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Journal of
Mire development across the fen–bog transition on
Blackwell Publishing Ltd.

Ecology 2003
91, 253 –264 the Teifi floodplain at Tregaron Bog, Ceredigion,
Wales, and a comparison with 13 other raised bogs
P. D. M. HUGHES and K. E. BARBER*
Palaeoecology Laboratory (PLUS), Department of Geography, University of Southampton, Highfield,
Southampton, SO17 1BJ

Summary
1 A literature comparison of 14 radiocarbon-dated macrofossil records of raised peat
bog initiation indicates that there is a relationship between the prevailing climate and
the character of the first ombrotrophic vegetation communities at peatland sites in Britain
and Ireland.
2 All that is required for ombrotrophy is the separation of the mire growing surface
from the influence of surface and subsurface waters. This could occur via vertical
accumulation of the peat mass or a lowering of the water table. The establishment of bog
species can be rapid once isolation occurs.
3 Peatlands may become ombrotrophic in a variety of water table conditions and
climatic regimes. There are at least two distinctive routes to ombrotrophy, via a ‘dry-
pioneer oligotrophic community’ or via a ‘wet-pioneer oligotrophic community’.
4 Tregaron South-east Bog does not fit the pattern suggested by the literature com-
parison. The Fen–Bog Transition (FBT) occurred in a period of increased effective
precipitation but the first ombrotrophic community was indicative of relatively dry,
‘hummocky’ bog and a deep or unstable water table.
5 The transitional poor fen communities at Tregaron South-east Bog were short-lived.
Sphagnum palustre mire lasted for 90 years compared to 300 years at Bolton Fell Moss
in Cumbria, and the FBT was synchronous across much of the bog.
6 The Tregaron peatland complex has a long history of water table fluctuations as
shown by the stratigraphy of the marginal peats.
7 Channel incision in the River Teifi could have contributed to the development of ‘dry
hummock’ pioneer bog in the humid climatic conditions at 7300 cal. BP, by reducing
the level of lagg streams.
Key-words: mire development, fen–bog transition, Holocene, hydroseral succession,
plant macrofossils, peat stratigraphy.
Journal of Ecology (2003) 91, 253–264

raised bogs have either been severely cut, re-exposing

Introduction
Although the Fen–Bog Transition (FBT) is one of the
key landscape changes of the Holocene it has been the
subject of little recent research. An understanding of
the mechanisms involved in this transition is required
for long-term conservation management, since many
Correspondence: Palaeoecology Laboratory (PLUS), Depart-
ment of Geography, University of Southampton, Highfield,
Southampton, SO17 1BJ (tel. +44 23 80592489; fax +44
© 2003 British 23 80593295; e-mail paul.hughes@soton.ac.uk).*E-mail:
Ecological Society keith.barber@soton.ac.uk
the underlying fen peats, or have been set back to an
early successional stage by changes in the local water
table. Conventional accounts of the development of
raised bogs from swamp, fen and fen carr communities
stress the role of autogenic processes (Walker 1970;
Bellamy 1972) at the FBT. The formation of the domed
bog surface is usually explained as a gradual process of
peat accumulation above the level of the regional water
table. Many, but not all, bog successions pass through
a fen carr stage, resulting in a substantial thickness of
wood peat. Acidification of the accumulating peat sur-
face is a consequence of leaching by rainwater, followed

254
P. D. Hughes
& K. E. Barber
Fig. 1 Location maps of Tregaron Bog, Ceredigion, Wales, showing the position of core TSE94 on the South-east Bog.
by the action of Sphagnum. The increasing elevation of
the growing surface would lead to the progressive
exclusion of ground and surface waters and eventually
to sole supply by base-poor rain water.
The autogenic model predicts that the water table
will rise gradually with peat accumulation, sustained
by a dynamic equilibrium between supply and discharge.
This suggests that the water table bog will remain close
to the surface throughout the fen–bog transition.
Damman (1986) has suggested that the surface of pioneer
raised bog would be wet, because the height of the
raised surface would be below the maximum possible
under the prevailing climatic conditions. This situation
is observed at the FBT in Abbeyknockmoy Bog near
Galway (Hughes 1997), where pool and mud-bottom
species dominate the first oligotrophic community
after the transition. Incipient raised bog patches in
modern fens also demonstrate that oligotrophic bog
mosses can survive just a few centimetres above the level
of mineral water influence, suggesting that little isolation
is required for oligotrophic bog formation to begin.
Modern observations of incipient raised bog condi-
tions are not necessarily a good guide to past mechan-
isms of bog formation because boundary conditions
have changed significantly over the duration of the
Holocene. Plant macrofossil investigations of raised
bogs in northern England (Hall et al. 1995; Hughes
1997; Leah et al. 1998) have shown that some bogs
forming over kettlehole fens (e.g. Walton Moss:
Hughes et al. 2000), have stratigraphies that differ
© 2003 British
significantly from that predicted by the autogenic
Ecological Society, model of hydroseral succession (sensu Walker 1970 and
Journal of Ecology, Damman 1986), and that ombrotrophic conditions were
91, 253–264 reached very early in the Holocene, sometime before
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9000 cal. BP. Many of the sites that developed from
thin fen peats record phases of highly humified Erio-
phorum vaginatum – Calluna peat, containing charcoal
and Cenococcum fruit bodies, after the FBT. This sug-
gests that the kettlehole mires became ombrotrophic
under conditions that promoted deep or fluctuat-
ing water tables. Although mire stratigraphies similar
to those in the kettlehole mires of northern England
have been reported by a number of researchers (e.g.
Casparie 1972; Svensson 1988; Leah et al. 1998), fur-
ther research is required to evaluate the importance of
lowered or fluctuating water tables at the FBT.
The Tregaron Bog (Cors Caron) peatland complex
in the Teifi valley (Fig. 1) is one of the largest near-
natural mires remaining in Great Britain, and offers an
opportunity to test ideas concerning the FBT in a peat-
land that has developed from deep lake and reedswamp
conditions in a floodplain setting. It contrasts greatly
with raised mires formed over thin kettlehole fen peats.
Although Walker (1970) used Tregaron Bog as a prime
example of an undisturbed autogenic sequence, there is
evidence to suggest that the South-east Bog in particu-
lar has a long history of fluctuating water levels. Godwin
& Mitchell (1938) first observed these changes in a
sequence of intercalated brushwood and fen peats from
the eastern margin of the peat complex (see Fig. 2).
This paper aims to examine the record of early
Holocene peatland development on the Teifi flood-
plain at Tregaron Bog, and to compare the timing and
character of the FBT with other radiocarbon-dated
macrofossil records of raised bog initiation from Britain
and Ireland. Specifically the paper will study the rela-
tionship between the character of the FBT and factors
affecting local water table dynamics.

255
Mire development
across the fen–bog
transition
Fig. 2 Lithostratigraphy of Tregaron South-east Bog showing (i) detail of the eastern margin, (ii) a typical central section and (iii)
the stratigraphy at the contact with the Teifi river channel. (After Godwin & Mitchell 1938). Open circles, till; wavy lines,
Sphagnum; vertical lines, Phragmites peat; cross-hatching, alluvium; V, wood peat; fine stipple, clay; coarse stipple, sands and silts.
 
Tregaron Bog NNR (Site code: TRG; NGR:
SN685622) is located 1.5 km north of Tregaron village
on the floodplain of the River Teifi (Fig. 1), covering an
area of 792 ha. The complex is composed of three
active raised mire domes, the West Bog, South-east Bog
and North-east Bog. Two further mires in the north
and south-western sectors of the system have been
destroyed by extraction for fuel and agricultural reclama-
tion, although significant quantities of early Holo-
cene peat probably survive under fringing fields. The
principal channel of the Teifi winds through the
middle of the peatlands, separating the West Bog from
the South-east and North-east Bogs. The anastomosing
character of the watercourse and the close associ-
ation of the river with the extensive floodplain wetlands
afford a rare opportunity to study a well-preserved,
near-natural floodplain environment. The wetland
communities of Tregaron Bog NNR consequently have
considerable conservation value.
Old Ordnance Survey maps show that the three
main raised domes were largely undisturbed prior to
1906. Much of the perimeter of the South-east Bog
was cut by hand during the first half of the twentieth
century, before the purchase of the site by the then
Nature Conservancy (now Countryside Council for
Wales (CCW)). Willow scrub has colonized the
easternmost margin of the South-east Bog adjacent to
the dismantled railway line. This area was regularly
managed to keep the line clear until its closure in the
1950s. Much of the central dome of the South-east Bog
supports Sphagnum-rich raised mire vegetation
described as M17 and M18 in the National Vegetation
Classification (NVC, Rodwell 1991). The dominant
Sphagna on the mires are Sphagnum papillosum,
© 2003 British
Ecological Society, Sphagnum magellanicum and Sphagnum capillifolium
Journal of Ecology, var. rubellum. Associated species include Sphagnum
91, 253–264 pulchrum.
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   
A peat core designated TSE94 was sampled from the
centre of the South-east Bog (see Fig. 1) after studying
the lithostratigraphic sections of Godwin & Mitchell
(1938). Samples were recovered using a wide-bore
(9 cm × 30 cm) modified Russian pattern corer (Barber
1984) and described using the Tröels-Smith sediment
classification (Tröels-Smith 1955). Peat samples were
taken alternately from two adjacent boreholes to
ensure that they were not disturbed by the nose cone of
the corer and sampling depths were overlapped by 5 cm
to provide a continuous record. Samples were stored at
4 °C to prevent fungal growth.
 
The surface layer of each core section was removed
before sampling to reduce contamination. Samples
with a volume of 4 cm3 were measured by displacement
and sieved using a 125-µm mesh and 5 L of water. Macro-
fossil remains were routinely examined at × 40 − × 150
magnification but identifications of Sphagnum branch
leaves and monocotyledon epidermal tissues were
made at × 200 − × 400 magnification. Vegetative plant
macrofossils were quantified using the quadrat and
leaf count (QLC) technique (Barber et al. 1994). Fruits,
seeds and other small macrofossils were assessed
using a five-point scale of abundance (Walker & Walker
1961; Barber 1981; 10 = rare, 20 = occasional, 30 =
frequent, 40 = very frequent, 50 = abundant). Nomen-
clature follows Stace (1991) for vascular plants, Daniels
& Eddy (1990) for Sphagna & Smith (1978) for brown
mosses.
  
Pollen samples with a volume of 1 cm3 were prepared
using standard procedures (Barber 1976; Moore et al.

256 1991). Samples were treated twice with potassium
P. D. Hughes hydroxide and washed through a 180-µm sieve. A 10-
& K. E. Barber µm sieve was used to remove the clay fraction prior to
treatment with acetylation mixture and staining with
alcohol-soluble safranin. The concentrated pollen was
mounted in 2000 cs density silicon oil and counted
using a pollen sum of 350 non-mire pollen (NMP), on
a light-transmitting microscope. Routine identification
was undertaken at × 400 magnification, whilst problem
grains were examined at × 600 or under oil immersion
at × 1000. Identifications were made using the South-
ampton Palaeoecology Laboratory (PLUS) pollen
type collection and the keys of Moore & Webb (1978),
Faegri et al. 1989) and Moore et al. (1991). Pollen type
nomenclature follows Bennett (1994).
  
Samples for conventional radiometric radiocarbon
dating, with a wet mass of 100 g, were analysed at the
NERC Radiocarbon Laboratory at East Kilbride. In
the ombrotrophic strata Sphagnum was sampled for
dating where possible and every effort was made to
date identifiable above-ground wood samples in the
Phragmites-rich fen peats. Obvious rootlets were
extracted from the samples prior to submission. Radio-
carbon dates were calibrated using INTCAL98 (Stuiver
et al. 1998) and the results are presented in Table 1.
Radiocarbon dates quoted in the text are presented
as the full 2 σ calibrated range before present (cal. BP).
Both calibrated and uncalibrated dates are quoted in
Table 1. Interpolated age estimates based upon the
radiocarbon chronology are denoted with the prefix
‘circa’ (c).
Results
  
Tregaron South-east Bog: core TSE94,
850 cm – 300 cm depth
The results of the macrofossil analyses for core TSE94
below 300 cm depth are shown in Fig. 3. Within the
macrofossil diagrams, unlinked histograms represent
data quantified using the five-point scale of abundance
technique, whereas linked histograms represent QLC
data. As far as possible, the order of appearance of spe-
cies in the diagram is sequential, with fen species pre-
sented to the left. Local macrofossil zones have been
Table 1 Radiocarbon dates from core TSE94
Depth below peat surface
Laboratory no. Site code Material (cm), in core TSE94
© 2003 British
SRR-5642 TSE94 Ombrotrophic peat 338 –346
Ecological Society,
SRR-5643 TSE94 Ombrotrophic peat 536 – 544
Journal of Ecology,
SRR-5644 TSE94 Fen carr peat 562 –570
91, 253–264
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defined with the aid of a stratigraphically constrained
cluster analysis with no data transformation, using
CONISS within TILIA version 1.12. The resultant
dendrograms have been omitted from the macrofossil
diagrams to maintain clarity. The local macrofossil
assemblage zones have been labelled using the follow-
ing system: TSE (Tregaron South-east Bog), M (local
macrofossil zone (lmz)), -zone letter(s). Upper case
letters denote main zones. Lower case letters denote
subzones.
    
The lowermost zones of the sediment sequence from
core TSE94 record a classic lake infilling sequence
beginning with weakly laminated lake clays containing
Chara oospores and Myriophyllum spicatum, through a
protracted reedswamp phase, composed principally of
Phragmites australis and Cladium mariscus, to a swamp
carr stage. In zone TSE94M-Da the reed peat contains
a significant proportion of woody roots, which were
probably deposited at this level as a result of root pene-
tration from the overlying Alnus glutinosa-dominated
zone, TSE94-Db.
   
In the latter part of TSE94M-Db the mire began to
develop a tussocky surface, as indicated by the appear-
ance of Carex paniculata. A major shift in the vegeta-
tion occurred during in Zone TSE94M-F, and by the
end an open poor fen containing Sphagnum palustre,
Aulacomnium palustre and a small amount of Eriophorum
vaginatum had replaced the swamp carr woodland. This
poor fen phase only deposited approximately 8 cm of
peat before a strongly humified peat type containing
the remains of oligotrophic bog vegetation (Zone
TSE94M-Ga) replaced it.
   
( ‘     ’)
A community indicative of deep mire water tables and
aerated surface conditions dominated the pioneer
raised mire in zone TSE94M-Ga. Eriophorum vagina-
tum, Calluna vulgaris and the soil fungus Cenococcum
were the main species recorded in this phase. Increased
levels of macroscopic charcoal indicate frequent local
burning of the mire surface throughout the zone. Godwin
& Mitchell (1938) identified this lower ombrotrophic
Calibrated range BP
14
C date BP (2 σ) (uncalibrated)
4035 ± 45 4805 – 4410
6480 ± 45 7400 –7240
6775 ± 50 7650 –7480
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257
Unlinked histograms denote abundance data quantified on a 5-point scale from rare = 10 to abundant = 50 (see text for full scale). Linked histograms denote data collected using the quadrat and leaf count method
Fig. 3 Macrofossil diagram of core TSE94, 300 – 850 cm, Cors Caron south east bog (Tregaron Bog), Ceredigion, Wales. Taxa arranged with rheotrophic species on the left and oligotrophic species on the right.

Mire development
across the fen–bog
transition
of Barber et al. (1994).
© 2003 British
Ecological Society,
Journal of Ecology,
91, 253–264
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258 Table 2 Summary of the pollen stratigraphy of core TSE94 presented in Fig. 4
P. D. Hughes
& K. E. Barber Local pollen zone Description

TSE94P-C Sphagnum spores are frequent, whilst Dryopteris-type declines to trace levels. The diversity of herb
430–550 cm species is much lower than zone TSE94P-B. Frequencies of Corylus-type and Alnus increase throughout
Betula/Quercus/Alnus the zone at the expense of other tree species.
TSE94P-B The pollen curves for most arboreal species remain stable from TSE94P-A to TSE94P-B but Alnus
550–580 cm displays a sustained rise from trace levels at 580 cm. TSE94P-B marks a significant increase in
Betula/Pinus/ Cyperaceae pollen accompanied by Dryopteris-type spores. By contrast Poaceae pollen declines from
Quercus/Alnus 40% to 10% NMP. These change coincide with the establishment of Carex paniculata in the macrofossil
record in zones TSE94M-D and TSE94M-E. The diversity of fen herbs increases significantly whilst the
upper zone boundary with TSE94P-C is defined by the rise of Sphagnum spores to 60% of NMP.
TSE94P-A The principal tree species in pollen zone TSE94P-A, Betula, Pinus and Quercus, display stable frequencies
580–670 cm throughout the zone. At 638 cm Alnus registers the first significant increase to c. 10% NMP. This increase
Betula/Pinus/Quercus coincides with the appearance of Alnus glutinosa macrofossils at the sample site in macrofossil zone
TSE94M-D. Poaceae pollen represents between 30 and 40% of NMP. Cyperaceae remains stable
throughout the zone at 15% of NMP. The zone also registers significant levels of Osmunda regalis and
Dryopteris-type fern spores. Acidophilous species such as the members of the Ericaceae occur at trace
levels within the zone whilst records of Sphagnum spores remain below 10% of the pollen sum.

horizon as a humified Sphagnum peat in their field-
based study of the mire’s stratigraphy, but the macro-
fossil diagram (Fig. 3) clearly shows that Sphagnum
is only an intermittent component of the early raised
mire peats. Barber et al. (1998) reported similar problems
with the identification of humified oligotrophic
peats at Bolton Fell Moss.
    -
  
The upper two zones of Fig. 3 (TSE94M-H and -I)
register the switch to dominance by Rhynchospora alba
and subsequently Sphagnum imbricatum cf. ssp. austini
(hereafter referred to as Sphagnum imbricatum). These
changes probably represent evidence for stabilization
of the mire water table at or near the peat surface, since
Rhynchospora alba is a species indicative of pool edges,
low lawns and mud-bottomed areas subject to regular
inundation. The wetter surface conditions are also
shown by the disappearance of Cenococcum fruit
bodies and the increase in Erica tetralix. Field strati-
graphy cores, sampled at the same time as the main macro-
fossil core, demonstrate that these changes occurred
over most of the mire. The upper 3 m of peat have not
been analysed in detail in the research presented here.
   
Tregaron South-east Bog: core TSE94,
850 cm – 300 cm depth
The results of the pollen study are presented in Fig. 4.
An unconstrained cluster analysis routine was used to
objectively define local pollen assemblage zones (lpz),
© 2003 British
using the CONISS option within the computer pro-
Ecological Society, gramme TILIA v1.12; the resultant dendrograms have
Journal of Ecology, been omitted from the pollen diagram to aid clarity. The
91, 253–264 local pollen assemblages zones have been labelled using
the following system: TSE94 (Tregaron South-east
Bog, cored 1994), P (local pollen zone), -zone letter.
The pollen stratigraphy is summarized in Table 2.
Discussion
 ‒      

Analyses of early and mid-Holocene raised bog depos-
its from Britain and Ireland show at least two distinc-
tive pathways across the fen–bog transition. At
Abbeyknockmoy Bog, near Galway in Ireland, the first
oligotrophic community began forming over fen peat
at 8200 cal. BP, in wet lawn and mud bottom condi-
tions (Hughes 1997). This stratum may be traced for
some considerable distance in the cut peat face that
dissects Abbeyknockmoy Bog (unpublished results),
and represents a mire-wide switch. The timing of the
FBT here is significant because it coincides with a
widespread and well-known climatic cooling event,
recorded in Greenland ice accumulation records
(O’Brien et al. 1995; Alley et al. 1997) and many other
palaeoclimate proxies, such as the oxygen isotope sig-
nal from lake sediments in Germany (Von Grafenstein
et al. 1998). Other FBTs with a ‘wet character’ occur
during subsequent phases of high effective precipita-
tion in the early to mid-Holocene (see Table 3). For
example, at Kirkpatrick Fleming Bog oligotrophic
conditions established at 7700 cal. BP with the
development of Sphagnum-filled pools (Tipping 1995).
Contemporaneous palaeoclimatic proxies – effective
precipitation records from peat bogs (Hughes et al.
2000), Swedish lake varve records (Snowball et al.
2002), and lake levels in the Jura Mountains of France
(Magny 1992) – all point to a major and prolonged
increase in effective precipitation commencing at
7800 cal. BP. At Fenton Cottage Bog, Lancashire
(Wells et al. 1997) the FBT occurs at 4300 cal. BP (tephra
13652745, 2003, 2, Downloaded from https://besjournals.onlinelibrary.wiley.com/doi/10.1046/j.1365-2745.2003.00762.x by Readcube (Labtiva Inc.), Wiley Online Library on [06/02/2023]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License
259
Fig. 4 Pollen diagram of core TSE94, Cors Caron south east bog (Tregaron Bog), Ceredigion, Wales, showing selected taxa for zones covering the fen–bog transition (430 – 670 cm depth). The pollen sum is based

Mire development
across the fen–bog
transition
on a count of 350 non-mire pollen grains (see text).
© 2003 British
Ecological Society,
Journal of Ecology,
91, 253–264
 13652745, 2003, 2, Downloaded from https://besjournals.onlinelibrary.wiley.com/doi/10.1046/j.1365-2745.2003.00762.x by Readcube (Labtiva Inc.), Wiley Online Library on [06/02/2023]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License
260 3 Summary of radiocarbon dated fen –bog transitions from Britain and Ireland
Table
P. D. Hughes
Raised bog initiation, Water table position after FBT Prevailing effective
& K. E. Barber
Authorship Cal. years BP* (dominant species) precipitation level†

Walton Moss, Cumbria Hughes et al. (2000) c. 9850 Deep (Eriophorum-Cenococcum) Low
Glasson Moss, Cumbria Hughes (1997) 10 040 –9910 (2 σ range) Deep (Eriophorum-Empetrum) Low
Bolton Fell Moss, Cumbria Hughes (1997) c. 9600 Deep (Eriophorum-Calluna) Low
Solway Moss, Cumbria Hughes (1997) c. 8840 Intermediate (Sphagnum sect. Low
Acutifolia)
Mongan Bog, Co. Hughes (1997) 5575 – 5315 (2 σ range) Deep (Pinus-Eriophorum) Low
Offaly, Eire
Borth Bog, Wales Hughes & Schulz (2001) 6280 – 5990 (2 σ range) Deep (Pinus-Eriophorum) Intermediate
Abbeyknockmoy Bog, Eire Hughes (1997) 8320 – 8020 (2 σ range) Near surface (Rhynchospora) High
Walkers Heath, Cheshire Leah et al. (1998) c. 8160 Surface (Sphagnum s. Cuspidata) High
Rawcliffe Moss A, Middleton et al. (1995) 8038 –7586 (2 σ range) Near surface (Eriophorum, S. s. High
North Lancashire Acutifolia and S. papillosum)
Kirkpatrick Fleming Bog, Tipping (1995) c. 7700 Surface (Aquatic local pollen types High
Scotland including Hydrocotyle vulgaris)
Sluggan Bog, Northern Ireland Smith & Goddard (1991) c. 7550 Near surface High
(Sphagnum imbricatum-Eriophorum)
Goldcliff, Gwent Levels, Wales Smith & Morgan (1989) c. 5750 Near Surface (Sphagnum) High
Tregaron Bog, Wales This paper 7400 –7240 (2 σ range) Deep (Eriophorum-Cenococcum) High
Fenton Cottage, Lancashire Wells et al. (1997) 4300 (tephra) Surface (Scheuchzeria palustris) High

*c. indicates that the age estimate is calculated by linear interpolation between two radiocarbon dates.
†Effective precipitation information is based on peat bog records from Barber et al. (1994) and Hughes et al.
(2000) (see Fig. 5) and the wider literature.

isochron), when the mire switched from fen to a wet
oligotrophic community dominated by Scheuchzeria
palustris, which suggests that the bog surface became
flooded with acidic water. This time period is well rep-
resented in the peat-based proxy palaeoclimatic record
as the start-date of another major increase in effective
precipitation (e.g. Barber et al. 1994; Hughes et al. 2000).
The transitions mentioned above accord with the
‘wet-mode’ of raised bog formation described by
Walker (1970) and Damman (1986), where the mire
water table evidently remains close to the ground sur-
face throughout the FBT and in the pioneer oligo-
trophic bog stage. The transition to ombrotrophy is
thus achieved through the rapid accumulation of peat
in a moist climate.
Stratigraphic records of raised bog development
from Cumbria and central Ireland show that there is
another route to ombrotrophy, via a dry hummocky
oligotrophic community. For example, at Solway Moss
(Hughes 1997), the pioneer oligotrophic plant macro-
fossil assemblage was dominated by Eriophorum vagi-
natum, Calluna, fungal sclerotia, Polytrichum commune
and later Sphagnum sect. Acutifolia. Hughes (1997)
and Hughes et al. (2000) have recorded similar strati-
graphies at Mongan Bog in Ireland and three other
sites in Cumbria, including Walton Moss. In many
cases the pioneer raised bogs developed during phases
of reduced effective precipitation (Table 3; Fig. 5). The
‘dry FBT’ is clearly possible only when effective pre-
© 2003 British
cipitation is marginal for bog development, resulting in
Ecological Society, a lowland mire community similar to the modern the
Journal of Ecology, ‘dry’ variant of ‘Pseudohochmoor’ found in central
91, 253–264 Europe and recognized by Rybnicek (1973). Other
raised bogs in Europe also display evidence for un-
stable or low water levels immediately after the FBT. For
example, Svensson (1988) noted that Store Mosse in
Sweden displayed a ‘dry’ stratigraphy at the transition
to raised peat. The summary of FBTs presented in
Table 3 suggests that in many mire sequences there is a
relationship between the prevailing level of effective
precipitation and the character of the pioneer raised
bog community. However, some FBTs are clearly influ-
enced by other factors that are not directly connected
to effective precipitation levels. For example, Casparie
(1972, 1993) uncovered evidence of desiccation cracks
in fen peat lying immediately below the contact with
highly humified ombrotrophic peat at Bourtanger
Moor in the Netherlands. He concluded that the fen
dried out and became oligotrophic as a consequence of
river capture. Kulczynski (1949) also noted that ombro-
trophic peat bogs developed in areas of the Pripet
marshes where river capture had deprived the peat
body of surface and subsurface water supplies.
     
At Tregaron South-east Bog there is no clear relation-
ship between the pioneer bog assemblage and prevail-
ing effective precipitation levels. Table 3 shows that the
bog responded in a different way when compared to the
other radiocarbon-dated macrofossil records avail-
able for sites in Britain and Ireland at this time (c.
7300 cal. BP). Whilst effective precipitation levels after
the FBT at 7300 cal. BP were relatively high, the first
oligotrophic community at Tregaron was hummocky
and indicative of deep or fluctuating water tables.

261
Mire development
across the fen–bog
transition
Fig. 5 Comparison of the timings of the Fen–Bog Transition (FBT) at (A) Abbeyknockmoy Bog (AKM95) (B) Mongan Bog
(MOG95) (C) Tregaron Bog (D) Glasson Moss (GLM95), and (E) Bolton Fell Moss (BFML), with the full Holocene proxy-
effective precipitation record from Walton Moss (Hughes et al. 2000). Graphs A-E represent a record of change in the fen–bog
gradient against calibrated radiocarbon years BP. The reconstructions are based on Detrended Correspondence Analysis (DCA)
sample scores × 100.
The original stratigraphic survey of Tregaron South-
east Bog, completed by Godwin & Mitchell (1938),
includes a detailed analysis of the deposits along the
eastern hill margin of the mire. The results, reproduced
in Fig. 2, record a sequence of three brushwood peat
stages intercalated with two bands of Phragmites peat.
Godwin & Mitchell (1938) interpreted this alternating
stratigraphy as evidence of early Holocene fluctuations
in the mire water table. Strata showing reedswamp
peats extending out onto the mineral soil at the edge of
the mire may represent high water table levels; con-
versely, the extension of brushwood peat out onto the
mire surface probably indicates periods when the water
levels fell. These fluctuations may be linked to the
dynamics of the River Teifi, since there is evidence of
several phases of channel incision in the stratigraphic
cross section of the South-east Bog. During a number
of undated periods in the Holocene, the river cut
through part of the peat sequence before incising into
the underlying Devensian lake deposits. (see river mar-
gin section in Fig. 2) The channel has since re-filled
with silts and clays that form the modern bed of the
Teifi.
The characteristics of the marginal peat stratigraphy
at the point of the FBT are of particular interest to the
present discussion. Figure 2 shows that the uppermost
© 2003 British
(75 cm thick) deposit of brushwood, extending at least
Ecological Society, 60 m out onto the edge of Tregaron South-east Bog,
Journal of Ecology, was recorded by Godwin and Mitchell as immediately
91, 253–264 overlain by ombrotrophic peat. Supplementary coring
13652745, 2003, 2, Downloaded from https://besjournals.onlinelibrary.wiley.com/doi/10.1046/j.1365-2745.2003.00762.x by Readcube (Labtiva Inc.), Wiley Online Library on [06/02/2023]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License
of the bog margin during the present study found
similar results in 12 more boreholes. This stratigraphy sug-
gests that the FBT occurred during a period when mire
water levels were relatively low.
Detailed analyses of plant macrofossils provide a
clearer picture of the sequence of events leading up to
the FBT. Figure 3 shows that the switch from swamp-
carr peat to dry Eriophorum vaginatum-rich peat
occurred via a short-lived mesotrophic fen phase dom-
inated by Sphagnum palustre. Two radiocarbon dates
placed just above (SRR-5643) and below (SRR-5644)
the FBT (see Table 1) show that the transitional fen
community could have lasted as little as c. 90 years.
This represents a relatively rapid rate of succession by
comparison with equivalent stages in other raised
mires. For example, Bolton Fell Moss passed through a
Sphagnum palustre-dominated transitional phase
that lasted c. 300 years prior to the emplacement of
Eriophorum vaginatum/Calluna mire and the average
lifespan of the Sphagnum palustre transitional stage at three
other mires in Cumbria was 290 years (Hughes 1997).
Not only was the FBT relatively rapid at Tregaron
South-east Bog, it was also synchronous across large
parts of the mire. This is suggested by three lines of evid-
ence. First, radiocarbon dates sampled at the FBT
from marginal and central peat cores, by Hibbert &
Switsur (1976) and Hughes (1997), respectively, both
give the same age estimate of c. 7300 cal. BP. Second,
the macrofossil and pollen diagrams (Figs 3 and 4)
both show that the first major increase in acidophilous

262 taxa occurred at 550 cm depth. Any significant lag
P. D. Hughes between the establishment of raised mire elsewhere on
& K. E. Barber the bog and its arrival at the core site should be evident
in a comparison of the two records, because microfossils
are readily transported and represent an extra-local
signal, whereas vegetative macrofossils mostly record
the vegetation at the point of deposition. Finally, raised
bog deposits first appear in the stratigraphic sequence
at the same altitude across all but the very margins of
the bog cross-section. This stratigraphy contrasts with
other bogs that also became ombrotrophic in periods
of high effective precipitation. For example, a cross-
section of Abbeyknockmoy Bog shows that there is a
significant difference in the altitude and age of the FBT
in different parts of the mire (Hughes 1997). Abbey-
knockmoy bog first became ombrotrophic near its
present geographical centre at 8200 cal. BP, whereas
marginal areas acidified over 1000 years later.
The first oligotrophic deposits recorded in Tregaron
South-east Bog (zone TSE94M-Ga of the macrofossil
diagram, Fig. 3) demonstrate that the pioneer raised
mire had a relatively deep or fluctuating water table.
This contrasts with the stratigraphy predicted by
Damman (1986). The growing surface was frequently
aerated, as shown by the prominence of the soil fungus
Cenococcum (Ferdinandsen & Winge 1925) and the
increased frequency of macroscopic charcoal produced
by local fires. The dominant vascular plants on the mire
were Eriophorum vaginatum and Calluna vulgaris, both
species that thrive on bog hummocks. Eriophorum
vaginatum is particularly well adapted to fluctuating
water levels, becoming abundant under conditions of
spring flooding followed by desiccation of the mire sur-
face in mid- to late-summer and it is tolerant of burning
(Kummerow et al. 1988). Aulacomnium palustre and
Sphagnum section Acutifolia dominate the bryophyte
flora present in the assemblage. These taxa can also tol-
erate a deep water table. Furthermore, the assemblage
contains no species that require a near-surface, stable
water table.
All that is required for ombrotrophic conditions to
develop is the separation of the mire growing surface
from the supply of surface and subsurface waters, so
that meteoric supply becomes the sole source of water
and nutrients. The separation can occur as a conse-
quence of vertical accumulation of the peat mass, but
equally a lowering of the bog water table would also
promote isolation and possibly speed up the ‘forward’
rate of succession to ombrotrophy.
In a bog subject to a moist climate, but where there is
a history of water table instability, such as at Tregaron
Bog, the combination of high atmospheric humidity
and a lowered water table might be expected to pro-
mote a swift succession towards raised bog. This
hypothesis may explain why the character of the FBT at
© 2003 British
Tregaron differs from many others in Britain and Ire-
Ecological Society, land that became ombrotrophic in similar climatic con-
Journal of Ecology, ditions (see Table 3). A lowered water table could also
91, 253–264 account for the synchronous transition to oligotrophic
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communities across the entire mire. Studies of de-
watered fen and carr peats have shown that peat chem-
istry alters dramatically as the peat dries out. Kazda
(1995) found that levels of nitrogen, organic carbon,
calcium and magnesium decrease, and that pH values
also decline in response to lowered water levels. H+
ions are liberated as nitrogen compounds are mineral-
ized. Thus acidification of the peat surface could
have been promoted by the combined affects of fen
peat humification and a change to a meteoric water
supply.
There are a number of possible factors that could
account for the changes in water level recorded in the
minerotrophic stages of the South-east Bog that culmi-
nated in the FBT; these include climate change, river
channel adjustment and human impact upon drainage
basin vegetation. Variations in effective precipitation
could have directly driven the inferred changes in the
mire water table. This hypothesis predicts that the drop
in water table at the FBT would coincide with a phase
of reduced effective precipitation at c. 7300 cal. BP.
Independent climate archives from raised mires and
lakes (Magny 1992; van Geel et al. 1996; Hughes et al.
2000) record relatively high water levels at about this
time, suggesting that direct meteoric inputs to the mire
were not the main control leading to the rapid replace-
ment of fen by relatively dry oligotrophic bog.
Adjustments in the plan-form and channel depth of
the Teifi could have had a major impact upon the
hydrology of the early Holocene Tregaron peatlands.
During the fen phase the mire surface is flat and sup-
plied by surface and subsurface waters, whereas the
later ombrotrophic mire stages are supported on a
domed body of catotelmic peat with a low hydraulic
conductivity. The raised mire peats are therefore more
likely to be protected from changes in the river channel.
The peat stratigraphy adjacent to the main channel
records at least two phases when the river incised into
the lower peat deposits and the Lateglacial lake clays
(see Fig. 2). The Teifi and its local tributaries effectively
act as the lagg streams for the entire northern and west-
ern margins of the South-east Bog. Incision along the
length of this reach could have been sufficient to lower
the mire water table, whilst renewed silting of the chan-
nel or accumulation of woody debris would have had
the opposite effect. There are many possible controls
upon channel incision, however, and a phase of
increased effective precipitation could have been an
important one.
Human impacts upon the vegetation in the sur-
rounding catchment could also potentially alter the
hydrology of the drainage basin, leading to flashy run-
off, channel incision and a consequent drop in the
water table of floodplain mires. Chambers (1991) has
argued that the peat humification record at Tregaron
Bog may hold a land-use signal because of the inter-
linkages between slope, vegetation and drainage basin
hydrology. There is little evidence of human impact in
the pollen diagram spanning the FBT from the South-east

263 Bog (Fig. 4), though Mesolithic activities were often
Mire development concentrated in marginal environments – such as at
across the fen–bog the tree line (Simmons 1993). Buckley & Walker (2001)
transition studied the early history of peat development in an
upland area at Bryniau Pica near Tregaron, but they con-
cluded that ‘irrefutable evidence of Mesolithic peoples
in the uplands remained elusive’. Further research into
the upland land-use history of the Teifi drainage basin
may be required to establish whether Mesolithic cultures
had a widespread impact upon the Tregaron peatland
system.
Conclusions
The comparison of 14 radiocarbon-dated macrofossil
records of raised peat initiation at peatland sites in Brit-
ain and Ireland indicates that there is a relationship
between the prevailing climate and the character of the
first ombrotrophic vegetation communities. This sug-
gests that some raised bogs may record palaeoclimatic
conditions in their early stages of development.
The plant macrofossil reconstruction from Tregaron
South-east Bog, however, is an example of a record that
does not conform to the pattern suggested by the liter-
ature comparison. At Tregaron, the FBT occurred dur-
ing a phase of increased effective precipitation, but the
first oligotrophic bog community was indicative of a
deep or fluctuating water table. Here the FBT may have
been affected by the proximity of the Teifi river chan-
nel, which has shifted and incised on a number of occa-
sions during the early to mid-Holocene.
The Tregaron peatland complex has a history of
water table fluctuations throughout the reedswamp
and fen stages, as shown by the stratigraphic cross sec-
tion produced by Godwin & Mitchell (1938). The FBT
was relatively rapid when compared to other similar
raised bogs and radiocarbon dates suggest that the
transition was synchronous across a substantial part of
the mire. These lines of evidence are consistent with the
bog being subject to allogenic influences that acceler-
ated the rate of forward succession, although autogenic
processes such as peat accumulation would still have
contributed to the succession.
All that is needed for ombrotrophy to develop is a
separation of the mire growing surface from the affects
of surface and subsurface water supplies. This could be
achieved by vertical peat accumulation or by lowering
of the water table. Many sites where the FBT developed
in high effective precipitation regimes have stratigraphies
that record raised bog pool or lawn species at the FBT
(e.g. Abbeyknockmoy Bog), suggesting that a near-
surface water table can be maintained across the FBT
in humid conditions. One possible explanation for the
contrasting pathway of development at Tregaron is that
river channel incision lowered the elevation of the lagg
© 2003 British
streams, depriving the mire growing surface of surface
Ecological Society, and subsurface water flow. Leaching of the peat surface
Journal of Ecology, in the humid climate of 7300 cal. BP could have aided
91, 253–264 a swift, mire-wide transition to ombrotrophy.
13652745, 2003, 2, Downloaded from https://besjournals.onlinelibrary.wiley.com/doi/10.1046/j.1365-2745.2003.00762.x by Readcube (Labtiva Inc.), Wiley Online Library on [06/02/2023]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License
It would appear that peatlands may become ombrotro-
phic in a variety of water table conditions and climatic
regimes, and that the FBT in many British and Irish
sites varies in date from early to mid-Holocene. There
are at least two distinctive routes to ombrotrophy,
a ‘dry-pioneer’ route and a ‘wet-pioneer’ route, and
these may only be recognized from palaeoecological
analyses. The development of ombrotrophic condi-
tions should not be used automatically to infer a
change in the climate towards higher effective precipi-
tation without careful consideration of the type of FBT
involved, and the other factors that can affect water
table levels in the early stages of raised peat development.
Acknowledgements
The authors would like to thank the University of
Southampton for providing the funding support for a
research studentship from which the work presented
here is drawn. We are also indebted to the Countryside
Council for Wales (C.C.W) for permission to work on
Tregaron South-east Bog, the C.C.W. warden, Mr Paul
Culyer, at Tregaron Bog NNR for helping with field-
work and providing site information. The Southampton
Cartographic Unit is thanked for the preparation of
diagrams. The support of the NERC radiocarbon com-
mittee and laboratory staff at the East Kilbride facility
is gratefully acknowledged. Dr Michael Proctor and
an anonymous referee are thanked for comments on
an earlier draft of this paper.
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Received 30 July 2002
revision accepted 6 January 2003