ 2)15–20 Poorly decomposed peat with moss

remains and a minerogenic component

(dark reddish brown, 5 YR 3 ⁄ 2)20–35 Dark brown highly decomposed peat

with some rootlets (brownish

black, 7.5 YR 2 ⁄ 2)35–40 Dark grey brown peaty silt; more

minerogenic and sandy towards the bottom

(brownish black, 7.5 YR 3 ⁄ 1)40–42 Sandy minerogenic deposit with a substantial

organic component; small stones

(brownish black, 7.5 YR 3 ⁄ 1)

*Colour chart by Oyama & Takehara (1970) was used for assign-

ing colours.



Fig.3.Percentagepollen

diagramsfrom

Cappanawalla(C

PW1)an

dGortaclare

(GTC7).Thecurves

are

plotted

tothesamehorizontalscale,exceptCalluna(horizontalscalereducedby10

relative

totheother

curves).Silhouettes(truncatedwhereappropriate)showvaluesexaggerated

·10.T

axanotedoutsideroutinecountingare

indicatedby+.



Fig.4.Plotsofthefollowingdata,p

rofilesCappan

awalla(C

PW1)andGortaclare

(GTC7):non-pollen

palynomorphs(N

PPs)andmicro-charcoalcurves

(percentagevalues),LOI,

14Cdates;m

a-

crofossils(abundance),andconcentrationandpollen

accumulationrate(PAR)curves.S

ilhouettesand‘+

’values

havesameconnotationsasinFig.3.



INTERPRETATION OF DATA

Interpretation of pollen analytical data is seldom straightfor-

ward and this is particularly true when the material under

investigation derives from sources that are rather unconven-

tional, as in the present instance. Estimating the pollen source

area is crucial, i.e. the relative contribution of pollen of local,

regional and extra-regional or long-distance origin (Jacobson

& Bradshaw 1981; Court-Picon et al. 2006). In this instance,

the local component is regarded as deriving from the upland

vegetation and, in particular, the area close to the sampling

site. As regards the regional component, its contribution is

expected to be strongly influenced by local conditions such as

wind speeds and direction, and dispersal capacity, which for

non-arboreal pollen (NAP) generally and especially for sev-

eral critical Burren taxa is low. In the case of percentage

curves, the inter-dependence of the curves must also be borne

in mind. In the present instance, Calluna (also other heath

taxa) is regarded as deriving mainly from vegetation at or

near the sampling sites and hence will tend to be strongly

over-represented (cf. Moore et al. 1986; Mulder & Janssen

1999). Calluna and other heath taxa are therefore excluded

from the pollen sum. To overcome the disadvantages inherent

in percentage data, concentration and pollen accumulation

rate (PAR) curves are also presented (Fig. 4; selected taxa

only).

Reconstruction of the history of typical Burren upland

species such as Gentiana verna, Dryas octopetala and

Arctostaphylos uva-ursi is difficult because of the severe

under-representation of most of the species in the pollen

record. Indeed, some of the more interesting species are com-

pletely silent, e.g. the orchids including Neotinea maculata.

Another hurdle is that some of the pollen taxa may include

pollen of several species; e.g. Mentha-type which, in the

present context, most likely consists more or less exclusively

of pollen of Thymus polytrichus. On the positive side, the

deposits used here are particularly well suited to archiving

the pollen of under-represented species, because many of the

species in question can be expected to have been present at or

near the point of sampling. Furthermore, the overall low

Table 3. Overview of the coprophilous fungal spore types (CFS) (Groups A and B) and indeterminate non-pollen palynomorphs (NPP)

(Group C)

Type Classification ⁄ description ⁄ ecology

Group A: identified coprophilous fungal types*

Sordaria-type (T. 55A and T. 205) Includes various species of the Sordariales (van Geel et al. 2003). Sordariales are mainly

coprophilous, but occur also on rotting wood and soil (cf. Lundqvist 1972)

Sporormiella-type (T. 113) Widely recognized coprophilous spore type indicative of grazing (e.g. Blackford & Innes 2006;

Davis & Shafer 2006; Raper & Bush 2009). The common species, S. intermedia, occurs on

the dung of most herbivores (Dennis 1978)

Tripterospora-type (T. 169) This type derives probably from a coprophilous Sordariales species and possibly the genus

Tripterospora (van Geel et al. 2003)

Arnium-type (T. 261) Spores of this type derive from the Sordariales and probably the genus Arnium

(van Geel et al. 2003). According to Dennis (1978), most Arnium species occur on dung,

but some species are also abundant on rotting herbaceous stems and wood

Podospora-type (T. 368) This type is regularly recorded in samples from archaeological sites (van Geel et al. 1981, 2003)

and appears to be connected with grazing (cf. Blackford & Innes 2006). Kirk et al. (2001)

regard Podospora species as coprophilous

Group B: additional possible coprophilous fungal types recorded in these investigations

BRN4 (cf. Delitschia sp.) Brown spore c. 29 · 13 lm with truncated base and an apical pore (c. 1.5 lm). The spore

as illustrated (Fig. 7) was probably originally two-celled (possibly a Delitschia species†)

BRN6 (cf. Podospora) Elliptical, non-septate, brown fungal spore, c. 23 · 45 lm with truncated base and a bent

tip with a small pore; resembles spores of Podospora (cf. Lundqvist 1972)

BRN7 (cf. Schizothecium conicum) Elliptical, non-septate, fungal spore, c. 16 · 25 lm with protruding apical pore. It resembles

spores of S. conicum, a common coprophilous species†

BRN8 (cf. Hypocorpa stercoraria) Ellipsoidal, dark brown spore, c. 45 · 21 lm, with longitudinal slit. It resembles

H. stercoraria, a Xylariales fungus that produces perithecia in stromata embedded in dung†

BRN9 (cf. Podospora ⁄Cercophora) Brown, ellipsoidal spore, c. 12 · 20 lm, truncated at one end and a slightly subapical pore

opposite; resembles spores of Podospora and Cercophora (cf. Lundqvist 1972)

Group C: regularly encountered indeterminate NPPs‡

BRN1 Globose microfossil (c. 20 lm diameter) with c. 1.5 lm long processes

BRN2 Spindle-shaped fungal spore (ascospore?), c. 18 · 37 lm, with pores at both ends; one end

tapered

BRN3 Bi-septate fungal spore c. 17 · 33 lm, with truncated pore-like endings at one or both ends.

It may derive from a dematiaceous hyphomycete (includes plant parasitic, saprophytic and

soil fungi)

BRN5 Brown, 2–3 septate fungal spore, c. 23 · 16 lm. This type probably derives from a

dematiaceous hyphomycete

*T. = Type as described in van Geel et al. 2003; †M. Richardson, pers. comm.; ‡Provisional type numbers with prefix BRN (= Burren).



input of arboreal pollen (AP), combined with substantial

pollen counts, increases the probability of recording critical

pollen taxa that have low representation.

Regarding the NPP and especially the CFS, these taxa often

represent plants and animals that were present at the coring

site. CFS are regarded as deriving from species that grew

mainly on herbivore dung. Spore production in these fungi is

usually high but dispersal is comparatively poor because of the

location of fruiting bodies close to the ground. CFS are there-

fore regarded as indicative of local grazing (cf. Blackford &

Innes 2006;Mazier et al. 2009).

The diversity and occurrence of coprophilous fungi is

influenced by various factors. Generally, they show prefer-

ences for dung of particular herbivore species (Richardson

1972, 2001). Changes in the CFS assemblage are known to

be influenced by the grazers, local environmental factors

and the local vegetation, i.e. the diet of the animal produc-

ing the dung can be important. Although it cannot be auto-

matically assumed that all the spore types classified as CFS

derive exclusively from dung, good agreement of the avail-

able records with NAP indicators of pastoral activity sup-

ports such an interpretation.

CHRONOLOGY — MONOLITHS

Age–depth curves for profiles CPW1 and GTC7 are presented

in Figs 6 and 7, respectively.

Three 14CAMSdates are used in constructing the age–depth

curve for CPW1. The 14C date, 440 ± 35 BP, was considered

to be unreliable by the 14C laboratory due to small sample size

and so was not used. At 10 cm depth, the so-called secondary

rise in Pinus is evident and AP shows some recovery; these fea-

tures suggest that this level dates to c.AD 1850. The top of the

monolith is assigned to AD 2005. The basal 14C date,

3290 ± 40 BP, derives from fine particulate charcoal. The

charcoal particles are probably broadly contemporaneouswith

the deposition of the minerogenic matrix in which the charcoal

occurs so the date is regarded as indicative of the age of deposi-

tion. Alternatively, the charcoal (also the pollen) may have

been worked into an existing deposit. In either scenario, the14C date suggests that the base of the pollen profile relates to

the mid Bronze Age (c. 1550 BC). The basal lithological unit

(36–45 cm) probably represents glacial drift.

The age–depth curve for profile GTC7 relies on the three

available 14C AMS dates, the date AD 1850, which marks the

Fig.5. Photomicrographs of selected non-

pollen palynomorphs (NPPs) including

coprophilous fungal spores (CFS). 1: Sorda-

ria-type (Type 55A), 2: Sordaria-type (Type

205), 3: Tripterospora-type (Type 169), 4:

Sporormiella-type (Type 113), 5: Podospora-

type (Type 368), 6: Arnium-type (Type 261),

7: Type BRN4 (cf. Delitschia sp.), 8: Type

BRN6 (cf. Podospora), 9: Type BRN7 (cf.

Schizothecium conicum), 10: Type BRN8 (cf.

Hypocorpa stercoraria), 11: Type BRN9

(cf. Podospora ⁄Cercophora), 12: Type

BRN2 (unknown fungal spore), 13: Type

BRN1 (unknown NPP), 14: Type BRN5

(unknown conidia), 15: Type BRN3

(unknown conidia).



point where AP begin to increase (cf. CPW1 above), and AD

2005, which is assigned to the top of the profile. The interval

between the dates 545 ± 45 BP and 1735 ± 35 BP has great-

est uncertainty because of changes in lithology and the basal

context. The assumption is made that an accumulation rate

defined by a straight line connecting the dates 255 ± 30 BP

and 545 ± 45 BP is best applied down to 35 cm, i.e. to the

point where peat begins to predominate. On this basis, depth

35 cm dates to c. AD 1200. The basal date, 1735 ± 35 BP,

suggests that the deposit began to accumulate aroundAD300.

The interpretations of the pollen profiles CPW1 and GTC7

follow. The chronology, including that suggested for each

PAZ, is the best approximation based on the relevant age–

depth curve.

CHRONOLOGY — CORINGS IN GRYKES

The 14C date, 1995 ± 35 BP, derives from charcoal fragments

in core GTC1, Gortaclare terrace. The core was 41 cm long

(uppermost c. 10 cm of rather unconsolidated deposit disre-

garded). The lower sediment, from which the dated charcoal

derives, wasminerogenic (silty) and yellowish brown in colour.

It is assumed that the charcoal does not have a large inherent

age and thus gives an accurate indication of the age of deposi-

tion. The 14C date indicates that colluvial accumulation began

at about the BC–AD transition (late IronAge; Table 4).

Cores RUA1–3 from Sliabh Rua yielded c. 70 cm of miner-

ogenic material, with charcoal fragments (fleck-like) common

in the lowermost c. 8 cm. Fragments from cores RUA2 and

RUA3 yielded 14C dates that suggest deposition in the late

11th to 12th and the late 4th to 5th century AD, respectively

(Table 4).

CAPPANAWALLA — RECONSTRUCTION OF

VEGETATION AND LAND-USE CHANGE

PAZ CPW1-1 (c. 1500–300 BC). This PAZ derives from a

highly minerogenic deposit. The basal sample had particularly

low pollen concentration and the pollen tends to be corroded

(6% of TTP compared with 2.4% and 0.9%, respectively, in

the overlying spectra). On the other hand, corrosion-resistant

fern spores are not particularly strongly represented (though

there is a substantial Botrychium curve). The latter suggests

that these spectra are not seriously distorted by differential

corrosion.

The relatively high Pinus and Corylus values indicate that

the uplands of Cappanawalla carried pine and hazel-domi-

nated woodland. Given the strong over-representation of

Alnus in pollen records, alder is assumed to have been of minor

importance andmay have been locally absent.

The considerable NAP component points to a relatively

open woodland and also open landscape. Bare limestone

Table 4. Details of radiocarbon dates fromupland Burren sites

14C laboratory

number

(GrA-)

Sample

depth (cm)

Age

(14C years BP)

1r range ⁄2r range

Median age†

(cal. years) Material dated

Monolith CPW1

n ⁄ a 15–16 440 ± 35* AD 1428–1468 AD 1450 Erica cinerea seed, Hypericum seed,

Carex fruit fragment,

charcoal fragments, charred moss stems

AD 1413–1616

35324 22–23 305 ± 30 AD 1521–1644 AD 1560 Carex fruit fragment, charcoal fragments,

charred moss stemsAD 1489–1650

35003 28–30 2115 ± 35 196–93 BC 140 BC Four Carex fruit, charcoal fragments

348–45 BC

35323 36–37 3290 ± 40 1615–1519 BC 1570 BC Charcoal fragments

1682–1464 BC

Monolith GTC7

33180 21–22 255 ± 30 AD 1532–1797 AD 1650 Charcoal fragments

AD 1521–1954


AD 1310–1438

33177 40–41 1735 ± 35 AD 249–344 AD 310 Carex fruit, charcoal fragments

AD 230–402

Gouge core GTC1

33176 36–39 1995 ± 35 40 BC–AD 30 AD 10 Charcoal fragments

91 BC–AD 80

Gouge core RUA2


AD 1039–1215

Gouge core RUA3


AD 358–544

*Date not reliable due to very small sample size; a 14C laboratory number was not assigned.†Median date as given by the calibration program Oxcal 4.0. Dates are rounded to the nearest decade.



pavement as exists today was probably not as extensive in

Cappanawalla uplands at this time, i.e. minerogenic soils have

probably undergone considerable erosion in the meantime (see

below). These soils would have provided suitable edaphic con-

ditions for pine–hazel woodland. The substantial Succisa and

Calluna curves suggest heathy vegetation that may have been

present as understorey in open pinewoods, as well as occurring

in open situations. Scilla-type pollen suggests Hyacinthoides

non-scripta, a species characteristic of north-western European

woodlands. Indeed, H. non-scripta can be regarded as ‘almost

endemic’ to Ireland and Britain (Cross 2006, p. 111).

The strong representation of taxa such as Plantago lanceol-

ata, Liguliflorae and Cerastium-type point to open, rather

species-rich grassland. The initiation of a Plantago maritima

curve in the uppermost spectrum — it peaks at 6% — coin-

cides with greatly reduced Pinus representation and suggests

replacement of pine-dominated woodland by grassland with

much Plantago maritima. This plantain is common in many

present-day Burren grasslands, even at considerable distance

from the coast (Webb& Scannell 1983).

PAZ CPW1-2 (c. 300 BC–AD 1100). This PAZ relates to

the dark, charcoal-rich clayey silt uponwhich peat rests. Pollen

concentration and especially PAR values are very high. This

suggests that the deposit accumulated slowly, under conditions

favourable for pollen preservation (cf. increased LOI values;

Fig. 4). The high PARvaluesmay be partly due to an over-esti-

mation of the rate of deposit accumulation.

During this time, hazel was still common on the upland pla-

teau though, undoubtedly, hazel from the wider region also

contributes to the Corylus curve. The tall canopy trees, birch,

oak and elm, were probably also present, at least in the wider

north-west Burren region.Hedera, Ilex and possiblyCrataegus

were probably also locally present. Pine seems to have become

extinct, but there is evidence that it survived in the eastern Bur-

ren until c.AD 1400 (see below). Given pine’s tolerance of kar-

stic conditions (Willis et al. 1998; Roche et al. 2009), it appears

justifiable to ascribe its extinction in the Burren during late

mediaeval times to human impact.

Increased importance of grassland is indicated by the

expansion of Poaceae, Plantago lanceolata and several of

the NAP curves. This may have been favoured by increased

grazing pressure (but cf. Court-Picon et al. 2006), although

CFS have yet to be recorded. Increased landscape openness

and a tendency towards soil acidification are indicated by a

substantial increase in ericoid pollen (Calluna, Erica cinerea,

Empetrum and Arctostaphylos uva-ursi (single tetrad)).

The heathlands appear to be similar to those of today but

D. octopetala was not important, at least locally, until the

next zone (Dryas is severely under-represented in pollen

records so failure to record its pollen does not necessarily

imply absence). The lithology and the micro- and macro-

charcoal records suggest frequent firing. Erica cinerea and

Potentilla erecta (this is the most likely contributor to the

Potentilla-type curve), which are tolerant of firing, expand.

The firing is presumably connected with farming activity on

the plateau.

A notable feature of this profile, and also profile GTC7, is

the substantial cereal-type curve (mainly >3%). While most

of the grains are in the range 40–44 lm, several larger grains

were also recorded, including grains ‡50 lm, in both CPW1

and GTC7. It is reasonable to assume, given the records for

pollen of weeds associated with arable habitats and also the

documentary evidence for cereal growing (e.g. the official agri-

cultural statistics), that at least some of the cereal-type pollen

derive from cereals (further consideration of cereal-type pollen

in an upland Burren context in Jelicic & O’Connell 1992; cf.

also Behre 2007). Whether they reflect cereal cultivation in the

uplands is rather uncertain. Given the present-day unsuitabil-

Fig.6. Age-depth curve for profile Cappanawalla (CPW1). Median

age of 14C dates is shown (a dot) and also the 2r age ranges.

Fig.7. Age-depth curve for profile Gortaclare (GTC7). Median age

of 14C dates is shown (a dot) and also the 2r age ranges. Mid point of

the 2r age range of 255 ± 30 BP is used in constructing the age-

depth curve to better accommodate expected changes in accumula-

tion rate based on considerations of peat decomposition.



ity of the Cappanawalla uplands for cereal growing, the exten-

sive fertile soils of the nearby valley floors where there is also

strong evidence for settlement, and the frequency of strong

winds and open landscape that inevitably favour pollen trans-

port to the uplands, it seems reasonable to assume that the

cereal-type pollen recorded in CPW1 derive, probably

exclusively, from arable farming in the lowlands.

As regards the single record of Solanum nigrum-type in

zone CPW1-2, it is doubtful if this derives from S. tuberosum

as potato growing did not become widespread in Ireland

until, at earliest, the 17th century (Fagan 2000; Bell &

Watson 2008). A native species such as S. dulcamara is a

more likely source.

PAZ CPW1-3 (c. AD 1100–1700). During this interval,

hazel declines so that as the zone ends it is no longer of signifi-

cance. There is a corresponding expansion of grassland and

heath. Notable are the high values for Poaceae and Plantago

lanceolata, and good representation of Bidens-type pollen —

this potentially includes pollen of not only common species,

e.g.Bellis perennis, Solidago virgaurea and Senecio vulgaris, but

alsoAntennaria dioica, a typical andwidespread Burren species

of both limestone grasslands and heaths (Webb & Scannell

1983). Vicia-type pollen is well represented, particularly given

the poor pollen production and dispersal in the Fabaceae. This

taxon may include not only common species such as Vicia

cracca and V. sepium, but also Lathyrus linifolius, which is

common today on heathy Burren uplands. Curves forMentha-

type (probably exclusively Thymus polytrichus) and Dryas are

initiated in this zone. Overall, the heathy communities seem to

have been comparable to those of today. Local presence of

Corylus, Betula and Salix (probably S. repens) cannot, how-

ever, be ruled out. Heathers and Potentilla erecta dominated

at the sampling site and Calluna probably acted as host to

Meliola cf. niessleana (Type 14).

High values for akinetes ofZygnema (Type 314) in the upper

spectrum suggest moist conditions, which is also supported by

records for the rhizopod, Assulina. Calluna probably provided

favourable, moist micro-climate (cf. Mulder & Janssen 1999).

A general shift towards cooler and wetter conditions is also

plausible at about this time, which corresponds with the more

severe phase of the Little Ice Age climatic deterioration (Fagan

2000).

Increased grazing on the uplands is supported by sub-

stantial representation of several CFS in the upper spec-

trum. If arable farming was important in the lowlands (as

seems to be the case; cf. cereal-type curve), then upland

summer grazing, if only to facilitate cereal-growing in the

lowlands, is highly plausible. Heavy grazing pressure would

have led to the decline in hazel.

PAZCPW1-4 (c. AD 1700–1850).The decline in AP to 4%

suggests lack of woody vegetation at both local and regional

levels. By the latter part of the 18th century the population in

Ireland had begun to expand rapidly, and this expansion con-

tinued at an accelerated rate until severely checked and ulti-

mately reversed by the Great Famine (1845–47). As a

consequence of increased human impact, the 19th century saw

the final demise of woody vegetation in the Burren, with recov-

ery beginning only towards the end of that century (Kirby

1981, Jelicic & O’Connell 1992). These developments are

reflected in this zone and they also help to secure the chronol-

ogy. Re-introduction of pine and planting of beech, which was

well underway by the late 18th century in western Ireland

(Molloy & O’Connell 2004), is reflected in a rise in Pinus

(so-called secondary rise) and initiation of a curve for Fagus.

Grasslands have expanded, both locally and regionally, at

the expense of hazel (cf. Poaceae, Plantago lanceolata and Lig-

uliflorae). Other noteworthy NAP curves include Anthemis-

type (potentially includes pollen of Achillea millifolium and

Leucanthemum vulgare), Campanula-type (presumably

C. rotundifolia; C. trachelium seems unlikely given its present-

day rarity in the Burren (Roden 2001) and in Ireland gener-

ally), Rhinanthus-type, Linum cartharticum and Polygala (may

include pollen of P. serpyllifolia and P. vulgaris). The species

mentioned are intimately associated with dry, semi-open grass-

land and ⁄or heathy grasslands in the present-day Burren

(Ivimey-Cook & Proctor 1966; Keane & Sheehy Skeffington

1995).

Calluna declines presumably in response to high grazing

pressure. Foot (1864) remarks that wood was so scarcity in the

Burren that Dryas and Pteridium were collected as firewood.

The woody stems of Calluna would also have been valuable in

this context and so may have been used for fuel. Firing in the

uplands seems to be at its minimum, probably because

Calluna, which is often the incentive to fire so that fresh growth

is stimulated, was no longer important.

There is an overall decline in CFS, although Arnium-type

(Type 261) and Podospora-type (Type 368) spores assume

importance. These changes may reflect a shift in pasture man-

agement, including changes in the grazing regime. A shift in

spore representation may also have come about as a result of

the decline in Calluna, which would have resulted in a change

in dung composition and hence, indirectly, the coprophilous

fungal flora (cf. Richardson 2001).

Type BRN1, which achieves 27% at 13 cm, seems to be

associated with complete clearance and an intensive grazing

regimewhich is also true for profileGTC7.

The slender curve for Chenopodiaceae probably reflects the

importance,withina regional context, of potato cultivationand

the associated weed flora which is invariably characterized by

Chenopodiaceae species (cf. alliance Polygono-Chenopodion,

White & Doyle 1982). Solanum-type pollen (a single record)

most likelyderives, in this instance, fromS. tuberosum.

PAZ CPW1-5 (c. AD 1850–1950). This zone reflects the

regeneration of hazel in response to a decline in population

and farming pressures, and the planting of pine and beech in

the wider region. The modest expansion of Corylus to 5%

reflects the regeneration of hazel at a regional level.

The NAP and heath taxa suggest that the upland vegetation

remained largely unchanged. There is, however, less Mentha-

type and Rhinanthus-type pollen, and Polygala representation

increases. These pollen taxa probably represent mainly local

vegetation at the sampling site and so the changes may not be

of major significance as regards regional vegetation and land-

use dynamics. On the other hand, the Artemisia curve is inter-



rupted, suggesting a decline in tillage which characterized the

20th century.

Coprophilous fungi are poorly represented which suggests

further changes in the grazing regime, including possibly less

emphasis on active management. Interestingly, micro-charcoal

is well represented. Firing has not been a feature of land-man-

agement in the Burren in recent times. This, and lack of both

macro-charcoal and indications of burning in the upper part of

the monolith, suggest that the micro-charcoal is of regional

origin and hence not indicative of local firing as in PAZ 2 (see

above).

GORTACLARE – VEGETATION AND LAND-USE CHANGE

PAZ GTC7-1 (c. AD 100–1650). Corylus at c. 18% suggests

the presence of hazel scrub, at least in the region and possibly

locally on the plateau. Slender curves for Betula, Quercus and

Alnus suggest local or at least regional presence of these trees.

Pine was a veryminor component, probably at a regional level,

until c.AD1400 (base of subzone 1c) when it became extinct.

As organic matter began to accumulate, heathy vegetation

with much Calluna and Succisa dominated locally (subzone

1a). Throughout the period, grasslands with much Plantago

lanceolata, Liguliflorae (the latter is expected to include Leon-

todon, Pilosella and Taraxacum), and also Vicia and Lathyrus

spp. (cf.Vicia-type)were important.TypicalBurrenplants such

as Campanula rotundifolia, Galium spp. (most likely G. verum;

possibly also G. boreale, G. saxatile and Rubia peregrina) were

well represented in theuplandflora.

In subzone 1c, Poaceae and Plantago lanceolata play a less

important role than hitherto (cf. percentage and concentration

curves) while the Liguliflorae andCampanula rotundifoliawere

favoured. This is presumably due to changes in farming activ-

ity. Whereas CFS are rather poorly represented at the begin-

ning of the zone, improved representation in subzone 1c (c.

AD 1400–1650) suggests an increase in local grazing. Greater

representation of the testate amoeba Hyalosphenia subflava

points to peat disturbance (van Geel 1978) and dry conditions

(Charman 2001). Micro- and macro-charcoal are important

throughout the zone, and Gelasinospora spores and Cenocco-

cum sclerotia, both from fungi that have a strong association

with firing (van Geel 1978), are well represented. It seems that

firing, which is most likely related to farming activity, was

important on the plateau, especially as the zone ends (cf. sharp

peak in micro-charcoal). Firing would also have favouredCal-

luna andErica cinerea.

A substantial cereal-type curve (average 4.2% in subzones

1b and 1c) suggests considerable emphasis on cereal growing.

Amoreor less continuousArtemisia curve supports this.Cereal

growing was probably confined to the fertile lowlands but the

uplands, even if rather unpromising for cereal growing from a

present-day perspective, cannot be completely excluded.

PAZ GTC7-2 (c. AD 1650–1850). Low AP representation

(average 3.5%) suggests a fully open landscape, more or less

devoid of trees and shrubs. Low AP, high NAP and increased

CFS values suggest intensive farming. The distinct reduction

in Calluna is probably attributable to high grazing pressure

and possibly use of the woody stems as firewood (cf. CPW1-4

above). Firing of vegetation during this time, however, was

greatly reduced or ceased (cf. charcoal data).

During subzone 2a, Plantago lanceolata was exceptionally

well represented in the local grasslands. Increased representa-

tion of Pteridium, coinciding with a pronounced decline in

Corylus, suggests local clearance of hazel scrub during the late

17th century. In subzone 2b, the grassland composition

changed so that Poaceae, Ranunculus and Rubiaceae spp.

(cf. Galium-type curve) were favoured. Interestingly

also tubular composites were favoured over ligulates

(cf. Bidens-type and Liguliflorae curves). As observed in

CPW1-4, Type BRN1 peaks when grazing pressures were

high. Mineral matter was conspicuous in the peat (cf. also

lower LOI) which points to increased local disturbance.

These changes are probably attributable to severe grazing

pressures associated with a sharp rise in population

(subzone 2b spans c. AD 1800–1850).

PAZ GTC7-3 (c. AD 1850–1950). This zone reflects the

regeneration of Corylus and other woody species as a result

of decline in farming that began in the mid 19th century.

Evidence from other sources (e.g. Kirby 1981) indicates that

woodland and scrub regeneration was largely confined to

steeply sloping parts such as the eastern side of Slievecarran

and the Glen of Clab. There is no evidence to support

regeneration on the plateau. Grazing, however, seems to

have declined; cf. decline in NAP and especially Plantago

lanceolata, Liguliflorae and Galium-type, and increase in

Calluna and other ericoids. The NPP records consist mainly

of Assulina and Meliola cf. niessleana which are probably

associated with Calluna (see CPW1). Hyalosphenia subflava,

which is indicative of disturbance, is also well represented.

On the other hand, CFS are poorly represented. Casual

observations over recent years suggest that grazing density

is relatively low on Gortaclare compared with Cappanawal-

la. This seems to have been also true during the late 19th

and earlier part of the 20th century. A peak in micro-char-

coal at 11 cm (also dark, charcoal-enriched peat) suggests a

phase of heath firing, a practice that seems to have been

completely abandoned in recent times.

PALAEOENVIRONMENTAL INFORMATION DERIVED

FROM GRYKES

Grykes on Gortaclare terrace. Though several grykes were

probed on the plateau and surrounding area, substantial min-

erogenic deposits in grykes were noted only on Gortaclare ter-

race (see Fieldwork inMethods and materials). These deposits

are regarded as a colluvium that has arisen through soil loss

from nearby sloping ground. The charcoal fragments from

near the base of sequences were presumably deposited with

eroding soil. The 14C date 1995 ± 35 BP, which derives from

the charcoal, provides a reliable indication that the erosion

dates to about the BC–AD transition. It is assumed that it was

triggered by woodland clearance or intensive grazing. The ero-

sionwas probably contemporaneous with, or shortly preceded,

the beginning of theGTC7 record.



Grykes at Sliabh Rua. The two available dates indicate

erosion taking place in the early and mid to late mediaeval

periods (late 4th to 5th and late 11th to 12th centuries

AD), respectively, i.e. periods noted for increased levels of

human activity in western Ireland. Again, it is assumed that

the erosion resulted from disturbance of the vegetation

cover due to farming.

Discussion and conclusions

OVERVIEW OF ENVIRONMENTAL CHANGE AND LAND

USE IN THE BURREN UPLANDS

The palaeoecological evidence presented above provides fresh

insights into the developmental history of two important and

species-rich Burren upland habitats, and, furthermore, new

information on past soil erosion. To get a better overall per-

spective, the new data are considered in the context of results

from another upland investigation, namely that at Lios

Lairthın Mor, which lies south-west of the Ballyvaughan low-

lands (grid ref. 53�04.8¢ N, 09�13.26¢ W; 270 m a.s.l.; Jelicic &

O’Connell 1992; Fig. 1). The profile (LLM II; 192 cm long)

derives from a peat mound (diameter <50 m) that lies at the

junction of karstic limestone and a Namurian shale outlier.

Summary diagrams of the three profiles, drawn to an age

rather than a depth scale, are presented in Fig. 8. In LLM II,

the bog and heath pollen are presumed to arise mainly from

the sampling site and to a lesser extent from the large adjoining

bog to the east, while the terrestrial component is expected to

arise mainly from the local karst, which has minimal drift

cover. The main relevant archaeological evidence for land use

at Lios Lairthın Mor are the many ancient field boundaries

and also ringforts, the nearest situated within 80 m of the cor-

ing location, LLM II (Fig. 1).

Profiles CPW1 and LLM II span the interval mid Bronze

Age tomodern times while profile GTC7 opens in the late Iron

Age. The basal part of CPW1 suggests open pine–hazel wood-

land with grassland and heath in the interval c. 1500–300 cal.

BC, i.e. mid Bronze Age to mid Iron Age. At Lios Lairthın

Mor, on the other hand, the landscape was open, and hazel

was the main woody species. The relevant pollen spectra from

CPW1 are, however, from highly minerogenic sediments, so

rather different taphonomic processes are at play and also the

dating may not be as secure as LLM II. These considerations

aside, the overall evidence points to contrasting woodland

composition and cover (at Lios Lairthın Mor, pine unimpor-

tant or absent and landscape more open), presumably largely

due to different farming regimes and especially grazing

intensity.

In CPW1, strong human impact registers first in zone 2

(decline in Pinus and charcoal-enriched minerogenic sedi-

ments) and then increases distinctly at c.AD 1700 (top of zone

3; decline in Corylus and further expansion of Poaceae and

Plantago lanceolata). This sets a pattern that is maintained

until the mid 19th century when regeneration of woody vegeta-

tion commences (zones 4 and 5). Changes over the correspond-

ing intervals, i.e. from c. 300 BC to the present day, are

recorded in greater detail in LLM II. A distinctive feature is

the regeneration of woody vegetation, involving mainly hazel,

ash and birch as a response to a steep decline in farming, i.e.

the so-called Late Iron Age Lull (cf. Molloy & O’Connell

2004), which, in this instance, is dated to c.AD 300–600 (zone

LLM II-2). Human impact, including arable farming, is partic-

ularly strong from about AD 1100, but the first intensive clear-

ances in the partially wooded landscape begin at about AD

1650, and final clearance of hazel scrub begins a century later

(see below).

Profile GTC7, although covering a shorter interval (c. AD

100–1950), has the advantage of a higher sampling resolution

than CPW1. The Gortaclare uplands carried hazel scrub until

c. AD 1650 or, alternatively, the nearby lowlands had more

scrub than today.

Near the base of GTC7 (zone 1a, 39 cm; c.AD 500), there is

a decline in Plantago lanceolata and high Calluna values. This

may be an expression, albeit in subdued form, of a Late Iron

Age Lull. The abandonment of active farming may have

resulted in conditions suitable for peat initiation (cf. Molloy &

O’Connell 1993).

At c.AD 1500, there is evidence for increased farming activ-

ity in both GCT7 (subzone 1c) and LLM II. This may be part

of a general recovery subsequent to the Black Death, which

reached Ireland in AD 1348 (Kelly 2001; Yeloff & van Geel

2007).

The charcoal records from GTC7 and CPW1 indicate that

burning of upland heaths, most likely connected with pastoral

farming, was important until at least the 17th century. Further

research into fire history and the long-term impact of firing

upland heaths in the Burren is desirable.

The evidence from all three sites points to major change,

particularly from the 17th century onwards, that finds expres-

sion mainly in the decline of Corylus in the context of strong

farming pressures (GTC7-2, top of CPW1-3 and CPW1-4,

LLM II-5). This coincides with the laying out of large field sys-

tems in the Burren uplands between c.AD1650–1750, possibly

in connectionwith a shift from subsistence farming to amarket

economy (Plunkett Dillon, 1985). During this time sheep rear-

ing became increasingly important. Dutton (1808), for

instance, reports that large flocks of sheep were being reared

on a year-around basis in the western Burren uplands. High

Potentilla-type values at about this time in profile CPW1 point

to increased abundance and flowering of Potentilla erecta, a

development probably favoured by intensive sheep grazing (cf.

Moore et al. 1986).

Exceptionally low Corylus representation at about c. AD

1800 in all three profiles highlights the extreme scarcity

of shrubby vegetation (even Calluna was adversely affected).

This corresponds with high representation of CFS which

emphasizes the importance of upland grazing in the farming

economy.

The relatively strong recovery in woody vegetation, and

especially hazel, at a regional level, as farming pressures

declined and tree-planting became more common, is

reflected in the uppermost zone of all three profiles. The

phenomenon of hazel-scrub expansion of the last few dec-



Fig.8.Selectedpercentagecurves

from

Gortaclare

(GTC7),Cappanawalla(C

PW1)an

dLiosLairthın

Mor(profileLLM

II)plotted

toatimescale.Ulm

usispoorlyrepresentedin

GTC7andso

hasbeenomitted.T

hecoprophilousfungalspore

curves

includethose

taxaclassified

ingroupsAandBin

Table3.



ades (cf. ERA-Maptec 2006) is probably not captured as

sampling stopped short of the uppermost deposits. A shift

in the composition of the upland grassland communities is

also notable (cf. NAP curves and especially Plantago

lanceolata), presumably as a result of changes in grazing

regimes (cf. decrease in CFS).

EVIDENCE FOR LOSS OF SOIL COVER IN THE BURREN

Apart from most of the lowlands, extensive smooth pavement

and ⁄or shattered limestone terrain with little or no soil cover

characterize much of the Burren. As a result of research in

recent decades, past soil erosion, once largely speculative, is

now underpinned by a considerable body of evidence. This

includes soil remnants sealed by archaeological structures

(Drew 1982, 1983; Plunkett Dillon, 1985; Lynch 1994; Jones

2004), colluvial deposits in grykes (Drew 1983) and karren fea-

tures, preserved on limestone slaps used in megalithic-tomb

construction, that suggest an acidic soil cover in nowadays

soil-free areas (Plunkett Dillon 1983). Drew (1982) reported

preliminary results from U ⁄Th dating of speleothems from

Burren caves that suggested cessation of calcite deposition in

the 2nd millennium BC as a result of loss of soil cover. Mean-

while, a speleothem from Polnagollum cave, south-western

Burren, has yielded an end-of-deposition date of

3230 ± 50 cal. BP (F. McDermott, pers. comm.). This points

to soil cover on the overlying bedrock until at least the mid to

late BronzeAge.

The present investigations have demonstrated the presence

of minerogenic soil in shallow peat-covered depressions on the

Cappanawalla upland and under shallow peat at Sliabh Rua.

In the latter Knockanes–Sliabh Rua area, mass soil movement

has been documented and buried charcoal has yielded the

AMS 14C date 3070 ± 35 BP, which suggests substantial soil

erosion in the late Bronze Age (Moles et al. 1999; Moles &

Moles 2002).

The data presented here indicate that soil erosion may

not be confined to the Neolithic and Bronze Age as is

sometimes postulated (e.g. Gibson 2007), but continued to

be of significance well into the mediaeval period and proba-

bly beyond. Erosion has probably characterized Burren soils

since the ice sheets retreated (c. 12 000 BC), with intense

erosion taking place at particular times, e.g. during climatic

downturns (cf. Younger Dryas; c. 10.6–9.5 BC) or during

phases of woodland clearance associated with intensive

farming.

It is puzzling, however, that lake sediments have failed to

yield conclusive evidence for soil erosion (cf. Watts 1984;

Schettler et al. 2006), even allowing for the possibility that

eroding soils may not readily reach lake basins where karst-

type, sub-surface hydrology prevails. Sediments from Molly’s

Lough, however, show elevated magnetic susceptibility values

that have been interpreted as indicative of sustained erosion

during the Neolithic and Bronze Age (c. 6000–3000 cal. BP;

Lamb&Thompson 2005). The lake, however, may not be typ-

ical for the Burren, lying, as it does, near the karst–shale geo-

logical divide.

While it is clear that soil erosion on a significant scale has

occurred in the Burren since the last glaciation and that the link

with human activity is strong (but cf. Wilson et al. (2008) for a

contrasting soil-erosion history from limestone terrain in

north-west England), details regarding extent, timing and

quantification of erosion, and its consequences for vegetation

dynamics, flora history and past human settlement patterns

have still to be elaborated.

FRESH INSIGHTS INTO THE DEVELOPMENT AND

ORIGIN OF THE UPLAND BURREN FLORA

The late Holocene history of key upland Burren species and

plant communities is considered in the light of the newly avail-

able evidence (see also Interpretation of data).

Dryas was not recorded in LLM II, there is only a single

record in GTC7 (33 cm) while, in CPW1, a continuous

curve is initiated at the base of zone 3 (c. AD 1100), i.e.

the record begins as peat accumulation starts. At CPW1,

the decrease in hazel and the beginning of peat accumula-

tion clearly provided locally favourable conditions for

expansion of Dryas in the mediaeval period. Failure to

record pollen of Dryas prior to this cannot be regarded as

proof of absence. The species appears to be so under-repre-

sented in pollen diagrams as to require local and consider-

able presence before it registers in pollen records (cf. Jelicic

& O’Connell 1992).

Records for Gentiana (cf. verna) are extremely scarce in all

profiles (CPW1-4, 16 cm: two pollen; and LLM II-1, 168 cm;

c. 400 BC: a single pollen). This is presumably due to poor pol-

len production and dispersal rather than rarity so that the

records, though sparse, are of some significance.

As regards Arctostaphylos uva-ursi, there is only a single

record (zone CPW1-2, 25 cm; c. AD 1500), prior to the

decline in Corylus on Cappanawalla. Arctostaphylos uva-ursi

seems to be very much under-represented and so no firm

conclusions regarding presence or absence in the past can be

drawn. Absence, however, is unlikely given the species’

frequency today in the north-west Burren uplands. The

CPW1 profile shows that other heath species such as Erica

cinerea (seed also recorded) and Empetrum nigrum were

important for at least the last two millennia at Cappanawalla

but are much less well represented at Gortaclare (especially

E. nigrum), which again corresponds to the present-day

situation. Interestingly, E. nigrum is abundant on nearby

Turlough Hill which is at similar elevation, but more compa-

rable to the north-west Burren uplands in being near the

coast.

The importance of Plantago maritima at Cappanawalla and

Lios Lairthın Mor, and rather poor representation in GTC7

are noteworthy. Plantago maritima is, however, frequent in the

eastern Burren and is locally abundant in limestone grasslands

on the lower slopes of Gortaclare, but it was not noted at or

near the sampling site, GTC7, which serves to emphasize the

local character of the pollen profiles.

Inter-site differences, as elaborated above, suggest that fac-

tors other than grazing and overall levels of human impact



have a major influence on distribution and composition of the

upland plant communities. Differences in wind exposure, sub-

strate (e.g. shallow minerogenic soils bearing heath on Cap-

panawalla; chert locally extensive on Gortaclare), humidity

levels and atmospheric nutrient input as determined by dis-

tance from the ocean probably all play an important role. To

what degree these are plagioclimax communities, especially the

arctic-alpine-like heaths as on Cappanawalla, remains to be

demonstrated.

The survival of arctic-alpine species in an eu-oceanic region

during the post-glacial, when woodlands largely dominated, is

another intriguing aspect of the Burren flora. The evidence pre-

sented here for open pine woodland with hazel on the Cappan-

awalla uplands until relatively late in the Holocene is

noteworthy, but not altogether surprising, given the role

played by pine during much of the Holocene in the Burren and

Aran Islands (Watts 1984; Molloy & O’Connell 2004; Feeser

2009). A modern-day analogue of these former plant commu-

nities is the open pine woodlands common on shallow alkaline

soils in Scandinavia (cf. Dierssen 1996; Kelly & Connolly

2000). TheMelico nutantis–Pinetumwoodlands on calcareous

rendzinas in southern Scandinavia include not only Sesleria

caerulea, Galium boreale, Geranium sanguineum and Campan-

ula rotundifolia,but alsoheathy species suchasCalluna vulgaris,

Arctostaphylos uva-ursi, Succisa pratensis and Potentilla erecta

(cf. Dierssen 1996). From mid Norway northwards, Dryas

octopetala is an important component in open birch–pine

woodlands on shallow alkaline rendzinas (Epipacto

atrorubentis-Betuletum). Given these striking floristic and

environmental similarities, we suggest that open Pinus

sylvestris-dominated woodland provided an important and

extensive habitat for the arctic-alpine flora of the Burren dur-

ing prolonged periods of more or less full woodland cover.

This would also help explain why some 39 species of putative

ectomycorrhizal fungi, normally intimately associated with

pine woodland, are confined in the Burren to these upland arc-

tic-alpine plant communities (Harrington & Mitchell 2005;

also Harrington 2003). Both the understorey and the charac-

teristic ectomycorrhizal flora of these open pine woodlands

have presumably survived, in situ, the demise of pine.

Acknowledgements

A. Bird and N. Higgins assisted with fieldwork, and A. Bird carried out pre-

liminary analyses on CPW1. J. van Leeuwen and M. Richardson advised on

identification of coprophilous fungal spores. F. McDermot provided informa-

tion on unpublished speleothem analyses. The paper has benefited from helpful

comments by referees (two anonymous and M.-J. Gaillard). The research was

financially supported by an EMBARK Initiative Postgraduate Research

Fellowship (IRCSET) to I.F.

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Handling Editor: Richard Bradshaw



Documents

Fresh insights into long-term changes in flora, vegetation, land use and soil erosion in the karstic environment of the Burren, western Ireland