The Holocene History of Taxus Baccata (Yew) in Belgium and Neighbouring Regions

R o y a l Botanical Society Belgium

THE HOLOCENE HISTORY OF TAXUS BACCATA (YEW) IN BELGIUM AND NEIGHBOURING REGIONS Author(s): Koen Deforce and Tan Bastiaens Reviewed work(s): Source: Belgian Journal of Botany, Vol. 140, No. 2 (2007), pp. 222-237 Published by: Royal Botanical Society of Belgium Stable URL: http://www.jstor.org/stable/20794641 Accessed: 10/01/2013 04:53

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THE HOLOCENE HISTORY OF TAXUS BACCATA (YEW) IN BELGIUM AND NEIGHBOURING REGIONS

Koen Deforce* and Jan Bastiaens Flemish Heritage Institute, Koning Albert II-Laan 19 bus 5, B -1210 Brussels, Belgium (* Author for correspondence; e-mail: [email protected] ) Received 5 September 2006; accepted 24 November 2006.

A bstract . — The current natural distribution of Taxus baccata L. in Belgium is limited to 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 the world. In Flanders, the northern part of Belgium, Taxus is considered not to be native and Taxus stands are interpreted there as being planted by humans or as garden escapes. The Holocene pollen and macrofossil data for Taxus, however, show a very different picture regarding abundance and geographical distribution, as well as habitat. It appears that during the Sub-boreal, Taxus grew in the coastal plain and the lower Scheldt valley, where it was 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 (raised) bogs. Belgium is not the only case where such observations have been made. In other 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 overview of the palaeobotanical finds ofTaxus baccata from Belgium is here given, supplemented with finds from neighbouring regions. The Holocene distribution and palaeo- ecology of Taxus baccata are discussed in a broader northwest European context.

K ey w ords . — Taxus baccata L., Belgium, Holocene, vegetation history, yew.

INTRODUCTION period characterised by a warm oceanic climate (Jessen et al. 1959,W est 1962, K elly 1964, During most of the Pleistocene interglacial G odwin 1975,W atts 1985). High percentages of periods, Taxus formed a much more important both Taxus pollen and macroremains have been element of the vegetation of northwestern Europe found at several northwest European sites where than during the Holocene (A verdieck 1971,G od Holsteinian sediments are preserved. High per win 1975,Z agwijn 1992,L ang 1994).Taxus has centages of Taxus pollen have also been found at been found in Belgium as early as the Tiglian-C5 several sites in Belgium, in sediments dating from interglacial period (ca. 2-1.8 million yrs BP; this period (D e G roote 1977,P onniah 1977, Pleistocene chronozones according to Lang Sommé et al. 1978). The most famous palaeo 1994)(K asse 1988) and seems to have had its botanical record of Taxus dating from the Holstein greatest expansion in northwestern Europe during interglacial, however, is a spear made ofTaxus the Holstein interglacial (400 - 367 ka BP), a wood found at Clacton (Essex, UK; G odwin

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1975), which is also one of the oldest known arte growing and long-living, reaching maturity only facts made of wood. at ca. 70 years. It is extremely shade tolerant but During the last interglacial period before the can withstand full exposure to the sun(T utin et Holocene, the Eemian (130 - 115 ka BP), Taxus al. 1964, Thomas & Polwart 2003). Taxus is also played a more important role in the vegeta normally dioecious, rarely monoecious and is tion in northwestern Europe than during the wind-pollinated (Tutin et al. 1964, Thomas & Holocene, although the pollen percentages are Polward 2003). It flowers from February to April much lower than during the Holstein interglacial (R ichard 1985). (B ehre 1962,A ndersen 1975,W oillard 1979, Taxus occurs throughout most of Europe and Lang 1994).Z agwijn (1983, 1992) even distin some parts of northern Africa, although its distri guished a Taxus subzone in his zonation of the bution is very scattered. Taxus thrives best in Eemian period in the Netherlands. In Belgium, regions with a mild, oceanic climate. Its distribu both pollen and wood of Taxus have been found tion is limited by low temperatures in Scandi in Eemian peat deposits at Beemem(D e Groote navia, a severe continental climate in eastern 1977,D eforce 1997,K linck 1999). Europe and aridity and high temperatures in During the present interglacial period, the Turkey and north Africa (Thomas & Polwart Holocene, Taxus seems to have played a less 2003). In the Mediterranean region, Taxus is con important role in the vegetation. However, during fined to the higher mountains (Tutin et al. 1964). the Sub-boreal (5 000 - 2 500 uncal. BP; Taxus does not form pure monospecies Holocene chronozones according to M angerud et stands (except in the Caucasus Mountains and on al. 1974), Taxus was more abundant and showed a chalk and limestone in England) but belongs to completely different distribution and ecology than diverse forest communities mainly composed of nowadays. The aim of this paper is to review the Abies, Fagus, Carpinus, Alnus and Picea (E llen - available data on the Holocene history of Taxus berg et al. 1991, Jahn 1991, Iszulo & B oratyn - and to discuss its past distribution and ecology. ski 2004). Furthermore, the existing hypotheses for the In Europe, most of the natural stands of Holocene Taxus decline will be evaluated in the Taxus grow on well-drained calcareous soils, light of the available palaeobotanical data. although the tree can grow on almost any soil, including silicious soils derived from igneous and sedimentary rocks (Thomas & Polwart 2003). In PRESENT-DAY ECOLOGY AND most countries, Taxus is a declining or even DISTRIBUTION threatened species. The reasons are thought to be deforestation, selective felling and grazing There is no scientific agreement on the exact (M uhle 1979,S venning & M agârd 1999, taxonomic position of the genus Taxus, which N avys 2000, H oltan 2001, Thomas & Polwart encompasses about seven closely related species 2003, M ysterud & 0 stbye 2004). Several pro scattered throughout the northern temperate tected areas have been established to conserve the region (V oliotis 1986,D empsey & H ook 2000, species (S valastog & H oland 1991,H artzell Thomas & Polward 2003). The species separa 1991,K ról 1993,B rande 2002), which also sur tion within the genus is equally disputed (van vives in a cultivated form as an ornamental tree in V uure 1990,D elahunty 2002, Thomas & Pol- parks, gardens, and cemeteries (K rüssm ann wart 2003), although recent genetic research 1972,S aintenoy -S imon 2006). (C ollins et al. 2003) shows that the current In Belgium, the current natural distribution species delimitations are well founded. of Taxus is limited to a few localities in the south Taxus baccata L. (subsequently referred to ern part of the country, namely the southern and as Taxus), the species native to Europe, is an ever western part of the Meuse district (Fig. 1; Lawal - green needle-leaved gymnosperm shrub or tree, rée 1952,D uvigneaud 1965,G aloux 1979,van growing up to 28 m high. The species is slow- R ompaey & D evosalle 1979,Saintenoy -S imon

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1983, 2006, Lambinon et al. 1998). At these petraea, Tilia platyphyllos and, sometimes, localities, Taxus predominantly grows on steep, Ulmus spp. andCorylus avellana (D uvigneaud calcareous slopes, which are believed to be its 1965,G aloux 1979). In Flanders, the northern natural habitat in the region (L ambinon et al. part of Belgium, Taxus is considered not to be 1998).Taxus grows there together with Fagus syl native and Taxus stands are interpreted there as vatica, Carpinus betulus, Quercus robur, Q. having been planted by humans or as escapes

THE NETHERLANDS

GERMANY

BELGIUM

FRANCE

0 - 5 m • potieri LUXEMBOUI 5 -200 m 4 seeds B 200-500 m ■ woocj/charcoai M + SOOm O current natural populations

F ig . 1. Map showing both the current natural distribution and subfossil finds of Taxus baccata in Belgium: white dots denote natural T. baccata populations (data fromL aw a l r ée 1952,v a n R o m p a e y & D e v o s a l l e 1979,S a in t e n o y -S im o n 2006, V a n L a n d u y t et al. 2006, and M a e s et al. 2006); black symbols denote Holocene palaeobotanical records of pollen, seeds and wood/charcoal ofTaxus. (1) Avekapelle 363 (B a e t e m a n & V e r b r u g g e n 1979); (2) Booitshoeke (B a e t e m a n & V e r b r u g g e n 1979); (3) Oudekapelle(S t o c k m a n s & V a n h o o r n e 1954); (4) Sint- Jacobs-Kapelle (S t o c k m a n s & V a n h o o r n e 1954); (5) Raversijde(D efo r c e & B a s t ia e n s , in press); (6) Leffinge (B a e t e m a n et al. 1981); (7) Wenduine(M er t e n s 1958); (8) Blankenbergse vaart - Zuid (A l l e m e e r sc h 1991); (9) Heusden (S t o c k m a n s 1945); (10) Laame - Damvallei(V e r b r u g g e n 1971); (11) Borsele(NL)(V a n R ijn 2001); (12) Ellewoutsdijk (NL)(V a n R ijn 2003, V a n S m e er d ijk 2003); (13) Baarland (NL)(D e Jo n g 1986); (14) Temeuzen (M u n a u t 1967a,b); (15) Waasmunster - Pontrave(M e r c k x 1996); (16) Weert(M in n a e r t 1982); (17) Verrebroek (D e fo r c e et al. 2005); (18) Doei (M in n a e r t & V e r b r u g g e n 1986);(D efo r c e et al. 2005); (19) Zand vliet (M u n a u t 1967a); (20) Oorderen(M u n a u t 1967a); (21) Kruisschans(V a n h o o r n e 1951); (22) La Karelslé (Waldbillig, eastern Gutland, Luxembourg) (P e r n a u d 2001).

This content downloaded on Thu, 10 Jan 2013 04:53:02 AM All use subject to JSTOR Terms and Conditions HISTORY OF TAXUS BACCATA IN BELGIUM 225 from gardens and parks (L ambinon et al. 1998, were present in the investigated peat sequence. M aes et al 2006). Similarly, pollen analysis of peat sequences In other parts of northwestern Europe, the from Kruisschans (Antwerp, Belgium; V a n natural distribution of Taxus is also mainly con hoorne 1951), Oudekapelle (Diksmuide, Bel fined to soils on limestone and other types of gium; S tockmans & V anhoorne 1954) and well-drained, base-rich soils (UK and Ireland: Sint-Jacobs-Kapelle (Diksmuide, Belgium; Tittensor 1980, Kelly & K irby 1982, Stace Stockmans & V anhoorne 1954) did not reveal 1997,P reston et al. 2002; The Netherlands: Taxus pollen during the palynological research, W eeda et al. 1985; Germany: Jäger & W erner while several Taxus seeds were recovered from 2002; France and Switzerland: Palese & A eschi - the same peat sequences investigated. m ann 1990; Scandinavia:Jonsell 2000, Moss- berg & Stenberg 2003). Holocene Taxuspollen records from Belgium and immediate surroundings PALAEOBOTANICAL RECORDS OF TAXUS Holocene records of Taxus pollen from Belgium are not very abundant. Out of a total of The pollen data 370 palynological studies from all over Belgium Characteristics of Taxuspollen that were reviewed (D eforce & B astiaens 2006), only 12 contained substantial records of Taxus pollen is spherical to obtusely angu Taxus. Sites where Taxus occurred in only one lar, its size ranging between 19.3 and 29.8 pm or a few samples, and with a frequency of less after acetolysis (A verdieck 1971,B eug 2004). than 1%, are not included in the overview pre Taxus pollen does not have sacci and is inapertu- sented here, as this might represent long-dis rate. The exine is intectate and has a scabrate or tance transport. On the other hand, it must be microgemmate sculpturing (M oore et al. 1991, remembered that in some of the older analyses, B eug 2004). Regarding identification, confusion Taxus pollen was very probably overlooked, as might be possible with pollen of Juniperus, Pop- already discussed above. Three additional ulus, Rhynchospora and Quercus (A verdieck records derived from the southwestern Nether 1971,Z oller 1981, M oore et al. 1991,B eug lands, close to the Belgian border, are included 2004). The pollen grains frequently split and are in the discussion. sensitive to corrosion (H avinga 1967, Together, the records show two remarkable A verdieck 1971, R ohr & K ilbertus 1977, characteristics: (1) they are all situated in the B radshaw 1978). Although Taxus is an coastal plain and the lower Scheldt valley (Fig. anemophilous tree, the pollen representation in 1), and (2) they can all be dated from the Sub- surface samples near Taxus stands seems to be boreal. Some of these dates only rely upon bio rather low and decreases sharply with increasing zonation as the interpretation of the older dia distance from the Taxus stand (H eim 1970, grams is not supported by radiocarbon dating. But N oryskiewicz 2003). These factors, in combina still, the available 14C dates (Table 1) allow the tion with the lack of distinctive features such as conclusion that Taxus appears in pollen diagrams sacci, apertures or a distinctive exine sculptur in Belgium and the southern Netherlands between ing, make it likely that Taxus pollen was not 4750 ± 140 uncal. BP (Oorderen) and 4280 ± 130 recognised in some of the earlier palynological uncal. BP (Temeuzen). At all sites Taxus percent studies (K üster 1994), or at sites where condi ages remain rather low, varying between 2 and tions for pollen preservation were poor. During 6%. The percentages are higher only at Zandvliet the palynological investigation of Wood Fen (10.1%) and Ellewoutsdijk (16%). Taxus disap (Ely, UK; G odw in et al. 1935), for example, no pears from the pollen diagrams between 4035 ± Taxus pollen was found while several trunks and 30 uncal. BP (Raversijde) and 3510 ± 45 uncal. even the pollen-bearing eone scales of Taxus BP (Baarland) (Table 1).

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Table 1. Radiocarbon dates of the start and end of the Taxus curve in Holocene pollen diagrams from Belgium and the southern Netherlands. Radiocarbon dates are calibrated using calib 5.0.2 (S tuiver & R eimer 1993) and the INTCAL04 calibration data set (R eimer et al. 2004). 4 •8 la ■g to toi O 0 1 ”3 .SP > Q o 00 to ^ H -H -H £ to -H p i?

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Holocene passestheintestinal canal undamaged id ad ml mmas wie h se itself seed mammals, small the while and birds Nevertheless,all findscan be attributed to the Late toxic part of Atlantic or Sub-boreal on the basis of their position radiocarbon dated by analysis. been havewhich have all been recovered from peat deposits, none of region that yielded subfossil pollen,i.e. the coastal bothalong the riverScheldt (Fig. 1). Kruisschans at and area, coastalwestern conspicuous reddish arii.This arii is the only non theyarelarge (6-8 mm) andellipsoid-ovoid with hoorne Kapelle ans m W hoorne (Z winter the into September from occurs which al. nua o bcne i scin ter ufc is surface their section; in biconvex orangular lsy ri os o peev, palaeobotanical preserve, not does arii fleshy T inthe lower part of theso-called surface peat (see area and thelower Scheldt valley.The sal finds of pce. hn rs, ed ae surrounded a arespecies. seeds fresh,by When taperinga upperend, androunded slightlyto tri elasticand resistant todecay silseeds have been found at Oudekapelle seeds of ute) o fple analysispollen thefurther),of carried on out or immediate surroundings smooth. Seeds cannotmistakenotheranybefor ae et sequences peat same Characteristics of Taxus seeds Characteristics f o he oller ood 1985). Birds are the main agent of seed disper

(Z seed These finds originate from the same restricted of records Holocene of number small A Theseeds of The wood of of wood The V & oller

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(S Taxus 1951, 1951) andHeusden Taxus

Taxus data 1981, tockmans

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anhoorne wood and charcoal anhoorne s ey es, hard, dense, very is V & t al. et P & (Z (S tockmans oller anhoorne 00. s the As 2000). olwart

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2003), 1945), 1954). seeds tock (V V The n a n a

HISTORY OF TAXUS BACCATA IN BELGIUM 227

sapwood is white to yellowish; the heartwood is time gap between the time of the construction and red and colours orange-brown after contact with the much older age of part of the construction the air. Owing to its good elasticity, it has been a wood is that at these sites during Roman times, very popular timber for the production of tools subfossil wood was used for the construction of and weapons, in particular bows (Z oller 1981, buildings (V an R ijn 2003). As inferred from the Lanting et al. 1999,G ale & Cutler 2000). pollen analysis of Ellewoutsdijk (V a n Smeerdijk Taxus wood and charcoal are easy to differentiate 2003) and the palaeogeographical map of the from that of other European gymnosperms, on region (Vos & V an H eeringen 1997), these sites account of the distinct spiral thickenings in the were situated in an almost treeless peat-bog envi tracheid walls and the absence of resin canals in ronment during the Roman Age, which might the former (G rosser 1977, S chweingrüber explain the use of subfossil material. 1990,G ale & Cutler 2000). The Taxus wood found at the Roman site of Wenduine (M ertens 1958) has not been dated but Holocene Taxus wood and charcoal records from it might represent subfossil wood too, given that Belgium and immediate surroundings Wenduine was also situated in a peat bog or a peri-marine environment during Roman Age Subfossil Holocene wood remains of Taxus (A llemeersch 1991,E r v y n c k ^ a /. 1999). from Belgium are only known from Blanken Charcoal from Taxus has been found in La berge (A llemeersch 1991) and Wenduine Karelslé, eastern Gutland (Luxemburg) (Fig. 1), (M ertens 1958). At Blankenberge, Taxus wood in archaeological cave deposits. A few fragments was found embedded in a peat deposit dating derive from deposits dating from the Middle from the late Atlantic or the Sub-boreal. At Wen Neolithic (4 500 - 3 500 BC; which corresponds duine, a fragment of Taxus wood was found in an to the Late Atlantic) while rather large amounts archaeological site dating from the Roman period have been recovered from Late Bronze Age (57 BC - 402 AD; archaeological periods accord deposits (1 100 - 800 BC, corresponding to the ing to Slechten 2004). Late Sub-boreal) (Pernaud 2001). This is rather In the southern part of the Netherlands, sub surprising as there is no evidence for the exten fossil wood of Taxus has been found at Temeuzen sion or even presence ofTaxus in any of the (M unaut 1967a,b), Borsele(V an R ijn 2001) and Holocene palynological records from Luxemburg Ellewoutsdijk (V an R ijn 2003). At Temeuzen, (Pernaud 2001), eastern Belgium and the Gaume several Taxus stems have been recovered from district (C outeaux 1969a,b), the Plateau des peat deposits but they have not been dated and Tailles area (M ullenders & Knop 1962) or the their Stratigraphie position has not been recorded. French Ardennes (M ullenders 1960,L efevre et However, it is very likely that the stems derive al 1993). from the same levels from which Taxus pollen All dated finds of Holocene Taxus wood and was recovered, which would place the wood frag charcoal are of Late Atlantic or Sub-boreal age ments in the Sub-boreal (M unaut 1967a,G od and, except for the charcoal from Gutland, all win 1968). At Borsele and Ellewoutsdijk, the finds were excavated in the Belgian coastal plain Taxus finds were partly preserved in the peaty soil or the Scheldt estuary. and consisted of wooden poles that were used as parts of Roman Age buildings. One Taxus pole Holocene palaeobotanical records of Ta x u s from Borsele was radiocarbon-dated at 4690 ± 60 FROM OTHER PARTS OF NORTHWESTERN EUROPE uncal. BP (several poles made from Pinus sylvestris gave similar dates) (S ier 2001). The Probably one of the earliest mentions of Taxus poles from Ellewoutsdijk were not dated by Taxus occurring in peat deposits was made by radiocarbon analysis but a pole from P. sylvestris Staring (1983, re-edition from 1856). The from the same buildings was dated at 4480 ± 25 author expressed his surprise about the presence uncal. BP. The only possible explanation for the of subfossil Taxus wood in Dutch peat deposits,

This content downloaded on Thu, 10 Jan 2013 04:53:02 AM All use subject to JSTOR Terms and Conditions 228 BELGIAN JOURNAL OF BOTANY 140 in contrast with the 19th century distribution of palaeobotanical records from Belgium and the Taxus in The Netherlands and elsewhere in southern Netherlands can be dated from the Sub- Europe. That Taxus grew on peat has been further boreal as well. Not only in Belgium, but also in demonstrated by the finds of subfossil wood at other parts of northwestern Europe (see 3.4), Ely (U K ), described by Miller and Skertchley Taxus shows a maximum in pollen diagrams dur (1878) and discussed by G odwin et al. (1935). ing the Sub-boreal. Another early observation of Taxus trunks recov ered from peat deposits was made at Ballyfin Distribution and habitat of Ta x u s d u rin g th e Bog (Ireland) by A dam s (1905), who stated that S u b -b o r e a l similar finds were so plentiful in former times Most of the finds of subfossil pollen, seeds that farmers in the neighbourhood used the wood and wood of Taxus in Belgium and the southern for gate posts, house roofs, etc. part of the Netherlands are situated in the coastal Firbas (1949) reviewed Holocene records of lowlands and more precisely in the area where the Taxus wood from Germany, both from natural so-called ‘surface peat’ or ‘Holland peat’ occurs peat deposits and from archaeological contexts in the subsoil. This surface peat was formed dur including several trunks from peat deposits from ing the mid Holocene when the postglacial sea- the coastal lowlands of Ostfriesland. Other finds level rise began to slow down and coastal barriers of Taxus wood recovered from peat deposits in could develop (B aeteman 1981, 1999, Vos & Germany are listed by H ayen (1960, 1966) and V a n H eeringen 1997). These coastal barriers A verdieck (1971). closed the coast almost completely and initiated Next to these finds of subfossil wood, there mire development. At the time of the maximal are also numerous pollen diagrams from north expansion of the peat, in the Sub-boreal, the mires western Europe, showing a distinct Taxus curve, stretched nearly continuously from Calais in mostly during the Sub-boreal (for northwestern northwestern France to southwestern Denmark, Germany, see A verdieck 1971, 1983,H ayen including the coastal plain of Belgium, the west 1960; for Ireland: O ’Connell et al. 1988, ern part of the Netherlands and the lower Scheldt M itchell et al. 1996,M olloy & O ’Connell valley (Pons 1992). 2004; for England: G odwin 1975, Peglar According to Pons (1992), this surface peat 1993a,b,G reig 1996,B atchelor et al. 2004; for shows more or less the same general development northwestern France: V a n Zeist 1964; for Swe in the coastal plain in Flanders and the southwest den: B erglund 1966, for Finland: Sarmaja - ern part of the Netherlands. On the salt marshes of K orjonen et al. 1991). the regression surface brackish fens developed, forming Phragmites (-Scirpus) peat. Gradually, désalinisation and decreasing amounts of avail DISCUSSION able nutrients resulted in Carex-Phragmites fens, which changed into mesotrophic Betula-Alnus From the Holocene palaeobotanical records carr, sometimes with some Pinus. This phase of of Taxus presented, it is clear that this tree was carr peat is followed by a transition to Sphagnum more abundant and had a different distribution peat and, in most places, the development of during the Sub-boreal in northwestern Europe raised bogs resulting in the formation of Sphag- compared to nowadays. «wm-Ericaceae peat. Peat growth ended because of Taxus occurs sporadically in pollen diagrams marine transgressions and fluvial sedimentation from England (B irks 1982, G odwin 1975), Ire between the Late Sub-boreal and the Late Middle land (H uang 2002) and Sweden (B erglund Ages, depending on the location, which resulted in 1966) from the Late Boreal or early Atlantic the covering of the peat with marine and alluvial onwards. For Belgium, the earliest post-glacial clay deposits (Janssens & Ferguson 1985, palaeobotanical records of Taxus date from the D enys & V erbruggen 1989,A llemeersch 1991, late Atlantic or early Sub-boreal. All the other

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Pons 1992). The upper part of the surface peat isoccurrence upon peat land, although natural com often lacking as a consequence of marine erosion munities of this kind can no longer be pointed or medieval and post-medieval peat extraction out” (G odwin 1968: 737). (B aeteman et al. 2002, B aeteman 2005). It has to be stressed here that palaeobotanical Macroremains of Taxus from the coastal plain of data are generally sparse for chalk regions, since Belgium and the southern Netherlands (Fig. 1), the geological and topographical conditions in except those associated with archaeological sites, these regions are unsuitable for the formation and have all been recovered from the part correspon preservation of stratified peat(T ittensor 1980). ding to the carr-phase of the peat profiles. They This could, of course, bias the reconstruction of were associated with seeds and other macroscopic the Holocene distribution of Taxus presented. remains of plants typical of an alder/birch carr or However, this objection may be valid for the fen vegetation composed of, e.g.,Alnus glutinosa , areas with actual natural Taxus populations in Betula sp., Comarum palustre , Carex paniculata , Belgium (Fig. 1) but this is not true for the areas Carex pseudocyperus, Lysimachia vulgaris , Lyco- both to the northwest and to the southeast of this pus europaeus and Thelypteris palustris (V a n region. For those areas, Holocene pollen dia H oorne 1951,S tockmans & V anhoorne 1954, grams and other palaeobotanical data are avail A llemeersch 1991). The occurrence ofTaxus in able, but no Taxus pollen were recorded outside pollen diagrams from these areas also corre the coastal area and the lower Scheldt valley, one sponds with the carr-phase of the analysed peat exception being the records of Taxus charcoal profiles (e.g., M unaut 1967,D eforce & B asti - from Gutland (Luxembourg). aens in press).

The fact that, at sites mentioned above, The Ta x u s d e c lin e Taxus was actually part of the local vegetation community has been ignored or even denied by At the above-mentioned sites from Belgium several authors as it does not seem to correspond and surrounding regions, Taxus disappears in the with the present day ecology and distribution of pollen diagrams during the second half of the this tree. V a n Smeerdijk (2003: 161-162), for Sub-boreal, around 3 500 uncal. BP (see Table 1). example, argued that the high percentages of Similarly, no botanical macroremains of Taxus Taxus pollen at Ellewoutsdijk and Baarland must have been found that are younger than the end of be explained by transport by the river Scheldt and the Sub-boreal. In pollen diagrams from other thus must originate from a more inland area. sites situated in the lowlands of northwestern However, the fact that Taxus wood has been dis Europe, Taxus disappears as well, or shows a covered at Ellewoutsdijk and at the nearby strong decline, before the end of the Sub-boreal. Temeuzen, does not support this hypothesis. The Holocene decline of Taxus in northwest Besides, the river Scheldt followed a more north ern Europe is generally attributed to competition ern route at that time and did not flow near Elle with Fagus and Carpinus , deforestation, selective woutsdijk or Baarland (Vos & van H eeringen felling and grazing (Firbas 1949, Averdieck 1997,D enys & V erbruggen 1989). Many other 1971,Z oller 1981, Svenning & M agârd 1999, records of pollen and macroremains of Taxus N avys 2000, H oltan 2001, Thomas & Polwart from peat deposits, often in areas without any flu 2003). However, these explanations are all based vial activity, indicate thatTaxus actually did grow on the actual ecology and distribution of Taxus, on peat. In fact, this is not an entirely new obser i.e., Taxus growing on well-drained calcareous vation as Godwin already remarked, based on his soils. For the decline of Taxus growing in a fen carr research at Wood Fen (Ely, UK; G odwin et al. environment, other explanations must be sought. 1935), at Woodwalton Fen (Hunts, UK; G odwin The most common explanation for the Sub- & Clifford 1938) and on the Taxus finds at boreal Taxus decline is competition with Fagus Temeuzen (The Netherlands; G odwin 1968), that and Carpinus (Firbas 1949,A verdieck 1971, “Taxus almost certainly had an extensive natural M uhle 1979). Recent research showed that a

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Taxus population in Denmark increased after cause tangible damage to the trees (M ysterud & thinning the tree canopy, especially by felling 0 STB ye 1995, 2004N avys 2000). Fully-grown beech (S venning & M agârd 1999). Other Taxus trees are largely resistant to the nibbling; research demonstrated, however, that regenera even after a tree is cut down, green shoots appear tion of Taxus could be rather successful under the from the stump. On the other hand, recent canopies of several broadleaved trees, including research has demonstrated that roe deer browsing Carpinus betulus (Iszkulo & B oratynski 2004). reduces Taxus recruitment (H ulme 1996,G arcia Moreover, as Fagus and Carpinus only grow on & Obeso 2003, M ysterud & O stbye 2004). In well-drained soils (E llenberg et al. 1991), light general, although roe deer occur in wetland habi competition with these two taxa cannot have tats (Danilkin 1996,B arancekova 2004), it is played a role in the decline of Taxus growing in very unlikely that the Sub-boreal Taxus decline wet conditions. can be explained by roe deer -browsing, as there Another explanation for the Taxus decline are no indications for an increase in their popula could be deforestation, but the pollen diagrams tion at that time. from Belgium and the southern Netherlands do One more possible explanation for theTaxus not indicate deforestation during the period of the decline would be a climate change. There is no Taxus decline. There are also no indications for evidence, however, for a major change of the cli agriculture or other forms of intensive human matic conditions in northwestern Europe around impact on the vegetation in the coastal lowlands 3 500 uncal. BP (Davis et al. 2003). As Belgium during the Sub-boreal (Vos & van H eeringen and the southern Netherlands are not situated near 1997,Ervynck et al. 1999). the limits of the natural distribution of Taxus, it is Selective felling of Taxus, for its valuable unlikely that a minor change in climatic condi wood or to avoid cattle poisoning, has been pro tions would have caused the Taxus decline. posed as another explanation for the Taxus In conclusion, although some of the above- decline at several sites in northwestern Europe mentioned explanations for the Taxus decline (S armaja -K orjonen et al. 1991,S venning & might hold true for Taxus stands growing on well- M agârd 1999,O ’Connell & M olloy 2001). drained, mineral soils or in regions where human From the Neolithic onwards, Taxus was probably impact was more intense (Tittensor 1980, the most used wood for the manufacture of bows O ’Connell & M olloy 2001), they are not suit (C lark 1963, Lanting et al. 1999,B euker able to explain the Sub-boreal decline of Taxus in 2002). Many other wooden implements were the coastal area of Belgium and the southern made from Taxus as well (G odwin 1975,C oles et Netherlands. As a more likely explanation for the al 1978,G ale & Cutler 2000). However, since decline of Taxus in Belgium and the southern human populations and activities were almost Netherlands, a change of the environment in which absent during the Sub-boreal, in the region under Taxus grew can be proposed. In most places in the consideration here, these explanations can also be coastal area and the Scheldt estuary, this environ rejected (Vos & van H eeringen 1997,E rvynck mental change could consist of the already men et al. 1999,Louwe K ooijmans et al. 2005). tioned transition from the carr peat phase, in which In several forests of northwestern Europe, it most of the palaeobotanical records of Taxus can has been observed that Taxus recruitment suffers be situated, to ombrotrophic moss peat and, in from browsing by roe deer, Capreolus capreolus most places, the development of raised bogs result (G arcia & O beso 2003, M ysterud & O stbye ing in the formation of Sphagnum-Ericaceae peat 2004). The branchlets, needles and seeds of Taxus (A llemeersch 1986, 1991,Pons 1992,V er contain a poisonous alkaloid, a lethal toxin for bruggen et al. 1996,D eforce & B astiaens in many species including horses, cows, goats and press.). Especially the pollen diagrams from humans (Jordan 1964, Schulte 1975). Only a Oorderen (M unaut 1967) and Raversijde few animals including roe deer are not sensitive (Deforce & B astiaens in press) show very clearly to it; they like to nibble the yew branchlets and that Taxus 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 IN BELGIUM 231 ombrotrophic conditions, illustrated by the A l l e m e e r sc h L., 1991. — Peat in the Belgian eastern increase of Sphagnum and Ca//w«a/Ericaceae. coastal plain. In: G u l l e n t o ps F. (ed.), Wetlands In the inland part of the lower Scheldt valley, in Flanders. Contributions to the palaeohydrol- where no ombrotrophic peat occurs on top of the ogy of the temperate zone in the last 15 000years. fen-carr peat deposits, the end of the peat growth Aardkundige Mededelingen 6, pp. 1-54. Leuven was caused by the deposition of alluvial loam and University Press, Leuven. A n d e r s e n S.Th., 1975. — The Eemian freshwater clay, as a consequence of agricultural practices deposit at Egemsund, South Jylland, and the (V erbruggen et al. 1996,H uybrechts 1999). Eemian landscape development in Denmark.Dan- marks Geologiske Undersogelse A 1974: 49-70. A v e r d ie c k F.-R., 1971. — Zur postglazialen CONCLUSIONS Geschichte der Eibe {Taxus baccata L.) in Nord westdeutschland. Flora 160: 28-42. All Holocene palaeobotanical records of A v e r d ie c k F.-R., 1983. — Palynological investigations Taxus from Belgium and the southern Netherlands of the sediments of ten lakes in eastern Holstein, show three remarkable characteristics: (1) they all North Germany. Hydrobiologia 103: 225-230. are situated in the coastal plain and the lower B a e t e m a n C., 1981. —De Holocene ontwikkeling van Scheldt valley, (2) they all date from the Sub- de westelijke kustvlakte (België), Ph.D. thesis, Vrije Universiteit Brussel, Brussel, Belgium. boreal and (3) they all indicate that Taxus grew on B a e t e m a n C., 1999. — The Holocene depositional his peat. This strongly contrasts with the recent distri tory of the Ijzer Palaeo-valley (western Belgian bution and ecology of Taxus, the current natural coastal plain) with reference to the factors con distribution of Taxus baccata L. in Belgium being trolling the formation of intercalated peat beds. limited to a few localities in the southern part of the Geológica Belg. 2: 39-72. country, all situated on steep, calcareous slopes. B a e t e m a n C., 2005. — How subsoil morphology and The Holocene occurrence of Taxus in the erodibility influence the origin and pattem of late coastal plain in Belgium, and probably in several Holocene tidal channels: case studies from the other lowland areas in northwestern Europe, cor Belgian coastal lowlands. Quatem. Sei. Rev. 24: relates with the carr peat phase of the surface or 2146-2162. Holland peat, which was mainly formed during B a e t e m a n C. & V e r b r u g g e n C., 1979. —A new approach to the evolution o f the so-called surface the Sub-boreal. The decline and disappearance of peat in the western coastal plain of Belgium. Pro Taxus in northern Belgium and the southern fessional Paper 167, Belgische Geologische Netherlands during the second half of the Sub- Dienst, Bmssel. boreal are most likely the result of the transition B a e t e m a n C., C l e v e r in g a P. & V e r b r u g g e n C., of these coastal marshes from a fen-carr environ 1981. — Het paleomilieu rond het romeins ment to ombrotrophic bogs. zoutwinningssite van Leffinge. Professional Paper 186, Belgische Geologische Dienst, Bmssel. ACKNOWLEDGEMENTS B a e t e m a n C., S c o tt D.B. & V a n S t r y d o n c k M., 2002. — Changes in coastal zone processes at a The authors wish to thank Nele Van Gemert for high sea-level stand: a late Holocene example help with the production of the figures and Otto from Belgium. J. Quatern. Sei. 17: 547-559. Brinkkemper for valuable comments. B a r a n c e k o v a M., 2004. — The roe deer diet: Is floodplain forest optimal habitat? Folia Zool. 53: 285- 292. REFERENCES B a tc h e lo r C.R., B r a n c h N., E l ia s S., H a w k in s D. S h a w P. & S id e l l J., 2004. —Middle Holocene A d a m s J., 1905. — The occurrence of yew in a peat environmental history of the lower Thames val bog in Queen’s County. Irish Naturalist 14: 34. ley. The history of Taxus woodland, pp. 79-80. A l l e m e e r sc h L., 1986. — Hochmoortorfe im QRA International Postgraduate Symposium, östlichen Küstengebiet Belgiens. Courier Royal Belgian Institute of Natural Sciences, Forsch-Inst. Seckenberg 86: 397-407. Bmssel, Belgium.

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