THE HOLOCENE HISTORY OF TAXUS BACCATA (YEW) IN BELGIUM AND NEIGHBOURING
REGIONS
Author(s): Koen Deforce and Jan Bastiaens
Reviewed work(s):
Source: Belgian Journal of Botany, Vol. 140, No. 2 (2007), pp. 222-237
Published by: Royal Botanical Society of Belgium
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Belg. J. Bot. 140 (2) :222-237 (2007)
? 2007 Royal Botanical Society ofBelgium
THE HOLOCENE HISTORY OF TAXUSBACCATA(YEW)
INBELGIUM AND NEIGHBOURING REGIONS
Koen Deforce*
Flemish
Heritage
Institute, Koning
(*Author
for correspondence;
and Jan Bastiaens
Albert
II-Laan
e-mail:
19 bus 5,
-1210
Brussels,
Belgium
koen.deforce@rwo.vlaanderen.be)
Received 5 September2006; accepted 24November 2006.
?
is limited to
The current natural distribution of Taxus taccata L. inBelgium
a few localities in the southern part of the country. In these localities, Taxus is predominantly
growing on steep, calcareous
slopes, which is believed to be its natural habitat in this part of
Abstract.
In Flanders, the northern part of Belgium, Taxus is considered not to be native and
stands are interpreted there as being planted by humans or as garden escapes. The
data for Taxus, however, show a very different picture
Holocene
pollen and macrofossil
distribution, as well as habitat. It appears that during
regarding abundance and geographical
theworld.
Taxus
the Sub-boreal, Taxus grew in the coastal plain and the lower Scheldt valley, where itwas part
of the carr vegetation on peat. Before the end of the Sub-boreal, Taxus seems to have disap
peared from this region, most probably because of the transition from the carr vegetation to
is not the only case where such observations have been made. In other
(raised) bogs. Belgium
areas of northwestern Europe, Taxus also seems to have had a completely different distribution
and ecology in the past, especially during the Sub-boreal.
An
finds of Taxus taccata
from Belgium
is here given,
of the palaeobotanical
distribution and palaeo
supplemented with finds from neighbouring regions. The Holocene
in a broader northwest European context.
ecology of Taxus taccata are discussed
overview
?
Key words.
Taxus taccata
L., Belgium,
INTRODUCTION
During most of the Pleistocene interglacial
periods, Taxus formed a much more important
element of the vegetation of northwestern Europe
than during theHolocene (Averdieck 1971, God
win 1975, Zagwijn 1992, Lang 1994). Taxus has
been found inBelgium as early as theTiglian-C5
interglacial period (ca. 2-1.8 million yrs BP;
Pleistocene
chronozones according to Lang
1994) (Kasse 1988) and seems to have had its
greatest expansion innorthwesternEurope during
the Holstein interglacial (400 - 367 ka BP), a
Holocene,
vegetation
history, yew.
period characterised by a warm oceanic climate
1962, Kelly
1964,
(Jessen et al 1959, West
Godwin 1975,Watts
1985). High percentages of
both Taxus pollen and macroremains have been
found at several northwest European sites where
Holsteinian sediments are preserved. High per
centages of Taxus pollen have also been found at
several sites inBelgium, in sediments dating from
this period (De Gro?te
1977, Ponniah
1977,
Somm? et al 1978). The most famous palaeo
botanical record of Taxus dating from theHolstein
interglacial, however, is a spear made of Taxus
wood found at Clacton (Essex, UK; Godwin
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HISTORY OF TAXUS BACCATA INBELGIUM
1975), which is also one of the oldest known arte
factsmade ofwood.
During the last interglacial period before the
Holocene, the Eemian (130 -115 ka BP), Taxus
also played a more important role in the vegeta
tion in northwestern Europe than during the
Holocene, although the pollen percentages are
much lower than during theHolstein interglacial
1975,Woillard
1979,
(Behre 1962, Andersen
Lang 1994). Zagwijn (1983, 1992) even distin
guished a Taxus subzone in his zonation of the
Eemian period in theNetherlands. In Belgium,
both pollen and wood of Taxus have been found
inEemian peat deposits at Be?rnem (De Gro?te
1977, Deforce
1997, Klinck 1999).
During the present interglacial period, the
Holocene, Taxus seems to have played a less
importantrole in the vegetation. However, during
the Sub-boreal (5 000 - 2 500 uncal. BP;
et
Holocene chronozones according toMangerud
was
more
abundant and showed a
al. 1974), Taxus
completely differentdistribution and ecology than
nowadays. The aim of this paper is to review the
available data on the Holocene history of Taxus
and to discuss its past distribution and ecology.
Furthermore, the existing hypotheses for the
Holocene Taxus decline will be evaluated in the
lightof the available palaeobotanical data.
PRESENT-DAYECOLOGY AND
DISTRIBUTION
There is no scientific agreement on the exact
taxonomic position of the genus Taxus, which
encompasses about seven closely related species
scattered throughout the northern temperate
1986, Dempsey & Hook 2000,
region (Voliotis
Thomas & Polward 2003). The species separa
tion within the genus is equally disputed (van
Vuure 1990, Delahunty
2002, Thomas & Pol
wart 2003), although recent genetic research
et al. 2003) shows that the current
(Collins
arewell founded.
delimitations
species
Taxus baccata L. (subsequently referred to
as Taxus), the species native toEurope, is an ever
green needle-leaved gymnosperm shrub or tree,
growing up to 28 m high. The species is slow
223
growing and long-living, reaching maturity only
at ca. 70 years. It is extremely shade tolerant but
can withstand full exposure to the sun (Tutin et
al 1964, Thomas & Polwart 2003). Taxus is
and is
normally dioecious, rarely monoecious
wind-pollinated (Tutin et al 1964, Thomas &
Polward 2003). Itflowers from February toApril
(Richard 1985).
Taxus occurs throughoutmost of Europe and
some parts of northernAfrica, although its distri
bution is very scattered. Taxus thrives best in
regions with a mild, oceanic climate. Its distribu
tion is limited by low temperatures in Scandi
a severe continental climate in eastern
navia,
Europe and aridity and high temperatures in
Turkey and north Africa (Thomas & Polwart
2003). In theMediterranean region, Taxus is con
fined to thehighermountains (Tutin et al 1964).
Taxus does not form pure monospecies
stands (except in theCaucasus Mountains and on
chalk and limestone in England) but belongs to
diverse forest communities mainly composed of
Abies, Fagus, Carpinus, Alnus and Picea (Ellen
berg et al 1991, Jahn 1991, Iszulo & Boratyn
ski 2004).
In Europe, most of the natural stands of
Taxus grow on well-drained calcareous soils,
although the tree can grow on almost any soil,
including silicious soils derived from igneous and
sedimentary rocks (Thomas & Polwart 2003). In
most countries, Taxus is a declining or even
threatened species. The reasons are thought to be
deforestation, selective felling and grazing
1979, Svenning & Mag?rd
1999,
(Muhle
Navys 2000, Holtan 2001, Thomas & Polwart
2003, Mysterud & 0stbye 2004). Several pro
tected areas have been established to conserve the
1991, Hartzell
species (Svalastog & Holand
Kr?l
Brande
1991,
1993,
2002), which also sur
vives in a cultivated form as an ornamental tree in
parks, gardens, and cemeteries (Kr?ssmann
1972, Saintenoy-Simon 2006).
In Belgium, the current natural distribution
of Taxus is limited to a few localities in the south
ern part of the country, namely the southern and
western part of theMeuse district (Fig. 1; Lawal
r?e 1952, Duvigneaud
1965, Galoux
1979, van
Rompaey & Devosalle
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1979, Saintenoy-Simon
224
BELGIAN JOURNALOF BOTANY 140
1983, 2006, Lambinon et al. 1998). At these
localities, Taxus predominantly grows on steep,
calcareous slopes, which are believed to be its
natural habitat in the region (Lambinon et al.
1998). Taxus grows there togetherwith Fagus syl
vatica, Carpinus
betulus, Quercus
robur, Q.
Tilia platyphyllos and, sometimes,
petraea,
Ulmus spp. and Corylus avellana (Duvigneaud
1965, Galoux
1979). In Flanders, the northern
of
part
Belgium, Taxus is considered not to be
native and Taxus stands are interpreted there as
having been planted by humans or as escapes
Fig. 1.Map showingboth the currentnaturaldistributionand subfossilfinds of Taxus baccata inBelgium: white
dots denote
tenoy-Simon
natural
& Devosalle
1952, van Rompaey
(data from Lawalr?e
populations
et al. 2006); black symbols denote Holocene
et al. 2006, and Maes
Landuyt
T. baccata
2006, Van
1979, Sain
palaeobotan
ical recordsofpollen, seeds andwood/charcoal ofTaxus. (1)Avekapelle 363 (Baeteman & Verbruggen 1979); (2)
Booitshoeke (Baeteman & Verbruggen 1979); (3) Oudekapelle (Stockmans & Vanhoorne 1954); (4) Sint
inpress); (6) Leffinge
Jacobs-Kapelle (Stockmans & Vanhoorne 1954); (5) Raversijde (Deforce & Bastiaens,
(Baeteman et al. 1981); (7)Wenduine (Mertens 1958); (8) Blankenbergse vaart Zuid (Allemeersch 1991); (9)
Heusden (Stockmans 1945); (10) Laarne - Damvallei (Verbruggen 1971); (11) Borsele (NL)(Van Run 2001);
(12) Ellewoutsdijk (NL)(Van Run 2003, Van Smeerdijk 2003); (13) Baarland (NL)(De Jong 1986); (14)
Terneuzen (Munaut 1967a,b); (15)Waasmunster - Pontrave (Merckx 1996); (16)Weert (Minnaert 1982); (17)
Verrebroek (Deforce et al. 2005); (18) Doel (Minnaert & Verbruggen 1986); (Deforce et al. 2005); (19) Zand
vliet (Munaut 1967a); (20) Oorderen (Munaut 1967a); (21) Kruisschans (Vanhoorne 1951); (22) La Karelsl?
(Waldbillig,easternGutland, Luxembourg) (Pernaud 2001).
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HISTORY OF TAXUS BACCATA INBELGIUM
from gardens and parks (Lambinon et al. 1998,
Maes et al 2006).
In other parts of northwestern Europe, the
natural distribution of Taxus is also mainly con
fined to soils on limestone and other types of
well-drained, base-rich soils (UK and Ireland:
Tittensor
1980, Kelly & Kirby 1982, Stace
et al. 2002; The Netherlands:
Preston
1997,
et al. 1985; Germany: J?ger & Werner
Weeda
2002; France and Switzerland: Palese & Aeschi
mann 1990; Scandinavia: Jonsell 2000, Moss
berg & Stenberg 2003).
PALAEOBOTANICALRECORDS OF TAXUS
The pollen data
225
were present in the investigated peat sequence.
Similarly, pollen analysis of peat sequences
from Kruisschans
(Antwerp, Belgium; Van
hoorne
(Diksmuide, Bel
1951), Oudekapelle
&
Vanhoorne
Stockmans
1954) and
gium;
(Diksmuide,
Sint-Jacobs-Kapelle
Belgium;
Stockmans & Vanhoorne
1954) did not reveal
Taxus pollen during the palynological research,
while several Taxus seeds were recovered from
the same peat sequences investigated.
Holocene Taxus pollen recordsfrom Belgium and
immediate surroundings
Holocene
records of Taxus pollen from
are
not
very abundant. Out of a total of
Belgium
370 palynological studies from all over Belgium
that were
Characteristics of'Taxus pollen
Taxus pollen is spherical to obtusely angu
its
size ranging between 19.3 and 29.8
lar,
after acetolysis (Averdieck 1971, Beug 2004).
Taxus pollen does not have sacci and is inapertu
rate. The exine is intectate and has a scabrate or
microgemmate sculpturing (Moore et al. 1991,
Beug 2004). Regarding identification, confusion
might be possible with pollen of Juniperus, Pop
ulus, Rhynchospora and Quercus
(Averdieck
et al. 1991, Beug
1971, Zoller
1981, Moore
2004). The pollen grains frequently split and are
to
corrosion
sensitive
1967,
(Havinga
Averdieck
1977,
1971, Rohr & Kilbertus
Taxus
is an
Bradshaw
1978). Although
the
tree,
pollen representation in
anemophilous
surface samples near Taxus stands seems to be
rather low and decreases sharplywith increasing
distance from the Taxus stand (Heim 1970,
Noryskiewicz
2003). These factors, in combina
tionwith the lack of distinctive features such as
sacci, apertures or a distinctive exine sculptur
ing, make it likely that Taxus pollen was not
recognised in some of the earlier palynological
studies (K?ster
1994), or at sites where condi
tions for pollen preservation were poor. During
the palynological
investigation of Wood Fen
et
al 1935), for example, no
Godwin
(Ely, UK;
Taxus pollen was found while several trunks and
even the pollen-bearing cone scales of Taxus
reviewed (Deforce & Bastiaens
2006), only 12 contained substantial records of
Taxus. Sites where Taxus occurred in only one
or a few samples, and with a frequency of less
than 1%, are not included in the overview pre
sented here, as this might represent long-dis
tance transport. On the other hand, itmust be
remembered that in some of the older analyses,
Taxus pollen was very probably overlooked, as
above. Three
additional
already discussed
records derived from the southwestern Nether
lands, close to the Belgian border, are included
in the discussion.
Together, the records show two remarkable
characteristics: (1) they are all situated in the
coastal plain and the lower Scheldt valley (Fig.
1), and (2) they can all be dated from the Sub
boreal. Some of these dates only rely upon bio
zonation as the interpretation of the older dia
grams is not supported by radiocarbon dating. But
still, the available 14C dates (Table 1) allow the
conclusion thatTaxus appears in pollen diagrams
inBelgium and the southernNetherlands between
4750 ? 140 uncal. BP (Oorderen) and 4280 ? 130
uncal. BP (Terneuzen). At all sites Taxus percent
ages remain rather low, varying between 2 and
6%. The percentages are higher only at Zandvliet
(10.1%) and Ellewoutsdijk (16%). Taxus disap
pears from the pollen diagrams between 4035 ?
30 uncal. BP (Raversijde) and 3510 ? 45 uncal.
BP (Baarland)(Table 1).
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BELGIAN JOURNALOF BOTANY 140
226
The seed data
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Characteristics of Taxus seeds
The seeds of Taxus are highly characteristic:
they are large (6-8 mm) and ellipsoid-ovoid with
a tapering upper end, and rounded to slightly tri
angular or biconvex in section; their surface is
smooth. Seeds cannot be mistaken for any other
species. When fresh, seeds are surrounded by a
conspicuous reddish aril. This aril is the only non
toxic part of Taxus, and it is eaten and digested by
birds and small mammals, while the seed itself
passes the intestinal canal undamaged (Weeda et
al. 1985). Birds are themain agent of seed disper
sal (Zoller
1981, Thomas & Polwart 2003),
which occurs from September into the winter
et al. 2000). As the
1981, Bouman
(Zoller
fleshy aril does not preserve, palaeobotanical
finds of Taxus seeds always lack thatpart.
Holocene Taxus seed recordsfrom Belgium and
immediate surroundings
A small number of Holocene
records of
seeds of Taxus are known from Belgium: subfos
sil seeds have been found atOudekapelle (Stock
mans & Vanhoorne
1954) and Sint-Jacobs
1954), in the
Kapelle (Stockmans & Vanhoorne
western coastal area, and at Kruisschans (Van
hoorne 1951) and Heusden (Stockmans 1945),
both along theriver Scheldt (Fig. 1).
These findsoriginate from the same restricted
region thatyielded subfossil pollen, i.e. the coastal
area and the lower Scheldt valley. The Taxus seeds
have all been recovered from peat deposits, none of
which have been dated by radiocarbon analysis.
Nevertheless, all finds can be attributed to theLate
Atlantic or Sub-boreal on thebasis of theirposition
in the lower part of the so-called surface peat (see
further),or of pollen analysis carried out on the
same peat sequences (Vanhoorne
1945, Van
hoorne 1951, Stockmans & Vanhoorne
1954).
Wood
and charcoal
data
PQ
Characteristics ?/Taxus wood and charcoal
The wood of Taxus is very dense, hard,
elastic and resistant to decay (Zoller
1981). The
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HISTORY OF TAXUS BACCATA INBELGIUM
sapwood iswhite to yellowish; the heartwood is
red and colours orange-brown after contact with
the air. Owing to its good elasticity, ithas been a
very popular timber for the production of tools
and weapons, in particular bows (Zoller
1981,
Lanting et al 1999, Gale & Cutler
2000).
Taxus wood and charcoal are easy to differentiate
from that of other European gymnosperms, on
account of the distinct spiral thickenings in the
tracheid walls and the absence of resin canals in
the former (Grosser
1977, Schweingr?ber
1990, Gale
& Cutler
2000).
Taxus wood and charcoal recordsfrom
Belgium and immediate surroundings
Holocene
Subfossil Holocene wood remains of Taxus
are only known from Blanken
from Belgium
1991) and Wenduine
berge
(Allemeersch
At
(Mertens 1958).
Blankenberge, Taxus wood
was found embedded in a peat deposit dating
from the lateAtlantic or the Sub-boreal. At Wen
duine, a fragment of Taxus wood was found in an
archaeological site dating from theRoman period
- 402
AD; archaeological periods accord
(57 BC
to
Slechten
2004).
ing
In the southern part of theNetherlands, sub
fossil wood of Taxus has been found at Terneuzen
(Munaut 1967a,b), Borsele (Van Run 2001) and
Ellewoutsdijk (Van Run 2003). At Terneuzen,
several Taxus stems have been recovered from
peat deposits but they have not been dated and
their stratigraphieposition has not been recorded.
However, it is very likely that the stems derive
from the same levels from which Taxus pollen
was recovered, which would place thewood frag
ments in the Sub-boreal (Munaut 1967a, God
win 1968). At Borsele and Ellewoutsdijk, the
Taxus finds were partly preserved in thepeaty soil
and consisted of wooden poles thatwere used as
parts of Roman Age buildings. One Taxus pole
from Borsele was radiocarbon-dated at 4690 ? 60
(several poles made from Pinus
sylvestris gave similar dates) (Sier 2001). The
Taxus poles from Ellewoutsdijk were not dated by
uncal.
BP
radiocarbon analysis but a pole from P. sylvestris
from the same buildings was dated at 4480 ? 25
uncal. BP. The only possible explanation for the
227
time gap between the time of the construction and
themuch older age of part of the construction
wood is that at these sites during Roman times,
subfossil wood was used for the construction of
buildings (Van Rijn 2003). As inferredfrom the
pollen analysis of Ellewoutsdijk (Van Smeerdijk
2003) and the palaeogeographical map of the
1997), these sites
region (Vos & Van Heeringen
were situated in an almost treeless peat-bog envi
ronment during the Roman Age, which might
explain theuse of subfossilmaterial.
The Taxus wood found at theRoman site of
Wenduine (Mertens 1958) has not been dated but
itmight represent subfossil wood too, given that
Wenduine was also situated in a peat bog or a
peri-marine environment during Roman Age
1991, Ervynck et al. 1999).
(Allemeersch
Charcoal from Taxus has been found in La
Karelsl?, eastern Gutland (Luxemburg) (Fig. 1),
in archaeological cave deposits. A few fragments
derive from deposits dating from the Middle
Neolithic (4 500 - 3 500 BC; which corresponds
to the Late Atlantic) while rather large amounts
been recovered from Late Bronze Age
- 800
BC, corresponding to the
deposits (1 100
Late Sub-boreal) (Pernaud 2001). This is rather
surprising as there is no evidence for the exten
sion or even presence of Taxus in any of the
have
palynological records from Luxemburg
(Pernaud 2001), eastern Belgium and theGaume
district (Couteaux
1969a,b), the Plateau des
Tailles area (Mullenders & Knop 1962) or the
French Ardennes (Mullenders
1960, Lefevre et
Holocene
al. 1993).
All dated finds ofHolocene Taxus wood and
charcoal are of Late Atlantic or Sub-boreal age
and, except for the charcoal from Gutland, all
finds were excavated in theBelgian coastal plain
or the Scheldt estuary.
palaeobotanical
records of Taxus
Holocene
from other parts of northwestern europe
Probably one of the earliest mentions of
Taxus occurring in peat deposits was made by
Staring
(1983, re-edition from 1856). The
author expressed his surprise about the presence
of subfossil Taxus wood inDutch peat deposits,
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228
BELGIAN JOURNALOF BOTANY 140
in contrast with the 19thcentury distribution of
Taxus in The Netherlands and elsewhere in
Europe. That Taxus grew on peat has been further
demonstrated by the finds of subfossil wood at
Ely (UK), described by Miller and Skertchley
(1878) and discussed by Godwin et al (1935).
Another early observation of Taxus trunks recov
ered from peat deposits was made at Ballyfin
Bog (Ireland) by Adams (1905), who stated that
similar finds were so plentiful in former times
that farmers in the neighbourhood used thewood
for gate posts, house roofs, etc.
Firbas (1949) reviewed Holocene records of
Taxus wood from Germany, both from natural
peat deposits and from archaeological contexts
including several trunksfrom peat deposits from
the coastal lowlands of Ostfriesland. Other finds
of Taxus wood recovered from peat deposits in
Germany are listed by Hayen (1960, 1966) and
Averdieck(1971).
Next to these finds of subfossil wood, there
are also numerous pollen diagrams from north
western Europe, showing a distinct Taxus curve,
mostly during the Sub-boreal (for northwestern
1971, 1983, Hayen
Germany, see Averdieck
et al
1960; for Ireland: O'Connell
1988,
et al 1996, Molloy
Mitchell
& O'Connell
for England: Godwin
2004;
1975, Peglar
et al 2004; for
1993a,b, Greig 1996, Batchelor
northwestern France: Van Zeist 1964; for Swe
den: Berglund
1966, for Finland: Sarmaja
Korjonen et al 1991).
DISCUSSION
From theHolocene palaeobotanical records
of Taxus presented, it is clear that this treewas
more abundant and had a different distribution
during the Sub-boreal in northwestern Europe
compared to nowadays.
Taxus occurs sporadically inpollen diagrams
from England (Birks 1982, Godwin 1975), Ire
land (Huang 2002) and Sweden (Berglund
1966) from the Late Boreal or early Atlantic
onwards. For Belgium, the earliest post-glacial
palaeobotanical records of Taxus date from the
late Atlantic or early Sub-boreal. All the other
records from Belgium and the
palaeobotanical
southernNetherlands can be dated from the Sub
boreal as well. Not only in Belgium, but also in
other parts of northwestern Europe (see 3.4),
Taxus shows a maximum in pollen diagrams dur
ing the Sub-boreal.
Distribution
and habitat of Taxus during the
Sub-boreal
Most of the finds of subfossil pollen, seeds
and wood of Taxus inBelgium and the southern
part of theNetherlands are situated in the coastal
lowlands andmore precisely in the area where the
so-called 'surface peat' or 'Holland peat' occurs
in the subsoil. This surface peat was formed dur
ing themid Holocene when the postglacial sea
level rise began to slow down and coastal barriers
could develop (Baeteman
1981, 1999, Vos &
Van Heeringen
These
coastal barriers
1997).
closed the coast almost completely and initiated
mire development. At the time of themaximal
expansion of thepeat, in the Sub-boreal, themires
stretched nearly continuously from Calais
in
northwestern France to southwestern Denmark,
including the coastal plain of Belgium, thewest
ern part of theNetherlands and the lower Scheldt
valley (Pons 1992).
According to Pons (1992), this surface peat
shows more or less the same general development
in the coastal plain inFlanders and the southwest
ern part of theNetherlands. On the saltmarshes of
the regression surface brackish fens developed,
forming Phragmites (Scirpus) peat. Gradually,
d?salinisation and decreasing amounts of avail
able nutrients resulted inCarex-Phragmites fens,
which changed into mesotrophic Betula-Alnus
carr, sometimes with some Pinus. This phase of
carr peat is followed by a transition to Sphagnum
peat and, in most places, the development of
raised bogs resulting in the formation of Sphag
Hwm-Ericaceae peat. Peat growth ended because of
marine transgressions and fluvial sedimentation
between theLate Sub-boreal and theLate Middle
Ages, depending on the location,which resulted in
the covering of the peat with marine and alluvial
1985,
clay deposits (Janssens & Ferguson
Denys & Verbruggen
1989,Allemeersch
1991,
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All use subject to JSTOR Terms and Conditions
HISTORY OF TAXUS BACCATA INBELGIUM
Pons 1992). The upper part of the surface peat is
often lacking as a consequence ofmarine erosion
or medieval and post-medieval peat extraction
et al. 2002, Baeteman
(Baeteman
2005).
Macroremains of Taxus from the coastal plain of
Belgium and the southernNetherlands (Fig. 1),
except those associated with archaeological sites,
have all been recovered from the part correspon
ding to the carr-phase of the peat profiles. They
were associated with seeds and othermacroscopic
remains of plants typical of an alder/birch carr or
fen vegetation composed of, e.g., Alnus glutinosa,
Betula
sp., Comarum
palustre,
Carex
paniculata,
Carex pseudocyperus, Lysimachia vulgaris, Lyco
pus europaeus and Thelypteris palustris (Van
Hoorne
1954,
1951, Stockmans & Vanhoorne
occurrence
in
Allemeersch
of
Taxus
The
1991).
areas
corre
from
these
also
pollen diagrams
sponds with the carr-phase of the analysed peat
1967, Deforce & Basti
profiles (e.g., Munaut
aens inpress).
The fact that, at sites mentioned above,
Taxus was actually part of the local vegetation
community has been ignored or even denied by
several authors as itdoes not seem to correspond
with the present day ecology and distribution of
this tree. Van Smeerdijk (2003: 161-162), for
example, argued that the high percentages of
Taxus pollen at Ellewoutsdijk and Baarland must
be explained by transportby theriver Scheldt and
thus must originate from a more inland area.
However, the fact thatTaxus wood has been dis
covered at Ellewoutsdijk
and at the nearby
Terneuzen, does not support this hypothesis.
Besides, theriver Scheldt followed a more north
ern route at that time and did not flow near Elle
woutsdijk or Baarland (Vos & van Heeringen
1997, Denys & Verbruggen
1989). Many other
records of pollen and macroremains of Taxus
from peat deposits, often in areas without any flu
vial activity, indicate thatTaxus actually did grow
on peat. In fact, this is not an entirely new obser
vation as Godwin already remarked, based on his
research atWood Fen (Ely, UK; Godwin et al.
1935), atWoodwalton Fen (Hunts, UK; Godwin
& Clifford
1938) and on the Taxus finds at
Terneuzen (The Netherlands; Godwin 1968), that
"Taxus almost certainly had an extensive natural
229
occurrence upon peat land, although natural com
munities of this kind can no longer be pointed
out" (Godwin 1968: 737).
Ithas to be stressed here thatpalaeobotanical
data are generally sparse for chalk regions, since
the geological and topographical conditions in
these regions are unsuitable for the formation and
preservation of stratifiedpeat (Tittensor 1980).
This could, of course, bias the reconstruction of
the Holocene
distribution of Taxus presented.
However, this objection may be valid for the
areas with actual natural Taxus populations in
Belgium (Fig. 1) but this is not true for the areas
both to the northwest and to the southeast of this
region. For those areas, Holocene pollen dia
grams and other palaeobotanical data are avail
able, but no Taxus pollen were recorded outside
the coastal area and the lower Scheldt valley, one
exception being the records of Taxus charcoal
from Gutland (Luxembourg).
The Taxus decline
At the above-mentioned sites from Belgium
and surrounding regions, Taxus disappears in the
pollen diagrams during the second half of the
Sub-boreal, around 3 500 uncal. BP (see Table 1).
Similarly, no botanical macroremains of Taxus
have been found that are younger than the end of
the Sub-boreal. In pollen diagrams from other
sites situated in the lowlands of northwestern
Europe, Taxus disappears as well, or shows a
strong decline, before the end of the Sub-boreal.
The Holocene decline of Taxus in northwest
ern Europe is generally attributed to competition
with Fagus and Carpinus, deforestation, selective
1949, Averdieck
felling and grazing (Firbas
1971, Zoller
1981, Svenning & Mag?rd
1999,
Navys 2000, Holtan 2001, Thomas & Polwart
2003). However, these explanations are all based
on the actual ecology and distribution of Taxus,
i.e., Taxus growing on well-drained calcareous
soils. For thedecline of Taxus growing in a fen carr
environment, other explanations must be sought.
The most common explanation for the Sub
boreal Taxus decline
and Carpinus (Firbas
Muhle
1979). Recent
This content downloaded on Thu, 10 Jan 2013 04:53:02 AM
All use subject to JSTOR Terms and Conditions
is competition with Fagus
1949, Averdieck
1971,
research showed that a
230
BELGIAN JOURNALOF BOTANY 140
Taxus population in Denmark increased after
thinning the tree canopy, especially by felling
beech (Svenntng & Mag?rd
1999). Other
research demonstrated, however, that regenera
tion of Taxus could be rather successful under the
canopies of several broadleaved trees, including
Carpinus betulus (Iszkulo & Boratynski 2004).
Moreover, as Fagus and Carpinus only grow on
well-drained soils (Ellenberg et al 1991), light
competition with these two taxa cannot have
played a role in the decline of Taxus growing in
wet conditions.
Another explanation for the Taxus decline
could be deforestation, but the pollen diagrams
from Belgium and the southernNetherlands do
not indicate deforestation during the period of the
Taxus decline. There are also no indications for
agriculture or other forms of intensive human
impact on the vegetation in the coastal lowlands
during the Sub-boreal (Vos & van Heeringen
1997, Ervynck et al 1999).
Selective felling of Taxus, for its valuable
wood or to avoid cattle poisoning, has been pro
posed as another explanation for the Taxus
decline at several sites in northwestern Europe
et al 1991, Svenning &
(Sarmaja-Korjonen
Mag?rd
& Molloy
O'Connell
1999,
2001).
From theNeolithic onwards, Taxus was probably
themost used wood for themanufacture of bows
et al
1963, Lanting
1999, Beuker
(Clark
other
wooden
2002). Many
implements were
made fromTaxus as well (Godwin 1975, Coles et
al 1978, Gale & Cutler 2000). However, since
human populations and activities were almost
absent during the Sub-boreal, in the region under
consideration here, these explanations can also be
1997, Ervynck
rejected (Vos & van Heeringen
et al 1999, Louwe Kooijmans et al 2005).
In several forests of northwestern Europe, it
has been observed thatTaxus recruitment suffers
from browsing by roe deer, Capreolus capreolus
(Garcia & Obeso 2003, Mysterud & Ostbye
2004). The branchlets, needles and seeds of Taxus
contain a poisonous alkaloid, a lethal toxin for
many species including horses, cows, goats and
humans (Jordan 1964, Schulte
1975). Only a
few animals including roe deer are not sensitive
to it; they like to nibble the yew branchlets and
cause tangible damage to the trees (Mysterud &
0stbye
1995, 2004 Navys 2000). Fully-grown
Taxus trees are largely resistant to the nibbling;
even after a tree is cut down, green shoots appear
from the stump. On the other hand, recent
research has demonstrated that roe deer browsing
reduces Taxus recruitment (Hulme 1996, Garcia
& Obeso 2003, Mysterud & Ostbye 2004). In
general, although roe deer occur inwetland habi
tats (Danilkin
1996, Barancekova
2004), it is
Taxus
that
the
Sub-boreal
decline
very unlikely
can be explained by roe deer -browsing, as there
are no indications for an increase in theirpopula
tion at that time.
One more possible explanation for the Taxus
decline would be a climate change. There is no
evidence, however, for a major change of the cli
matic conditions in northwestern Europe around
3 500 uncal. BP (Davis et al 2003). As Belgium
and the southernNetherlands are not situated near
the limits of the natural distribution of Taxus, it is
unlikely that a minor change in climatic condi
tionswould have caused the Taxus decline.
In conclusion, although some of the above
mentioned explanations for the Taxus decline
might hold true forTaxus stands growing on well
drained, mineral soils or in regions where human
intense (Tittensor
1980,
impact was more
O'Connell
& Molloy
2001), they are not suit
able to explain the Sub-boreal decline of Taxus in
the coastal area of Belgium and the southern
Netherlands. As a more likely explanation for the
decline of Taxus in Belgium and the southern
Netherlands, a change of the environment inwhich
Taxus grew can be proposed. Inmost places in the
coastal area and the Scheldt estuary, this environ
mental change could consist of the already men
tioned transitionfrom the carr peat phase, inwhich
most of the palaeobotanical records of Taxus can
be situated, to ombrotrophic moss peat and, in
most places, thedevelopment of raised bogs result
ing in the formation of Sphagnum-E?ca.ce&e peat
1992, Ver
1986, 1991, Pons
(Allemeersch
bruggen et al 1996, Deforce & Bastiaens
in
the pollen diagrams from
press.). Especially
and Raversijde
Oorderen
1967)
(Munaut
Bastiaens
in
&
show
very clearly
press)
(Deforce
thatTaxus indeed disappears with the transition to
This content downloaded on Thu, 10 Jan 2013 04:53:02 AM
All use subject to JSTOR Terms and Conditions
HISTORY OF TAXUS BACCATA INBELGIUM 231
ombrotrophic conditions, illustrated by the
increase of Sphagnum and GzZ/ima/Ericaceae.
In the inland part of the lower Scheldt valley,
where no ombrotrophic peat occurs on top of the
fen-carrpeat deposits, the end of the peat growth
was caused by the deposition of alluvial loam and
clay, as a consequence of agricultural practices
(Verbruggen
et al.
1996, Huybrechts
1999).
Allemeersch
1991.
L.,
?
Peat
eastern
in the Belgian
F. (ed.), Wetlands
In: Gullentops
coastal
plain.
in Flanders.
Contributions
zone
ogy of the temperate
to the palaeohydrol
in the last 15 000years.
AardkundigeMededelingen 6, pp. 1-54. Leuven
Press, Leuven.
University
?
Andersen
1975.
The
S.Th.,
Eemian
freshwater
deposit at Egernsund, South Jylland, and the
Eemian
landscape
inDenmark.
development
Dan
marks Geologiske Undersogelse A 1974: 49-70.
Averdieck
CONCLUSIONS
Flora
?
westdeutschland.
peat. This stronglycontrastswith the recent distri
bution and ecology of Taxus, the current natural
distribution of Taxus baccata L. inBelgium being
limited to a few localities in the southernpart of the
country, all situated on steep, calcareous slopes.
The Holocene occurrence of Taxus in the
coastal plain inBelgium, and probably in several
other lowland areas in northwestern Europe, cor
relates with the carr peat phase of the surface or
Holland peat, which was mainly formed during
the Sub-boreal. The decline and disappearance of
Taxus in northern Belgium and the southern
Netherlands during the second half of the Sub
boreal are most likely the result of the transition
of these coastal marshes from a fen-carr environ
ment to ombrotrophic bogs.
ACKNOWLEDGEMENTS
Averdieck
postglazialen
comments.
1983.
F.-R.,
160:
28-42.
Palynological
investigations
of ten lakes in eastern Holstein,
of the sediments
North Germany. Hydrobiologia
Baeteman
C,
1981.
C,
1999.
?
De Holocene
103: 225-230.
ontwikkeling
van
depositional
his
de westelijke kustvlakte (Belgi?), Ph.D. thesis,
Vrije UniversiteitBr?ssel, Br?ssel, Belgium.
Baeteman
?
The Holocene
toryof the Ijzer Palaeo-valley (westernBelgian
coastal
with
plain)
con
to the factors
reference
trollingthe formationof intercalatedpeat beds.
Geologica Belg 2: 39-72.
Baeteman
2005.
C,
?
How
subsoil morphology
and
erodibilityinfluencetheoriginand patternof late
Holocene
tidal
channels:
case
lowlands.
Quatern.
coastal
Belgian
2146-2162.
Baeteman
C.
&
Verbruggen
studies
from
Sci.
1979.
C,
to the evolution
24:
A
new
?
of the so-called
the
Rev.
approach
in the western
coastal
fessional Paper
167, Belgische Geologische
peat
plain
surface
Pro
of Belgium.
Br?ssel.
Dienst,
Baeteman
Cleveringa
C,
1981. ?
P. &
Het paleomilieu
van Leffinge.
zoutwinningssite
Verbruggen
C,
rond het romeins
Professional
Paper
186,Belgische Geologische Dienst, Br?ssel.
Baeteman
The authorswish to thankNele Van Gemert for
help with the production of the figures and Otto
for valuable
Zur
Geschichte der Eibe (Taxus baccata L.) inNord
All Holocene
records of
palaeobotanical
Taxus from Belgium and the southernNetherlands
show three remarkable characteristics: (1) they all
are situated in the coastal plain and the lower
Scheldt valley, (2) they all date from the Sub
boreal and (3) they all indicate thatTaxus grew on
Brinkkemper
?
1971.
F.-R.,
2002.
Scott
D.B.
& Van
Strydonck
M.,
zone processes
in coastal
at a
Changes
sea-level
stand: a late Holocene
example
C,
?
high
from Belgium.
J. Quatern.
Sci. 17: 547-559.
?
Barancekova
The roe deer diet: Is flood
M., 2004.
plain forestoptimalhabitat?Folia Zool. 53: 285
292.
REFERENCES
Adams
J., 1905.
?
Batchelor
The
occurrence
of yew
in a peat
bog inQueen's County. IrishNaturalist 14: 34.
Allemeersch
L.,
1986.
?
Hochmoortorfe
?stlichen K?stengebiet
Forsch.-Inst.
CR,
Branch
N.,
Elias
S., Hawkins
Shaw P. & Sidell J.,2004.?Middle
Seckenberg
Belgiens.
86:
397-407.
im
Courier
environmental
history
D.
Holocene
of the lower Thames
val
ley.The history of Taxus woodland, pp. 79-80.
QRA International Postgraduate Symposium,
Royal Belgian Institute of Natural Sciences,
Br?ssel,
Belgium.
This content downloaded on Thu, 10 Jan 2013 04:53:02 AM
All use subject to JSTOR Terms and Conditions
BELGIAN JOURNALOF BOTANY 140
232
Behre
?
.-E., 1962.
Pollen-
und diatomeenanalytis
an letztinterglazialen
Kiezel
che Untersuchungen
Davis
Berglund
?
1966.
B.E.,
in eastern
Late-Quaternary
southeastern
Blekinge,
II.
study.
analytical
Opera Bot. 12: 1-190.
?
.-!., 2004.
Leitfaden
pollen
Beug
vegetation
A
Sweden.
Beuker
J.R., 2002.
Nieuwe
?
Drentse
H.J.B.,
1982.
Roudsea
Wood
Birks
der Pollenbestimmung
in het moeras.
Een boogschutter
Volksalm?nak
119: 113-122.
?
Mid-Flandrian
National
forest history of
Cum
Reserve,
Nature
90: 339-354.
bria. NewPhytol.
Bouman
F., Boesewinkel
D., Bregman
?
N. & Oostermeijer
G., 2000.
van
1978. ?Modern
R.H.W.,
Ph.D.
England.
Cambridge,
Brande
A., 2002.
thesis, University
U.K.
?
Internationalen
1-36.
Deforce
history in S.E.
of Cambridge,
Clark
1963.
J.G.D.,
?
Neolithic
and Ireland. Proc.
Collins
D.,
Mill
Prehist.
R.R.
&
44:
Soc.
M?ller
1-45.
M.,
2003.
their reputedhybrids inferredfrom RAPD
cpDNA
Couteaux
M.,
and
,4m. J. Bot. 90: 175-182.
?
Recherches
1969a.
palynologiques
data.
en Gaume,
au pays
d'Arlon,
en Ardenne
m?rid
ionale (Luxembourgbelge) et au Gutland (Grand
Duch? de Luxembourg). Acta Geographica
Lovaniensia,
Leuven,
Couteaux
vol.
Belgium.
1969b.
M.,
9.
Leuven
?
University
Formation
Press,
et chronologie
palynologiques des tufscalcaires du Luxembourg
Ass. Fran?aise
belgo-grand-ducal.
ternaire 3: 179-206.
Danilkin
A.,
1996. ?Behavioural
and European
London.
roe deer.
?tude
du Qua
of Siberian
300 p. Chapman
& Hall,
ecology
In: Cromb?
vol.
ects,
University
De
De
?
T. canaden
of Taxus baccata,
separation
T. cuspidata
and origins of
(Taxaceae)
Species
sis and
1992-2002.
Verbruggen
V.,
?
2005.
L.,
The
gium).
western
Proc.
Soc.
29: 50-98.
Prehist.
Europe.
?
Heal
&
S.V.E.
Orme
The
1978.
B.J.,
J.M.,
use and character of wood
in prehistoric Britain
Noordzeegebied.
Belgi?)
Pollen
Ph.
(ed.),
C.
&
and
The
phytolith
last hunter
gatherer-fishermeninSandy Flanders (NW Bel
England, and theprehistoryof archery in north
Coles
Gelorini
K.,
analyses.
Eibenfre
bows from Somerset,
(Oostende,
Vrydaghs
im polischen
2002. Der
Eibentagung
voor
van 10 jaar opgraven, Arche
Opgravingsverslag
in
6. V.I.O.E.,
Vlaanderen,
Monograf?e
ologie
Br?ssel.
Verspreiding
Eibe-Vorkommen
Inventarisatie
bodemarchief
in het zuidelijke
Raversijde
Tiefland - eine ?bersicht zur Exkursion der 9er
und9.
het
vissersmilieu
represen
?
J., 2006.
Gent,
Pieters M. (ed.), Een laatmiddeleeuws landelijk
Devente
R,
pollen
recent woodland
and
Bastiaens
Gent,
en bescherming,
CAI
II.
2:
35-42.
rapport
VIOE-rapporten
?
Paleob
Deforce
K. & Bastiaens
J., in press.
van
het oppervlakteveen.
In:
onderzoek
otanisch
van zaden: 240 p. KNNV uitgeverij,Utrecht,The
tation factors
thesis, Universiteit
paleo-ecologisch
onderzoek
archeologisch
Netherlands.
Bradshaw
M.Sc.
Be?rnem.
Belgium.
K. &
Deforce
f?rMitteleuropa und angrenzendeGebiete, 542 p.
Pfeil, M?nchen.
data. Quatern.
Sci. Rev. 22:
1701-1716.
pollen
en paleo-ecolo
Deforce
1997. ?Paleobotanisch
K.,
van een Pleistocene
afzetting in
gisch onderzoek
time.
Post-Glacial
Contributers,
from
Europe during theHolocene reconstructed
gurlagernderL?neburgerHeide. Flora 152: 325
370.
J.&
S., Stevenson
A.C., Guiot
?
2003.
The temperature of
Brewer
B.A.S.,
Data
excavation proj
and Doel
Ghent
1, Archaeological
Reports
Press, Gent,
3, pp. 108-126. Academia
Belgium.
Gro?te
V,
?
1977.
van Midden-
Pollenanalytisch
en Boven Pleistocene
Vlaanderen,
Ph.D.
Gent, Belgium.
?
Jong J., 1986.
onderzoek
in
afzettingen
Universiteit
Gent,
thesis,
in verband
Onderzoek
met
het
voorkomen
van Pinus
Baarland.
Rapport 991, Rijks Geologische
Dienst.
De
Verrebroek
Jong
J., 1987.
bepalingen
?
hout
Uitkomsten
in het Hollandveen
van C14-ouderdoms
aan Hollandveen
bij Baarland.
Rap
port 991a, Rijks Geologische Dienst.
Delahunty
of the yew
?
war, and changing
Religion,
an historical
account
and ecological
tree (Taxus baccata L).
Ph.D.
thesis,
J., 2002.
landscapes:
bij
University of Florida,Gainesville, USA.
Dempsey
-
D. & Hook
chemical
I., 2000.
?
Yew
and morphological
(Taxus)
variations.
species
Pharm.
Biol. 38: 274-280.
Denys L. & Verbruggen C, 1989.? A case of
drowning The end of Subatlanticpeat growth
and
related
palaeoenvironmental
changes
in the
lower Scheldt basin (Belgium) based on diatom
and pollen
analysis.
7-36.
This content downloaded on Thu, 10 Jan 2013 04:53:02 AM
All use subject to JSTOR Terms and Conditions
Rev. Palaeobot.
Palyno.
59:
HISTORY OF TAXUS ACCATA INBELGIUM 233
DuviGNEAUD
?
J., 1965.
tion:
le Franc
Un
Bois
H.,
Werner
&
D?ll
H.E.,
Paulissen
Wirth
R,
?
1991.
D.,
erte der Gefasspflanzen
18: 1-122.
V.,
Zeigerw
Scripta
Mitteleuropas.
Baeteman
A.,
hollevoet
?
1999.
sion,
Demiddele
C,
., PlETERS
because
occupation
or the cause of a transgression?
on
the
events
and
human
interaction
H.,
J. et al.,
of a regres
schelvis
M.,
Human
review
und
a
critical
nacheiszeitliche
n?rdlich der
Waldgeschichte Mitteleuropas
Fischer,
Alpen: 256 p. Gustav
R. & Cutler
D., 2000. ?Plants
Gale
?
Garcia
Gilot
neuve.
Studia
Praehistorica
ven, Belgium.
?
., 1968.
Terneuzen
and buried
.
Clifford
post-glacial
history
., 1938.
of British
?
in southern Fenland.
Philos.
229: 323 -406.
Godwin
.,Godwin
M.E.
&
Fen near Ely. J. Ecol.
23: 509-535.
?
Greig
1996.
Great
Britain
J.,
B.E.,
zowa M.
& Wright
Birks
H.E.
England.
Ralska-Jasiewic
H.J.B.,
?
inEurope,
Chichester,
?
Universit?
Bel
L.
baccata
1996.
landscape
develop
in the Connemara
impact
Ireland. J. Biogeogr.
29: 153
?
baccata
i
Holocene
Western
P.E.,
Huybrechts
W.,
sediments
Natural
microsite,
L.):
of yew
regeneration
seed or herbivore
1999.
?
Post-pleniglacial
in central Belgium.
Geologica
29-37.
IszKULO
G.
BoRATYNSKi
&
2004.
A.,
In:
floodplain
Belg.
?
2:
Interaction
between canopy tree species and European yew
J?ger
baccata
Pol.
(Taxaceae).
E.J. & Werner
von Deutschland.
2002.
K,
Band
J. Ecol.
?
52:
523
Exkursionsflora
Kritis
4, Gef??pflanzen:
cher Band 9: 980 p. Rothmaler, Heidelberg Berlin.
Jahn
1991.
G.,
Europe.
perate
Janssens
?
deciduous
Temperate
In: Rohring
E. & Ulrich
deciduous
forests,
pp.
W.
&
Ferguson
D.K.,
forests
.
(eds.),
377-502.
1985.
?
of
Tem
Elsevier,
The
palaeo
ecology of theHolocene sedimentsatKallo, north
(eds.), Palaeoecological
pp. 15-76. Wiley,
Taxus
en
actuelle
Louvain-la-Neuve,
Amsterdam.
eventsduring the last 15 000 years: regional syn
theses of palaeoecological studies of lakes and
mires
Hulme
531.
., 1935.
Untersuchungen
Barlinda
human
and
Uplands,
165.
Taxus
Controlling factors in the formation of fen
deposits, as shownby peat investigationsatWood
Berglund
ment
T. Roy. Soc.
.
Clifford
2002.
C.C.,
of the
gin and stratigraphyof Fenland deposits near
of
Woodwalton, Hunts. II.Origin and stratigraphy
deposits
?
205.
Huang
I. Ori
Studies
vegetation.
Olden
limitation?/.Ecol. 84: 853-861.
forests of the
CambridgeUniversityPress, Cambridge.
Mooren.
la v?g?tation
Ph.D.
thesis,
de Louvain,
2001.
(Taxus
East Anglian fenland.New Phytol. 67: 733-738.
Godwin
., 1975.? The historyof theBritishFlora. A
factual basisfor phytogeography,2ndedn.: 383 p.
.&
occidentale.
D.,
7, Li?ge-Leu
Belgica
thousand
Moreog Romsdal
p? veg ut? Blyttia 59: 197
4-5:
Godwin
Godwin
Holtan
et
r?cents
polliniques
catholique
nurse plants in a threatened tree,
Taxus baccata:
local effects and landscape
level
26: 739-750.
consistency. Ecography
?
Index g?n?ral
des dates Lv. Labora
E., 1997.
toire du
14 de Louvain/Louvain-la
carbone
tree: a
In: Jankuhn H.
Aus
(ed.), Neue
lungst?tigkeit.
inNiedersachsen
und Forschungen
3,
grabungen
Lax, Hildesheim,
pp. 280-307.
August
Germany.
?
entre les spectres
Heim
Les
relations
J., 1970.
gium.
bivore-mediated
Germany.
The yew
zum Verlauf des Niederschlagklimas und seiner
Verkn?pfung mit der menschlichen Sied
Europe
commun
L'if
Ein
Mitteleuropas.
cata
in oldenburgischen
L.)
59: 51-67.
burger Jahrbuch
?
Hayen
1966.
Moorbotanische
H.,
Jena, Germany.
in archaeology:
en Belgique.
Les
113-132.
Belges
?
D. & Obeso
Facilitation
J.R., 2003.
by her
1979.
D.,
Naturalistes
H?lzer
320 p. Hulogosi,
Oregon.
?
1967.
and pollen preser
Palynology
2: 81-98.
vation. Rev. Palaeobot.
Palyno.
?
Hayen
Vorkommen
der Eibe
., 1960.
(Taxus bac
512 p.Westbury Publishing,Kew, U.K.
Galoux
Die
whispers:
Havtnga
A.J.,
geological
in the Belgian
occupation
26: 97-121.
?
Sp?t-
Nordseegebiet
1949.
F.,
?
Remagen,
?
., Jr., 1991.
Hartzell
between
coastal plain during the firstmillennium AD.
Probleme der K?stenforschung im s?dlichen
Firbas
1977.
D.,
Lehratlas: 217 p. Verlag
mikrophotographischer
Naturalistes
Geobot.
Ervynck
Grosser
Dr. Kessel,
Weber
W.
Les
Lompret.
Belges IQ: 441-461.
Ellenberg
de destruc
site menac?
U.K.
Jessen
?
ern Belgium.
Rev. Palaeobot.
Palyno.
S.Th. & Farrington
,Andersen
way,
46:
81-95.
A.,
1959.
The interglacialdeposit near Gort, Co. Gal
Ireland. Proc.
This content downloaded on Thu, 10 Jan 2013 04:53:02 AM
All use subject to JSTOR Terms and Conditions
Roy.
Irish. Acad.
60:
1-77.
BELGIAN JOURNALOF BOTANY 140
234
?
Flora
., 2000.
Polygonaceae.
JoNSELL
aceae
vol
Nordica,
1. Lycopodi
The Bergius
holm.
1964.
Yew
(Taxus
baccata)
poisoning
in pheasants (Phasianus colchicus). Tijdschr.
89:
187-188.
lands over
Klinck
1999.
B.,
J. Life Sei. Roy. Dublin
limestone.
?
De
van
samenstelling
Kr?l
Maes
Belgium.
?
S., 1993.
The
present-day
of yew-tree
reserve
Nienartowicz
(Taxus
A.
&
Yuchola
G.,
396 p. Paul
K?ster H.,
baccata
Boinski
value
1972. ?Handbuch
1994.?
Verwendung
R?VEKAMP
Ch., Van
Inheemse
bomen
en
O., Deforce
K.,
?
P. et ai, 2006.
en
in Nederland
struiken
B.E. & Don
J.,Andersen
S.T., Berglund
?
J.J., 1974.
Quaternary
stratigraphy of Nor
for terminology
and classifica
den, a proposal
tion. Boreas
3: 109-128.
?
van de
Merckx
1996.
V,
Landschapsreconstructie
De
Site Waasmunster-Pontrave.
Gallo-Romeinse
Mangerud
J., De
Duvigneaud
Groothertogdom
Miller
(eds.),
conservation
LWF
in der
and present:
?
S.B.J.,
1878.
601
p. Leach
und
vor-
Bericht
Landesanstalt
G. &
The Fen
and
Son,
Mitchell
F.J.G.,
und
10,
de Belgique.
g?n?rale
170 p. Minist?re
de l'A
griculture JardinBotanique de l'?tat,Bruxelles.
Verbruggen
Belgium.
?
1986.
C,
Palynolo
van een veenprofiel
uit het Doel
gisch onderzoek
van de Archeologische
dok te Doel.
Bijdragen
Dienst Waasland
1: 201-208.
die
f?r
Gent, Gent,
Minnaert
Bradshaw
O'connell
?
1996.
H.J.B.,
R.H.W.,
PlLCHER
M.,
Ireland.
In:
Hannon
J.R. &
G.E.,
watts
Berglund
W.A.,
.E., Birks
M & Wright
Ralska-Jasiewiczowa
events during
(eds.), Palaeoecological
H.E.
the last 15
000 years: regional synthesesofpalaeoecologi
inEurope,
of lakes and mires
cal studies
Wiley, Chichester,U.K.
Molloy
K. & O'Connell
etation
M.,
2004.
?
pp.
Holocene
1-14.
veg
in the karstic envi
dynamics
Ire
of Inis Oirr, Arran
Islands, western
and
ronment
land-use
land: pollen analytical evidence evaluated in the
light of the archaeological
record.
113: 41-64.
42: 7-10.
1:
land past
Archae
naar
1982. ?Palynologisch
onderzoek
G.,
en jysische
van de
oorzaken
antropogene
M.Sc.
thesis,
vorming van het Scheidealluvium.
Quart?re
Vegetationsgeschichte
Methoden
und Ergebnisse:
462 p. Gus
Europas.
tav Fisher, Jena
Stuttgart, Germany.
Lanting
WA.
& van Hinte
J.N.,
B.W., Casparie
?
1999.
the
J. Soc.
Bows
from
Netherlands.
R,
vol.
Skertchley
de Vlaamse
p?riode.
de
Belgium.
?
1994.
G.,
Spermatophytes,
en
Oudenburg
tijdens de Romeinse
39: 5-23.
Minnaert
Coperni
?
Flore
?
Wisbech.
de aangrenzendegebieden (Pteridofytenen Sper
Meise,
matofyten):1091p. Nationale Plantentuin,
Archer-Antiquaries
?
Lawalr?e
1952.
A.,
1958.
ologiaBelg.
S.H. &
M.
L. &
J.-E., Delvosalle
van Belgi?,
Flora
het
en
Luxemburg, Noord-Frankrijk
J., 1998.
J.,
Kustvlakte
Universiteit
Waidenentwicklung
in
Holzes
ihres
Langhe
Aardrijfakunde2: 45-52.
Mertens
in the
L.)
Die Stellung der Eibe
zur Eibe. Bayerische
Beitrage
Walt und Forstwirtschaft.
Lang
J., Brinkkemper
den Bremt
B., Bastiaens
der Nadelgeh?lze:
fr?hgeschichtlicher Zeit,
Lambinon
op
van
of the
Parey, Berlin.
nacheiszeitlichen
in
het bos
condition
natural
forests,
future, pp. 69-78. Nicholas
problems
cus University
,Poland.
Press,
Kr?ssmann
Bakker,
verspreiding, geschiede
Herkenning,
nis en gebruik:
376 p. Boom, Amsterdam.
im
L.
Staropolskie
inWierzchlas. In: Rejewski
Cisy
Wycz?lkowskiego
M.,
P.W.,
Ned
Vlaanderen.
afzettingen
thesis, Universiteit
Gent, Gent,
M.Sc.
population
BrOEKE
?
(eds.), 2005.
842 p. Bert
in de prehistorie:
erland
Soc.
onderzoek
paleoecologisch
van de
houtanalyse
van
basis
van Gijn A.
r?sul
premiers
ner
het laat-Eemiaam.
Be?rnem.
et
Amsterdam.
Philos.
T. Roy. Soc.
247: 533-592.
Birmingham.
?
.& Kirby E.N.,
Kelly
1982.
Irish native wood
3: 181-198.
H. &
Fokkens
and northern Belgium.
Ph.D.
thesis, Vrije Univer
The Netherlands.
siteit, Amsterdam,
?
The Middle
of North
1964.
Pleistocene
M.R.,
Kelly
?
s?dimentaires
la Meuse
(Ardennes,
France):
tats. Quaternaire
4: 17-30.
van
den
louwe
kooijmans
L.P.,
tidal and fluvi
Early-Pleistocene
environments
in the southern Netherlands
atile
environnements
J., 1993.
biologiques ? Holoc?ne dans la plaine alluviale
de
?
Diergeneesk.
?
Kasse
., 1988.
E. & Mouthon
J.,Gilot
des
?volution
Foundation,
The Royal SwedishAcademy of Sciences, Stock
Jordan W.J.,
D., Heim
Lefevre
Moore
P.D., Webb
JA. &
Collinson
Quatern.
M.E.,
1991.
Int.
?
Textbook of pollen analysis, 2nd edn: 216 p.
Blackwell,
Oxford.
This content downloaded on Thu, 10 Jan 2013 04:53:02 AM
All use subject to JSTOR Terms and Conditions
HISTORY OF TAXUS BACCATA INBELGIUM 23 5
.&
MossBERG
Stenberg
?
2003.
L.,
Den
nya
Peglar
nordiskafloran: 928 p.Wahlstr?m & Widstrand,
Sweden.
Muhle
?
1979.
O.,
von
R?ckgang
Eiben-Waldge
sellschaftenund M?glichkeiten ihrerErhaltung.
In: Wilmanns
T?xen
O. &
R.
von Pflanzengesellschaften,
pp. 483
Vergehen
501. J. Cramer, Vaduz.
?
? l'?tude paly
Mullenders
1960.
Contribution
W,
nologique des tourbi?resde laBar (Departement
Belg. 94: 163-175.
Munaut
A.-V,
Lovaniensia,
Geogr.
Press,
A.V.,
gium.
Pons
A. & 0STBYE
olus
capreolus)
1995.
E.,
A. & 0stbye
olus
E., 2004.
Roe
(Taxus
reserves
Navys E., 2000.?
Roe
deer
(Capre
affects yew
120:
Conserv.
Biol.
and
States
Reimer
Noryskiewicz
A.M.,
tion in the Taxus
?
2003.
Modern
pollen
in theWierzchlas
reserve
O'Connell
M.
& Molloy
Biology
and Environment:
Proc.
IrishAcad. 101B: 99-128.
O'Connell
M.,
Molloy
K.
Post-glacial
landscape
western
with
Ireland
woodland
Kaland
scape
-
and
Farming
dynamics in Ireland during the
woodland
Neolithic.
?
2001.
K.,
history.
P.E. & Moe
past,
&
Bowler
evolution
M.,
present
The
(eds.),
1988.
and future,
pp.
to
land
487-514.
CambridgeUniversityPress, Cambridge.
Palese
R. & Aeschimann
de Gaston
pays
Bonnier.
voisins,
D.,
1990. ?La
France,
vol. 4. Belin,
Suisse,
Paris.
2002.
?
Bard
E.,
et al,
C.J.H.
Bayliss
?
A.,
Int
2004.
de
?
?
a?ropalynologique
sur la
facteurs climatiques
Contribution
des
l'action
floraison de l'orme (Ulmus campestris) et de l'if
(Taxus
R. &
pollen de Taxus
ismes du sol. Nat.
Saintenoy-Simon
et Spores 27: 53-94.
?
du
1977.
G.,
D?gradation
L. par les microorgan
baccata
Pollen
baccata).
Kilbertus
Can.
?
104:
Ben-Ahin
377-382
L'if, Taxus
J., 1983.
27:
(Huy). Dumortiera
J. (with collaboration
Saintenoy-Simon
L.-M.,
Dufr?ne
M.,
baccata
L.,
?
37-38.
of Barbier
Gathoye
Y.,
J.-L.
& Vert?
H.J.B.,
cultural
T.D.,
?
Premi?re
liste des esp?ces
P.), 2006.
et prot?g?es
de la R?gion Wal
rares, menac?es
?
reference
Birks
P., 1985.
Delescaille
in Connemara,
particular
In: Birks H.H.,
D.
Roy.
J.W., Bertrand
l'?tude
Rohr
(north
ernPoland). 16thINQUA Congres,Reno.
& Dines
D.A.
sity Press, Oxford.
P.J., Baillie
M.G.L.,
Richard
for
deposi
Pearman
Cal04 Terrestrialradiocarbonage calibration,26 0 ka BP. Radiocarbon 46: 1029-1058.
itsdistinction fromLithuania. Baltic Forestry 6:
41-46
and con
dynamics
18. Kluwer,
Dor
Geobotany
pp.7-79.
CD.,
Beck
545-548.
reasons
the main
bogs
nutrient
history,
In: Verhoeven
in the Netherlands:
and
(ed.), Fens
in the
formation
New atlas of theBritish and Irishflora. An atlas
of thevascular plants ofBritain, Ireland, theIsle
ofMan and theChannel Islands. OxfordUniver
nature
within
Holocene
peat
of the Netherlands.
drecht.
(Capre
in
baccata
English yew (Taxus baccata L.) in
of Baltic
forests
?
browsing
recruitment
baccata)
inNorway.
parts
servation,
deer
pressure
capreolus)
lower
?
Vegetation,
relation to bilberry(Vacciniummyrtillus)density
and snow depth.WildlifeBiol. 1: 294-253.
Mysterud
1992.
Preston
?
on yew Taxus
feeding
L.J.,
J.T.A.
17: 7-27.
Bodemonderzoek
Pollen
thesis,Vrije Universiteit Br?ssel, Br?ssel, Bel
gisement tourbeuxsitu? ? Terneuzen (Pays Bas).
Berichten van de Rijfadienst voorOudheidkundig
Mysterud
vege
analytic studies of theHol
in the Izenberghe
M.Sc.
area, Belgium.
stenian
d'un
pal?o-?cologique
late-Holocene
10: 219-225.
Archaeobot.
?
J., 1977.
Ponniah
Leuven,
Belgium.
?
1967b.
?tude
and
of laKarelsl? (Waldbillig, easternGutland). Veg.
Recherches
pal?o
etMoyenne
Acta
Belgique.
vol. 6. Leuven
University
en Basse
?cologiques
Munaut
?
1967a.
annual
a year-by
The
laminations.
U.K.
interpreted using recurrent groups of taxa. Veg.
2: 15-28.
Hist. Archaeobot.
?
Pernaud
J.-M., 2001.
history
Postglacial
vegetation
new charcoal data from the cave
in Luxembourg:
Hist.
I.-La
belges.
Soc. Roy. Bot.
dans
les Ardennes
palynologiques
Bull
tourbi?re du Grand Passage.
Ulmus
mid-Holocene
Norfolk,
tationhistoryof Quidenham Mere, Norfolk,UK
et Spores 2: 43-55.
?
1962.
Recherches
C,
Pollen
des Ardennes).
W. & Knop
Mullenders
The
Mere,
year stratigraphy from
Holocene
3: 1-13.
?
Peglar
Mid1993b.
S.M.,
und
(eds.), Werden
at Diss
decline
?
1993a.
S.M.,
grande flore
et
Belgique
lonne (Pt?ridophytes et Spermatophytes).Ver
sioni (7/3/2006), available at http://rnrw.wal
lonie.be/dgrne/sibw/especes/ecologie/plantes/list
aspx?ID=5 94
erouge/fiche2.
Sarmaja-Korjonen
A.,
1991.
?
K., Vasari
Taxus
baccata
Y. & Haeggstr?m
and
influence
C
of Iron
Age man on thevegetation inAland, SW Finland.
Ann. Bot. Fennici
This content downloaded on Thu, 10 Jan 2013 04:53:02 AM
All use subject to JSTOR Terms and Conditions
28:
143-159.
BELGIAN JOURNALOF BOTANY 140
236
Schulte
1975.
T.,
?
Lethal
tree (Taxus
yew
158.
schweingr?ber
1990.
F.H.,
with
intoxication
Arch.
baccata).
?
leaves
Toxikol.
Anatomy
of
Van
153
34:
?
of European
and Landscape
land.
Sier M.M.,
veen;
een
Borsele,
bewoningsgeschiedenis
in het
opgraving
uit de Romeinse
?
K., 2004.
saurus project.
noemen:
Namen
1,1AP-Rapporten
CAI-rapport
J., Paepe R, Baeteman
C, Beyens
L., Cunat
?
R. et al,
1978.
La formation d'
N., Geeraerts
un nouveau
Herzeele:
stratotype du Pleistoc?ne
main
la
de
Mer
du Nord. Bull. Assoc. Fr.
moyen
Stace C, 1997.? New flora of theBritish Isles: 1165
p. CambridgeUniversityPress, Cambridge.
Nederland.
Stockmans
Deel
1945.
F.,
de
I. Dekker
?
&
Mededel.
Huis,
Wildervank.
Hout.
In: Sier M.M.
(ed.),
een opgraving
in het veen; bewonings
uit de Romeinse
tijd, pp. 51-61.
geschiedenis
d'Heusden-Lez-Gand
Konink.
Rijn
(Bel
Museum
Natuurhist.
F. & Vanhoorne
Konink.
M.
Stuiver
&
R.
Van
Belgium.
Smeerdijk
and revised
CALIB
radiocarbon
calibra
35: 215-230.
?
1991.
Localities
ern part of eastern Norway
J.-C. & Mag?rd
Van
including Aust-Agder.
E.,
and conservation
Biol.
P.A. &
Ecol.
Conserv.
Polwart
Zeist
Taxus
88:
A.,
1980.?
baccata
L.
1999.
tourbi?re
edel.
T.G.,
D.H.,
Heywood
Walters
Population
status of the last natural
173-182.
?
2003.
Taxus
Cambridge.
onderzoek.
Palynologisch
A
91 p.
Stichting
palaeobotanical
Kri
of
study
Biol.
V.H.,
S.M.
& Webb
N.A.,
D.A.,
R,
1951.
alluviale
gique).
Mededel.
Verbruggen
C,
?
Evolution
d'une
Med
21:
tourbi?re
au Kruisschans
Konink.
27:
1971.
geschiedenis
sis, Universiteit
?
de
Bel
(Anvers,
Inst. Natuur
Belg.
1-20.
van Zandig
landschaps
Ph.D.
the
Postglaciale
Vlaanderen.
Gent, Gent,
Belgium.
?
L. & Kiden
P., 1996.
Denys
gium. In: Berglund
Jasiewiczowa
M.
B.E.,
&
Birks
H.J.B.,
Wright
H.E.
Bel
Ralska
(eds.),
Palaeoecological events during the last 15 000
years: regional syntheses of palaeoecological
studies
of lakes and mires
Wiley, Chichester.
Con
D.,
1986.
?
inEurope,
Historical
and
pp. 553-574.
environmental
significanceof theyew (Taxus baccata L.). Israel
Valentine
1964.
(Belgique).
Inst. Natuurwetensch.
Belg.
plaine
Voliotis
Burges
?tude pollinique d'une
? Heusden-les-Gand
wetenschappen
Verbruggen
C,
Ecological historyofYew
in southern England.
158-180.
1945. ?
Konink.
Vanhoorne
L.
baccata
?
Flora Europaea, vol. 7, Cambridge University
Press,
Meise,
1-11.
?
serv. 17: 243-265.
Tutin
?
1964.
W.,
PalaeohistorialO:
91: 489-524.
Tittensor RM.,
?
Plantentuin,
naaldboomsoort:
Vanhoorne R,
population ofEnglish yew Taxus baccata inDen
mark.
Nationale
2003.
D.,
inheemse
wor
NINA Oppdragsmelding 64: 1-58. [InNorwegian]
Thomas
van
1979, ?Atlas
L.,
tischBosbeheer,Utrecht,The Netherlands.
14C
thyof conservationforyew and holly in thewest
ecology
E. & Delvosalle
schoten, The Netherlands.
?
van Vuure
taxus (Taxus baccata L):
De
T., 1990.
en bevordering
van een
bescherming
ekologie,
130.
Inst. Natuurwetensch.
?
Extended
P.J., 1993.
tion program. Radiocarbon
H. & Holand
Svalastog
K.,
Svenning
Sier
In: SierM.M. (ed.),Ellewoutsdijk inde Romeinse
tijd,pp. 148-166,ADC rapport200, ADC, Bun
of
(with collaboration
Belg.
Reimer,
database
In:
houtonderzoek.
some bogs inwesternBrittany(Finist?re),France.
?
Vanden Berghen C), 1954.
?tude botaniquedu
gisementde tourbede la r?gionde Pervijze. Ver
handel.
Het
en Spermatofyten.
Belgi? 21: 1-7.
Stockmans
?
2003.
P.,
de Belgische en Luxemburgseflora. Pter?dofyten
et feuilles
branchettes
Graines,
la tourbe holoc?ne
gique).
Rijn
The Netherlands.
?
van kienhout.
De wording
en de vindplaatsen
van kienhout. Uit
1856. De bodem van
treksel uit Staring W.C.H.,
1983.
ontstaan
Het
Plantentuin,
Belgium.
?
P., 2001.
van Rompaey
1-3: 81-149.
Quatern.
W.C.H.,
en het
M.M. (ed.),Ellewoutsdijk inde Romeinse tijd,pp.
104-138. ADC rapport200, ADC, Bunschoten,
14:
49-54.
Staring
2006.
D.,
Vlaanderen
1008 p. Nationale
Gewest:
lands.
Van
het CAI-the
Somm?
?tude
van
Beer
ADC rapport76.ADC, Bunschoten, The Nether
tijd.ADC rapport76.ADC, Bunschoten.
Slechten
De
Borsele,
?
2001.
E. &
den
L., Van
L, Vanhecke
P., Vercruysse
van de Flora
Atlas
Meise,
Van
Switzer
Bern,
Research,
Hoste
W.,
Bremt
Brussels
woods: 800 p. Swiss Federal InstituteforForest,
Snow
Landuyt
J. Bot.
Vos
35: 47-52.
P.C. & Van Heertngen
RM.,
1997.
?
Holocene
geology and occupationhistoryof theProvince of
This content downloaded on Thu, 10 Jan 2013 04:53:02 AM
All use subject to JSTOR Terms and Conditions
HISTORY OF TAXUS BACCATA INBELGIUM
Zeeland (SW Netherlands).Mededel Ned. Inst.
Wetensch.
TNO 59. 5-109.
Toegepaste
?
Watts
1985.
W.A.,
Quaternary
vegetation
cycles.
In: Edwards
K.J. & Warren
W.P.
(eds.), The
Quaternary historyof Ireland, pp. 154-185.Aca
demic
Weeda
Press,
London.
Ch. & Westra
T.,
E.J., Westra
R., Westra
?
1985.
Nederlandse
Wilde
oecologische
flora.
planten en hun relaties 1. IVN in collaboration
withVARA andVEWIN, s.l.
West
R.G.,
Hoxnian
1962.
?
A
Interglacial.
note
New
on Taxus
Phytol.
in the
pollen
61:
189-190.
WoiLLARD
G.,
237
?
The
last interglacial-glacial
France. Bull.
pile inNortheastern
G?ol. 88: 51-69.
?
Sea-level
in The
1983.
changes
1979.
at Grande
cycle
Soc. Belge
Zagwijn
W.H.,
Netherlands during the Eemian, Geologie en
Mijnbouw 62: 437-450.
Zagwijn
1992.
?
of vegetation
dur
Migration
in Europe.
Courier Forsch.
ing the Quaternary
Inst. Senckenberg
153: 9-20.
?
L. In: Markgraf
F. (ed.),
Zoller
1981.
Taxus
H.,
von Mitteleuropa,
Gustav Hegt
Illustrierte Flora
W.H.,
Band 1, Teil 2, pp. 127-134.Verlag Parey,Berlin.
This content downloaded on Thu, 10 Jan 2013 04:53:02 AM
All use subject to JSTOR Terms and Conditions