Studia Quaternaria, vol. 32, no. 1 (2015): 19–29.

DOI: 10.1515/squa-2015-0002

ANTHROPOGENIC TRANS FOR MA TION OF THE VEG E TA TION IN THE IM ME DI ATE VI CIN ITY OF THE SET TLE MENT COMPLEX AT (MR¥GOWO LAKELAND, NE )

Marta Szal1, Miros³awa Kupryjanowicz1, Mariusz Wyczó³kowski2 1Insti tute of Biol ogy, Uni ver sity of Bia³ystok, Œwierkowa 20b, 15-950 Bia³ystok, Poland; e-mail: [email protected], [email protected] 2Wojciech Kêtrzyñski Museum, Plac Zamkowy 1, 11-400 Kêtrzyn, Poland; e-mail: [email protected]

Ab stract The re sults of pol len, non-pol len palynomorph and microcharcoal par ti cle anal yses of de posits from a small pond in north east ern Po land are presented. The study fo cused on hu man-in duced veg e ta tion changes that oc curred in a close vi cin ity of the set tle ment complex at Poganowo dur ing the Mid dle Ages (ca 10th–16th cen tu ries). We dis tin guished three phases of hu man impact. First and third phases cor re spond to in ten sified set tle ment ac tiv ity. The sec ond phase was a pe riod when hu man ac tiv ity de creased and woodland re gen er a tion took place. The high in ci dence of the par a - sitic fun gus Kretzschmaria deusta in a lo cal for est stand dur ing the third phase was simul ta neous with nu merous spores of coprophilous fungi (Sordaria-type and Cercophora-type). We con sider that Kretzschmaria deusta in hab ited the roots and bases of tree trunks dam aged by dig ging and graz ing an i mals.

Key words: human impact, pollen analy sis, fungal spores, Masuria, Middle Ages.

Mansuscipt received 1 December 2014, accepted 1 June 2015

IN TRO DUC TION for ma tion of veg e ta tion caused by hu man ac tiv ity. More - over, by the appli ca tion of NPP and microcharcoal parti cle A use of in di cator charac ter istics of some plant taxa, as anal y sis we tried to char ac ter ize farm ing mod els, as sum ing well as of other organ isms, is a funda men tal method in the possi bil ity of attrib ut ing the presence of spores of copro- palaeo ec ol ogy, en abling re con struc tion of past en vi ron - philous fungi (e.g. Sordaria-type, Sporormiella-type) to ments (e.g. Birks, 2003; Lata³owa et al., 2013). Among the land-use prac tices, par tic u larly hus bandry. microfossils that are observed in pollen slides, fungi, algae and other non-pollen palynomorphs (NPP) have become par- tic u larly im por tant, be cause of their po ten tial to im prove re - AR CHAE O LOG I CAL AND HIS TOR I CAL construc tion of past ecosys tems (van Geel, 2001; van der Lin den et al., 2012). Van Geel and Aptroot (2006) under - BACK GROUND lined that fungal remains are of high po tential in this context. The settle ment com plex at Poganowo is lo cated in the In par tic u lar, these au thors em pha sized the im por tance of northeast ern part of the Mr¹gowo Lakeland. The earli est fun gal taxa with well-char ac ter ized bi o log i cal and eco log i - traces of human activ ity in the region have been dated to the cal re quire ments, e.g. Sporormiella sp. and Sordaria sp., Neo lithic pe riod. Nev er the less, gen er al iza tion of per ma nent which are indi ca tors for dung, or Kretzschmaria deusta, a set tlements occurred with the begin ning of the West Balt par a site on de cay ing wood, in di cat ing pres ence of their host Barrow culture in the early Iron Age (ca 7th cen tury BC). The trees. How ever, a palaeo eco logi cal po ten tial of fun gal re - first set tle ment phase in the Poganowo hillfort is linked to mains has not yet been fully explored. We present results of this cul ture (Wyczó³kowski et al., 2013), and the charcoal palynological anal y sis in the frame work of in ter dis ci plin ary and ash layer (perhaps from burnt tim ber construc tions of re search pro ject con ducted in the set tlement complex at ramparts or pali sades) of this phase are dated to the 6th–4th Poganowo (Szal et al., 2013; Wyczó³kowski et al., 2013). century BC. Other sites, located a short distance from Poga- This set tle ment complex is lo cated en tirely in a for est, and nowo, include the forti fied settle ment at , the set - nearby there are nu merous small peat bogs and ponds suit - tlement in Wólka, and the barrow cem eter ies in able for lo cal-scale veg e ta tion re con struc tions (com pare and Nakomiady, also dated to the early Iron Age (Hoffmann, Sugita et al., 2007). Our main objec tive was to study trans- 1999; Wyczó³kowski, 2011). 20 M. SZAL et al.

Fig. 1. Lo ca tion of the study area pro duced by using the dig i tal ter rain model from LIDAR data (a) and Google Maps (b) with PG6 cor ing site (54°00'11.2"N 21°23'04.6"E); 1 –Poganowo hillfort, 2 – sac ri fi cial place, 3 – set tle ments.

An increase in a number of sites related to the popu la tion performed by using an Instorf peat sampler. The core con - of the Bogaczewo culture during the Roman Pe riod (1st–4th tained sandy peat (0.9–0.78 m depth) and fen shrub peat century AD) has been observed (Szymañski, 2005). At that (0.78–0.2 m depth; D. Drzymulska, personal comm.). For time, two settle ments ex isted in the vicin ity of the Poganowo techni cal reasons the highly hydrated topmost part of the de- hillfort. There is a ceme tery in Turw¹gi (Voß, 1886), and a posit could not be retrieved. set tle ment and sac ri fi cial bog site at Wólka (Raddatz, 1992– 1993). Pol len anal y sis A sec ond set tlement phase in the Poganowo hillfort has been dated to the late 10th–12th centu ries AD. In north eastern Sam ples of 1 cm3 were pre pared us ing the standard pro- Poland this pe riod is called the Early Middle Ages or the Vi- cedure of Erdtman’s acetolysis (Berglund and Ralska-Jasie- king Age (e.g. Nowakiewicz, 2006, 2010; Nowakiewicz and wiczowa, 1986). De pending on the content of mineral matter Wróblewski, 2010). At that time, a sac ri fi cial place ex isted, the samples were treated with a heavy liquid (CdI2+KI). The and an ex ten sive set tlement has been ex cavated (Wyczó³- palynological analy sis was performed on 35 samples from kowski et al., 2013). Ra diocar bon dating (1338±58 AD) the PG6 profile. Pollen anal ysis was carried out with an from the hearth of the sacri fi cial place may in di cate a use of Olym pus BX43 light mi cro scope with mag ni fi ca tion of this place as late as in the 13th and 14th cen tu ries. 600×; a larger magni fi ca tion was used to identify problem - In the second half of the 14th cen tury the for est where the atic and small palynomorphs. The identi fi ca tion of ce- Poganowo hillfort is located was given to the city of Kêtrzyn real-type pollen was based on phase contrast studies under (Rastenburg). From the first half of the 15th century the vil- 1000× mag ni fi ca tion. For tax o nom i cal iden ti fi ca tion pol len lages began to develop (Beckherrn, 1880). Finally, in 1571 keys (e.g. Beug, 2004) and a refer ence collec tion of modern AD, the vil lages Prêgowo, S³awkowo and Poganowo, lo- pollen slides were used. On aver age 900 terres trial pol len cated in the northern and western part of the forest, included grains were counted and identified in each sample. 93 hufen (1 hufe is ca 16.8 ha) that were managed by 43 farm- ers (Visi ta tion, 1571). The num ber of farms exist ing on the Anal y sis of non-pol len palynomorphs edge of Poganowo did not change signif i cantly until the end th and microcharcoal of the 18 century (Goldbeck, 1785). Non-pollen palynomorphs (NPPs) were counted along with the pollen analy sis, and identified accord ing to van Geel MA TE RIAL AND METH ODS (1978, 2001), Bell (1983), Jankovská and Komárek (2000), van Geel et al. (2003) and van Geel and Aptroot (2006). Cor ing and li thol ogy Microcharcoal par ticles were counted in the pol len sli- The sed iment core (PG6 pro file) was collected in win ter des and grouped into four size classes: 10–30 µm, 30–70 µm, of 2012 from a small pond located ca 150 m to the northeast 70–100 µm and >100 µm. Accord ing to Tinner and Hu of the settle ment complex at Poganowo (Fig. 1). Coring was (2003) the larger par ticles better re flect the in cidence of lo cal ANTHROPOGENIC TRANS FOR MA TION OF THE VEG E TA TION 21 fires, whereas the smaller par ticles are more in dic ative of re- Ta ble 1 gional fires. Par ticles <10 µm were ig nored, as they cannot be AMS radio car bon dates from the PG6 core used safely identi ed (Blackford, 2000). One tablet contain ing for chro nol ogy 20,848 marker spores of Lycopodium was added to each Cal i brated age range Cal i brated age range sample to enable a quanti ta tive analy sis of microfossil con- Depth Age 14C Lab. No. 68% 95% cen tra tions (Stockmarr, 1971). Char coal par ticles were re - (cm) (BP) (cal en dar age, AD) (cal en dar age, AD) corded with a light micro scope at 400× magni fi ca tion. Char- 1266 (78.4%) 1308 48 GdA-3545 695±25 1274 (68.2%) 1296 coal se lection was restricted to frag ments that were black, 1362 (17.0%) 1385 completely opaque and angu lar (Whitlock and Larsen 2001). Cal cu la tions and the pre sen ta tion of palynological and microcharcoal data were performed with POLPAL for Win- First phase (LPAZ PG6-1) dows (Nalepka and Walanus, 2003). The AP+NAP sum was used for percent age calcu la tions, both in the pollen and the NPP Vege ta tion changes in the study area at the begin ning of dia grams. The pollen di agram was stratigraphically ordered and this phase repre sent a period of strong human im pact. Pollen zoned with Con strained Clus ter Anal ysis (CONISS), and di- spec tra illus trate oc cur rence of a pine-birch for est, where the vided into lo cal pollen assem blage zones (LPAZ), which un dergrowth was dom inated by ha zel. The ex istence of open were named ac cording to the specific com po sition of pollen surfaces and/or good light condi tions on the forest floor is in- spectra. The human impact groups follow Behre (1981), di cated by the pres ence of Juniperus communis pollen. Berglund and Ralska-Jasiewiczowa (1986), and Gaillard and More over, Juniperus communis indi cates that dry and mod - Berglund (1988). er ately moist soils oc curred near the study site, what can be Princi pal com ponent analy sis (PCA) served for the de- deduced from edaphic require ments of this species (Za- tection of interspecies re lation ships. The anal ysis was based rzycki, 2002). The abun dant oc currence of microcharcoal on data from all pollen sam ples; sporad i cally occur ring arbo - par ticles, in all size classes, may re ect hu man impact on the real taxa and non-ar bo real taxa were ex cluded. PCA was car - en vi ron ment. The pres ence of Melampyrum and Pteridium ried out using XLSTAT 2014 trial version software. aquilinum, which are con sidered in dica tors of soil enrich - ment by ash (Lata³owa, 2007), addi tion ally confirms the oc- Chro nol ogy currence of fires in the area. Most proba bly, it was inten tional use of fire, point ing to ac tiv ity within the set tle ment and to The age of one sedi ment sample from a depth of 0.48 m 14 woodland clearing. The palynological data reveal the exis - was de ter mined by ra dio car bon dat ing ( C/AMS) in the tence of large open ar eas near the sample site, as in di cated by Gliwice Ra dio car bon Lab o ra tory, In sti tute of Phys ics, Sile- nearly 20% of NAP. A con sid er able area was cov ered by sian Uni ver sity of Tech nol ogy (Ta ble 1). mead ows and pas tures, rep re sented mainly by Poaceae, Cy- peraceae, Filipendula and Centaurea nigra-type, and ruderal RE SULTS com mu ni ties with Ar te mi sia and Chenopodiaceae. Only small plots were used as fields. Lo cal pol len as sem blage zones In the inves ti gated water body aquatic macrophytes like The pollen dia gram was divided into three LPAZ (from Potamogeton, Nymphaea alba, Nymphaea candida and Nup- LPAZ PG6-1 to LPAZ PG-3), which corre spond to the sub- har occurred. Along the lakeshore reed swamp com mu nities se quent phases of veg e ta tion de vel op ment in the im me di ate with Typha latifolia and Typha angistifolia occurred. Nu- vicin ity of the sampling site (Fig. 2). The pollen zones are de- mer ous green al gae, in clud ing Tetraedron, Pediastrum and scribed (Ta ble 2). Botryococcus, devel oped in the water body. These algae, as well as cyanobacteria of the Gloeotrichia-type, play an im- Prin ci pal Com po nent Anal y sis (PCA) portant role in in ferring past lake con ditions, and almost all are known to be dom inant in more eutrophic sys tems (e.g. It de scribes sta tis ti cally pos si ble re la tion ships be tween van Geel and Grenfell, 1996; Jankovská and Komárek, arbo real pollen and the fun gal spores re cord (Fig. 3). The re- 2000). More over, Gloeotrichia is one of the most im portant sults in di cated that there may be a dis tinct pos itive cor re la - organ isms today causing so called ‘water blooms’, linked to tion between the pol len record of forest species, e.g. Quer- the en rich ment of the aquatic envi ron ment with phospho rus cus, Alnus, Picea abies, Carpinus betulus, and the spores of and nitro gen (Burchardt and Pawlik-Skowroñska, 2005). We the par a sitic fun gus Kretzschmaria deusta and the coprophi- may assume that the rel a tively high trophic sta tus of the in - lous fungi Sordaria-type and Cercophora-type. vesti gated water body was proba bly due to a large supply of nu tri ents in re sponse to the in ten sive set tle ment ac tiv ity in IN TER PRE TA TION AND DIS CUS SION this area. Re con struc tion of veg e ta tion dy nam ics Linking the first phase of anthropogenic vege ta tion chan- ges with a de nite cul tural event is not pos sible due to the avail - and hu man impact abil ity of only one radio car bon date from a depth of 0.48 m in The changes in the com posi tion of the pollen assem - this sec tion. How ever, it could be re lated ten ta tively to the sec - blages recorded in the PG6 profile enabled to distin guish ond phase of occu pa tion of the hillfort at Poganowo, dated to three phases of veg eta tion dynam ics connected with human the early Mid dle Ages (10th–12th centu ries AD). Archaeobo- ac tiv ity (Fig. 2). tanical anal ysis of the early me di eval layers and hearths 22 M. SZAL et al. aw – H ,deta vi tluc – G ;sdeew dna slaredur – F ,stnalp deni fe dnu ylla c igo loce dna wodaem – E ,ts erof debru tsid fo stnalp – D :sburhs frawd dna sbreh ;stnalp suolihpoileh – C ,stnalp gn ivol-edahs gn ,stnalp C – suolihpoileh ;stnalp sbreh dna frawd :sburhs D – stnalp fo tsid debru erof ,ts E – wodaem dna loce igo c ylla dnu fe deni ,stnalp F – slaredur dna ;sdeew G – tluc vi ,deta H – aw - –

B within the settle ment com plex at Poganowo con-

; s t firmed a cul ti va tion of ce re als in this time. Re mains a t i t

b of barley (Hordeum vulgare), millet (Panicum mi- a h liaceum), wheat (Triticum), rye (Secale cereale) h s

e and oats (Avena) were found (K. Piñska, personal r f

d comm.). Accord ing to Wyczó³kowski and Mako- n a

t wiecki (2009), it can be claimed that the inhab it ants e

w of the early mediaeval set tlement at Poganowo

f o raised cat tle and horses. s t n a l p

- Second phase (LPAZ PG6-2) A

: s

b The pe riod of in tensive exploi tation of the en- u r

h viron ment during the first phase was followed by a s

d phase of de creased hu man ac tiv ity (de crease in n a

s NAP frequency and concen tra tion of microchar- e e

r coal par ti cles) lead ing to the for est re gen er a tion. T

.

e Then, con sid er able changes in for est com po si tion r o

c 6GP eht morf dro morf eht 6GP c oc curred. An in crease of Quercus pollen is syn- chronous with an increase in pollen of the other main ar bo real taxa: Tilia, Ulmus, Carpinus, Picea, Fraxinus, and expan sion of Salix shrubs. The vicin - ity of the studied site be came almost completely forested. Dry morainic hills were overgrown by

cer lao crahc dna shpromonylap ne llop-non ,axat ydoow dna breh detce les htiw mar g aid eg atne crep ne lloP lloP ne crep atne eg aid g mar htiw les detce breh dna ydoow ,axat llop-non ne shpromonylap dna crahc lao cer mixed pine forests. Patches of fer tile, fresh soils were cov ered by oak-hornbeam forest, with an ad- mixture of spruce. Consid er able areas in the lake- shore zone and in other wet places were cov ered by

.taep htiw dnas – 2 ,taep burhs nef – 1 :yg oloh tiL .ignuf – J ,ea gla neerg – I :sPPN ;stnalp gn ivol-ret gn ;stnalp :sPPN I – neerg gla ,ea J – .ignuf tiL oloh :yg 1 – nef burhs ,taep 2 – dnas htiw .taep alder woods and carrs. At this phase, forest regen er - ation relates to a very low level of human exploi ta- tion of the landscape and possi bly reflects the time when the Poganowo settle ment complex was aban- doned. There is an assump tion that this could have happened in the early 13th century (M. Wyczó³kow- ski, per sonal comm.). Ac cord ing to his tor i cal sources (Saage and Woelky, 1860), in the surround - ings of the Poganowo hillfort there was a large for- ested area called Tauro (“Walt Tauro”), which was mentioned in docu ments dated to 1231–1340 AD. Nev er the less, pollen spec tra from the PG6 sec tion in di cate that some pas to ral ag ri cul ture seems to have been practiced at this time, as well as ar able farming. Accord ing to Vuorela (1973), Cerealia- type pollen grains are not well dispersed in air- streams, and there fore their poor rep re sen tation cannot be consid ered as a guide to the impor tance of crop farming in the area. Admit tedly, some weeds of ar a ble fields were pres ent in the pol len re cord, but in low frequen cies only. The appear ance of Centaurea cyanus was proba bly connected with the culti vation of winter crops. Other weeds docu - mented in the PG6 pollen profile include Chenopo- diaceae, Papaver and Polygonum aviculare. Can- na bis/Humulus-type pollen most proba bly indica- tes that Humulus lupulus grew in this area as a na- tive plant on damp soils. The area of meadows signif icantly de creased,

. but a low-level pres ence of Plantago lanceolata, 2

. Rumex acetosella-type, Rumex acetosa-type and g i

F Cichoriaceae pollen grains in di cates that some open ANTHROPOGENIC TRANS FOR MA TION OF THE VEG E TA TION 23

Ta ble 2 Local pollen assem blage zone descrip tion from PG6 core dia gram

Name of L PAZ and depth Description Dom i nance of Pinus sylvestris-type (14-43%) and Betula alba-type (15-45%). High val ues of Corylus avellana (to 19%) with short-last ing peak (27%) in the middle part of zone. Several per centage frequency of Alnus (to 2%) and Quercus (to 1.5%). Sporadic presence of other tree taxa. Rela tively fre quent Juniperus communis. Con sider able partic i pa tion of NAP PG6-1 (9–18%). The herba ceous spectra dom inated by Poaceae (to 15%). Regu larly present Filipendula. Among the anthropo- Pinus-Betula-Poaceae genic in di ca tors, sys tem atic pres ence of Ar te mi sia, Chenopodiaceae and Cerealia-type. Frequent pollen grains of 90-80 cm limnophytes, such as Potamogeton and Typha latifolia. Reg u lar oc cur rence of green algae (Botryococcus, Pediastrum, Tetraedron) and sin gle re mains of cyanobacteria (sheaths of Gloeotrichia).Very high con cen tra tion of mi cro scopic char - coal par ti cles. In crease of Alnus above 20%. Signif i cant de crease of Betula alba-type. Dom i nance of Pinus sylvestris-type (20–47%). High val ues of Corylus avellana (to 14%). Increas ing curves of Tilia cordata (to 7%), Ulmus (to 3%), Quercus (to 12%), Carpinus betulus (to 12%), Picea abies (to 4%) and Salix (to 4%). Less numer ous NAP (6–9%). Gradual decrease of PG6-2 Poaceae (to 3%). Frequent Filipendula, Cyperaceae, Apiaceae and Cichoriaceae. Reg u lar oc cur rence of Arte mi sia . In sev- Alnus-Pinus-Quercus-Tilia eral spec tra, pres ence of Cerealia-type, Centaurea cyanus-type, Plantago lanceolata, Rumex acetosella-type, Rumex 78–58 cm acetosa-type and Chenopodiaceae. Decreas ing values of green algae. Single occur rences of coprophilous fungi (Sporormiella-type, Sordaria-type and Cercophora-type). Spo radic Kretzschmaria deusta. Rel a tively low con cen tra tion of mi cro scopic char coal par ti cles. PG6-3 Betula-Quercus-Carpinus- Dom i nance of Betula alba-type, Pinus sylvestris-type and Alnus. Constant presence of Tilia cordata, Quercus, Carpinus Alnus betulus and Ulmus. The PG6-3 LPAZ was divided into three subzones. 56–20 cm Fast in crease of Betula alba-type (to 48%). High val ues of Corylus avellana (to 20%). Frequent Populus. Grad ual de - PG6-3a crease of Pinus sylvestris-type (to 8%). Rela tively high values of Alnus (10–13%), Quercus (4–10%) and Carpinus betulus Betula-Corylus (5–11%). More fre quent Poaceae (to 7%) and Cyperaceae (to 4%). Decreas ing val ues of Apiaceae, Filipendula and 56–50 cm Cerealia-type. Lower fre quency of Arte mi sia . Rela tively frequent Cercophora-type. Spo radic oc cur rence of Sporo- rmiella-type, Sordaria-type and Kretzschmaria deusta. Rel a tively high con cen tra tion of mi cro scopic char coal par ti cles. Strong increase of Betula alba-type to 63%, then decrease to 24%. In creasing per centage val ues of Pinus sylvestris-type (to 21%). Lower values of Corylus avellana (4–12%) and Populus (to 1%). Fluctu at ing increase of Alnus (to 28%). Grad - PG6-3b ual de crease of Poaceae. More fre quent Arte mi sia and Can na bis/Humulus-type. Reg u lar oc cur rence of Cerealia-type and Betula-Alnus sin gle oc cur rences of Secale cereale and Hordeum-type. Spo radic Centaurea cyanus-type, Plantago lanceolata, Rumex 48–36 cm acetosella-type, Rumex acetosa-type. More fre quent Cercophora-type and Kretzschmaria deusta. Sporadic Sporormiella-type and Sordaria-type. Much lower con cen tration of mi cro scopic charcoal par ti cles. The age of the sed i - ment at 0.48 m depth was esti mated at 1266–1385 AD (Table 1). High val ues of Quercus (to 16%) and Pinus sylvestris-type (to 20%). Increas ing tendency of Alnus (to 34%). Gradual de- PG6-3 crease of Betula alba-type (to 15%). Carpinus betulus oscil lat ing around 7–11%. Constant presence of Corylus avellana Quercus-Carpinus- (to 12%). Higher values of Ar te mi sia. Reg u larly occur ring Poaceae, Cyperaceae and Apiaceae. More frequent Plantago Kretzschmaria lanceolata, Rumex acetosa-type and Ranunculaceae. Spo radic Rumex acetosella-type. Tem po rary de creas ing ten dency of 34–20 cm Cerealia-type. Strong in crease of Kretzschmaria deusta to 12%. In creasing tendency of Cercophora-type. Spo radic occur - rence of Sporormiella-type and Sordaria-type. Low con cen tra tion of mi cro scopic char coal par ti cles. her ba ceous plant com mu ni ties oc curred as fod der for do mes- ti cated her bi vores. LPAZ PG6-2 repre sents a period of gener ally low an- thropogenic activ ity in the catchment area and, based on de- creased per cent ages of Botryococcus, Pediastrum, Tetrae- dron and Gloeotrichia, eutrophication dimin ished. How- ever, the de crease of algae and some aquatic plants co in cides also with sedimentological changes. It should be un derlined that lo cal terrestrialization, a tran si tion from wetland to drier peat, could be an alter native expla na tion for the decline in al- gae and cyanobacteria.

Third phase (LPAZ PG6-3) The pol len record from this core inter val en abled to de- ter mine three sub-phases that in dicate response of local vege - ta tion to a vari abil ity of hu man ac tiv ity dated to 13th–16th cen tu ries AD.

Fig. 3. PCA plot illus trat ing the interspecies rela tionships in the pol len spec tra from the PG6 core. 24 M. SZAL et al.

In the oldest part (LPAZ PG6-3a) the most charac ter is tic Secale, Hordeum and Fagopyrum were among the culti vated phe nom e non was wood land clear ance and the ex pan sion of plants. However, the spectrum of herba ceous taxa mainly birch. Mi cro scopic char coal con cen tra tions steeply increa- points to the exis tence of meadows (pollen of Poaceae, sed, indi cat ing the presence of fire events in the vicin ity. Centaurea nigra-type, Filipendula). In the surround ings of Moreover, the presence of spores of Pteridium aquilinum, the inves ti gated site, there occurred also ruderal plants, such which easily colo nizes disturbed ground, includ ing burnt ar- as Ar te mi sia, Urtica and Chenopodiaceae, which, accord ing eas (Hartig and Beck, 2003), may also point to fires. The fires to Behre (1981), are asso ci ated with nitro gen-rich habi tats. could have served as a silvicultural tool for defor es ta tion, or They may also have grown along the ni trogen-rich banks of just to create openings in a tree canopy. Lo cal devel op ment the lake. In tree-less parts of the land scape grazed grass lands of birch woodland and shrubs is perhaps asso ci ated with were pres ent, as in di cated by Plantago lanceolata, Plantago Betula, fea tur ing as a prom i nent pi o neer spe cies af ter fires ma jor/me dia, Potentilla-type, Cerastium-type, Rubiaceae, (Connor et al., 2012; Lindbladh et al., 2003). Moreover, at and Cichorioideae, which are consid ered to indi cate pas toral the begin ning of this phase Alnus also in creases, which plau - ac tiv i ties (Mazier et al., 2006; Hjelle, 1999). Moreover, Ma- si bly il lus trates its fire-re lated be hav iour. Al der was clas si - zier et al. (2006) concluded that Plantago lanceolata, Plan- fied by Tinner et al. (2000) to the group of taxa whose abun- tago ma jor/me dia and Chenopodiaceae could be also treated dance largely depends on fire. The rapid increase in the pol- as indi ca tors of woodland grazing. This may indi cate that do- len values of the light-de manding Corylus, ac com pa nied by mes ticated her bi vores grazed close to the sampling site. In the more frequent Populus, is evi dence for forest opening. ad di tion, pol len grains of Polygonum aviculare and Rumex Corylus as an understory spe cies produces little pollen under acetosella-type were recorded. These weeds thrive on tram- a heavy tree canopy, and large quan tities of ha zel pol len may pled, disturbed ground, and were classi fied by Pokorná et al. result from bushes growing at the wood land margin and in (2014) as re li able in di ca tors of tram pled veg e ta tion. clearings (Rackham, 1988). In our in vesti gation high fre- In the youngest sub-phase (LPAZ PG6-3c) a sig nifi cant quen cies of Corylus pollen, up to 22%, suggest that hazel had spread of Quercus was observed. This pro cess was ac compa - a dom i nant po si tion in the veg e ta tion, in di cat ing that pol len nied by an in crease of Carpinus betulus and Pinus sylvestris dis persal was hardly hin dered by trees. How ever, there were con tem po ra ne ous with a de crease in Betula and Picea abies. more changes in the woodland compo si tion that occurred as The pol len record shows that the surround ings of the study con se quence of for est fires. A re duc tion of de cid u ous tree site were overgrown by mixed pine-oak for ests and by rich, pol len was ob served, par tic u larly of Tilia, Quercus and Car- de cid u ous lime-oak-horn beam for ests. Ar eas oc cu pied by pinus, which grew on more fertile soils and this may indi cate hygrophilous Alnus in creased, sug gest ing the de vel op ment that more land was needed for ara ble fields. The im portance of swampy al der forest with Fraxinus ex cel sior. We suppose of Pinus and Ulmus gradu ally decreased. However, elm was that the observed changes in species compo si tion, mainly the still present on fresh, fertile and moder ately mesotrophic dom i nance of Quercus, de vel oped un der the sub stan tial in soils. Locally, wet alder woods may have grown in depres - uence of human ac tiv ity, and to some ex tent were as so ciated sions, around the lake and on mires. There were slightly with mod erate livestock grazing in woodland. The oc cur- larger areas covered by meadow veg eta tion, in the PG6 pol - rence of trampled and grazed ar eas near the study site is illus - len profile mainly rep re sented by Poaceae, Cyperaceae, trated by pres ence of Polygonum aviculare, Plantago lance- Apiaceae, Brassicaceae, Rubiaceae, Silene-type, Thalictrum olata-type, Potentilla-type, Cerastium-type, Rumex acetosa- and Centaurea nigra-type. Proba bly, there were also rel a - type and Rumex acetosella-type. It is worth mention ing that tively small-cul tivated ar eas and grazed grass lands sit u ated in mediaeval doc u ments re lated to the study area there are close to the lake. It is also pos si ble that de for es ta tion took several place names in dicat ing open oak woods with wood- place be cause of the need to ac quire new res iden tial areas. pas ture practices. For exam ple, the name of the village Nako- How ever, it is dif fi cult to de ter mine the causes of for est de - miady (‘Eichmedien’) (eiche – an oak tree; median – the Pru- struc tion, es pe cially be cause there is no ar chae o log i cal ev i - ssian forest), may in dicate the exis tence of oak wood (Geru- dence for construc tion mate ri als and no anthracological data llis 1922). Addi tion ally, Beckherrn (1885) mentioned the from the Poganowo settle ment complex. area named ‘Damerau’, sit u ated close to the study site since Pollen spectra from LPAZ PG6-3b indi cate the predom i- 1393. Gross (1935) claimed that in medi eval times this name nance of for est com mu ni ties, with Betula play ing the most was commonly used to describe decid u ous forest, mainly signif i cant role, as suggested by its pollen values of up to 60%. with oak trees, where do mes ticated pigs were fat tened on Huntley and Birks (1983) assumed that Betula pollen values acorns, a practice known as pannage. Moreover, accord ing to larger than 50% indi cate Betula-dom i nated wood lands in the Beckherrn (1889), and Wyczó³kowski and Makowiecki land scape. Szal et al. (2014) discussed the problem of the (2009), it can be claimed that the inhab it ants of the medi eval long-last ing dom i nance of Betula in the Iron Age and in the Poganowo settle ment complex raised cattle and horses. Mi gra tion Pe riod, par tic u larly in the area of Lake Salêt, sit u - It is not clear to what degree herbi vores can influ ence ated close to the site inves tigated here. It was con cluded that long-term patterns of plant abundance, dynam ics and dis tri- Betula ex pan sion was con nected with ag ri cul tural prac tices bution (Maron and Crone, 2006). Despite this, some scien - within the settle ment region. In the case of the Poganowo tists (e.g. Flannery, 2003) claim that there is a clear corre- area, the high Betula par tic i pa tion may in di cate that this land lation between oak forest and the intro duc tion of grazing was cleared repeat edly in order to prevent the re-growth of herds into the prime val forests in medi eval times. Faliñski woodland and to enable effec tive crop farm ing. Cereal culti - (1986) described the dom inance of Quercus in Bia³owie¿a vation indi ca tors recorded in the PG6 profile show that For est as a re sult of graz ing. How ever, dif fer ent an imal spe - ANTHROPOGENIC TRANS FOR MA TION OF THE VEG E TA TION 25 cies are likely to have dif fer ent ef fects, with larger an imals The spores not only survive passage through the diges tive likely to cause more physi cal damage by breaking branches, sys tem of an imals, but also in most cases the me chan ical and eat ing sap lings and seedlings, or tram pling the herba ceous chem i cal di ges tion pro cesses also ben e fit the ger mi na tion of veg eta tion. Cat tle and other large her bi vores may have cre - the spores on the dung substrates (Bell, 1983). Krug et al., ated clearings, or at least more open woodland, allow ing light (2004) under lined that ger mina tion, growth, and sporulation to reach the ground and to support a growth of herba ceous occur on freshly depos ited dung. In the next phase, the spores plants on the woodland floor. In the PG6 section there are are dis charged force fully into the air and readily adhere to clear signs of small landscape openness and reduc tions in a nearby vege ta tion. Then, they will be consumed by herbi - tree can opy, leading to understory de velop ment with light- vores, pass through their intes tines, and again will be depos - de mand ing Pteridium aquilinum and Calluna vulgaris. ited with faeces (Bell, 2005). Ac cording to Krug et al. Moreover, in some ar eas where live stock graz ing was sus - (2004), most of the coprophilous fungi are found on mamma - pended in creased flow er ing/re-sprout ing of Corylus avel- lian dung from do mes ticated farm an imals, such as cattle, lana and Populus could occur. horses, and sheep; from wild mam mals, both herbiv o rous Faliñski (1986) noticed that in Bia³owie¿a Forest chan- and carniv o rous; and from birds. ges in spe cies com po si tion oc curred that were as so ci ated From the vari ous stud ies of fossil and mod ern spores of with a reduc tion in grazed areas. The species of open habi tats coprophilous fungi (e.g. van Geel, 2001; Blackford and grad u ally de clined and mesophytic Carpinus re placed Quer- Innes, 2006; Graf and Chmura, 2006; van Geel and Aptroot, cus. This led to the loss of spe cies as semblages typ ical for 2006) it be came clear that the re corded spores in most cases sub con ti nen tal oak woods (Kwiatkowska et al., 1997). The were of strictly lo cal oc cur rence. They were fos silized at, or mainte nance of oak woodlands in the past was also discussed near, the place where they had been produced, or the spores by Szabó (2013). Ac cording to this author, tradi tional forest were depos ited only a short distance from the place where manage ment was not a driving factor in a distri bu tion of vari - sporulation took place. Conse quently, spores of coprophi- ous forest types, while oak wood might be an excep tion. Be- lous fungi indi cate nearby the former pres ence of dung pro- cause of the economic poten tial of acorns, woodland owners duced by herbi vores (van Geel et al., 2003). Accord ing to may have decided to conserve oak woods or even to trans- Lundqvist (1972), rep resen ta tives of the Sordariaceae (As- form other types of forests into oak woods. In ad dition, if comycetes) are mostly coprophilous, partic u larly the genera such oak woods were used as pastures, such manage ment led Sporormiella, Podospora, Sordaria, and Cercophora, which to subse quent changes in woodland compo si tion and struc- are the most common ones among dung fungi (e.g. Krug et al. ture, connected with the preven tion of forest succes sion. It is 2004; van Geel et al., 2007). Moreover, these fungi produce worth mention ing that accord ing to Bobiec (2012), very high thick-walled spores that are well preserved in palynological graz ing pres sure by large un gu late her bi vores sub stan tially prepa ra tions, and can be identi fied and counted alongside lim its oak re gen er a tion. His ob ser va tions of con tem po rary pollen, and this makes it feasi ble to use them in the cate gory feeding sta tions lo cated in Bia³owie¿a Forest in di cate that high of sec ond ary anthropogenic in di ca tors (van Geel et al., 2003; con cen tra tions of bi son ef fec tively pre vent oak re cruit ment. López-Sáez and López-Merino, 2011). More over, López-Sánchez et al. (2014) observed reduc tion in Ejarque et al. (2011) analysed mod ern as sem blages of density of oak seed lings and sap lings, but not of adult trees in pollen and NPP, and found useful rela tion ships with vege ta - ar eas grazed by cat tle. They con cluded that the long-last ing tion and envi ron men tal patterns, as well as with grazing ac - pres ence of ungulates neg a tively af fected oak re gen er a tion, tiv i ties. The study in cluded sam ples from her biv o rous excre- espe cially in pastures with a high stocking rate density. ment in or der to iden tify po ten tial graz ing in di ca tors. These authors empha sized the significant role of coprophilous taxa, Palaeo eco logi cal con text of the fun gal spores par tic u larly of Sporormiella-type and Sordaria-type, as lo cal indi ca tors of herbi vores. The spore cells of Sporormiella spe - re cord cies, which include ob li gate coprophilous species growing In cen tral Eu rope, many hu man ac tiv i ties, e.g. de for es ta- on herbi vore dung (Ahmed and Cain, 1972), are actu ally tion, forest grazing and agri cul ture, significantly altered most often used to study the mid-to- late Ho locene history of landscape struc ture and forest com po sition in the past (e.g. pas to ral ac tiv i ties (e.g. Mighall et al., 2006; Currás et al., Behre, 1988). An im por tant anthropogenic en vi ron men tal 2012). change at or near set tle ment sites is eutrophication, re sult ing Re cently, López-Me rino et al. (2014) indi cated that Cer- from the produc tion of dung by high densi ties of domes ti - cophora-type spores should be also regarded as good indi ca - cated her bi vores (van Geel et al., 2003). Further more, man tors of dung and grazing pressure. Fossil asco spores of the and domes ti cated ani mals are re sponsi ble for a range of new coprophilous Cercophora-type were first rec og nized by van hab i tats, also, the mycoflora of set tle ment sites and the sur- Geel (1978). Accord ing to Ralska-Jasiewiczowa and van rounding ar able land, pas tures and hay-meadows will have Geel (1992), the occur rence of Cercophora-type may be re- had dif fer ent fun gal as sem blages rel a tive to nat u ral hab i tats. lated to increased popu la tions of large wild, and possi ble do- Fungi that in habit or are asso ci ated with the dung of ani - mes ti cated, her bi vores. Cercophora species occur not only mals are known as coprophilous fungi (Krug et al., 2004). on dung, but also on decay ing wood, culms, stems and leaves These fungi are uniquely adapted to herbi vore dung. Bell (Lundqvist, 1972). Moreover, Cugny et al. (2010) reported (1983) concluded that the spores of many coprophilous taxa that Cercophora-type is more frequent in forested sites than are in gested by ani mals feeding on the plants, and then ger- elsewhere, and thus can be used as an indi ca tor of grazing in a mi nate af ter pass ing through the her bi vores’ di ges tive track. wood land. 26 M. SZAL et al.

In our study the record of spores of coprophilous fungi taken into account that over 4,500 fragments of ani mal bones (Sporormiella-type, Sordaria-type and Cercophora-type), were found dur ing the ar chae o log i cal in ves ti ga tions of the in combi na tion with pollen data, was used to identify ani - medi eval sac ri fi cial place at Poganowo (Wyczó³kowski and mal-re lated ag ri cul tural prac tices in the vicin ity of the stud - Makowiecki, 2009) which might have functioned here even ied site. In the first phase of human impact on the vege ta tion in the 13th and 14th centu ries AD (M. Wyczó³kowski, per- (LPAZ PG6-1, Fig. 2) frequently noted pollen of human im - sonal comm.). Almost 90% were horse bones and about 10% pact in di ca tors co in cided with ev i dence of de for es ta tion. The cat tle bones. The au thors esti mated the mini mum num ber of rel atively high peaks of Poaceae, Apiaceae, Ranunculaceae, horse in divid u als at 60. We suppose that cattle and horses as well as nu mer ous Filipendula and Centaurea nigra-type were the domi nant domes ticated spe cies in the study area. pollen (Fig. 2), docu ment a large area of meadows rather than More over, in writ ten sources (Beckherrn, 1885), a place pas tures. There were no Plantago lanceolata, Rumex aceto- called ‘Stiermarkt’ (bo vine/cat tle market) is men tioned and it sella-type, and almost no Rumex acetosa-type pollen, con- is lo cated a few kilo metres from the sampling site. This sid ered by Gaillard et al. (1992) as char ac ter istic of grazed toponym may confirm that grazing cattle played a significant and mown grasslands, and indi ca tors of ani mal husbandry. role in this area in the past, but we cannot exclude an influ - However, the presence of numer ous charred parti cles, high ence of game popu la tions on the vege ta tion. However, typi - fre quen cies of Ar te mi sia, which is an indi ca tor of ruderal cally, the spores of coprophilous fungi are occa sion ally com mu nities and certain amounts of Chenopodiaceae point present in the sedi ments and become very com mon only after to sig nif i cant ac tiv ity within the set tle ment. In spite of the the in tro duc tion of do mes ti cated pas ture an i mals (Graf and fact that human activ ity was intense at this phase, fungal Chmura, 2006). Accord ing to Ejarque et al. (2011), dung-re- spores were very scarce. Only Cercophora-type was pres ent, lated fungi appear in much higher abundance in the excre - but in small num bers and in only a few sam ples. It should be ment of do mes ticated an imals than in the faeces of wild taken into consid er ation that wet local condi tions may have spe cies, but the ex per i men tal re search by Rich ard son (2001) limited both the local devel op ment of fungi and the frequen - showed that it is not possi ble to differ en ti ate between fungal tation of the site by ani mals (therefore, almost no dung may spec tra from do mes tic (cattle, sheep, goat and horse) and have been depos ited). However, based on the pollen and fun- wild (deer, elk) spe cies. gal spore record from the first phase, it is possi ble that agri - It seems logi cal that the in crease in Cercophora-type, culture and ani mal hus bandry did not play a key role in the Sordaria-type and Sporormiella-type that occurs in LPAZ close vi cin ity of the PG6 site. This is under stand able if this PG6-3 (Fig. 2) implies a signif i cant enhance ment in grazing first phase relates to the second phase of occu pa tion of the inten sity and thus an increased popu la tion density of live- hillfort at Poganowo, when an early me di eval sac ri fi cial stock near the inves ti gated site. The higher frequen cies of the place ex isted there, and mainly mag i cal-re li gious rit u als spores of coprophilous fungi coin cide with a notice able in- were performed. Agri cul ture was the most im portant hu man crease of the spores of the para sitic fungus Kretzschmaria ac tiv ity in the me di eval pe riod, and there fore we can say with deusta, reaching a maxi mum frequency of ca. 12%. The data a high de gree of cer tainty that farmlands oc curred at some from Poganowo may show that ani mals dam aged the root dis tance from the in ves ti gated site. This in ter pre ta tion is in system and the lower part of the trees by scraping bark away, agree ment with the ar chae o log i cal ev i dence of an early me - hence al low ing in fec tion by this par a sitic fun gus. More over, dieval human habi ta tion in the Poganowo area. the re sults of the PCA (Fig. 2) show that there is a cor re la tion The first phase was followed by a phase of decreased hu- between dung-related fungi (Sordaria-type and Cercopho- man activ ity (second phase, Fig. 2). However, pollen data ra-type), the wood-rot fungus Kretzschmaria deusta and tree from the PG6 profile indi cate that some new land-use prac- spe cies such as Alnus, Carpinus betulus, Picea abies and tices ap peared in the sec ond phase, but on a small lo cal scale. Quercus. This may in di cate that an i mals browsed mainly in The oc cur rence of Plantago lanceolata, Rumex acetosella- woodland dom inated by the above-mentioned tree species. type, Rumex acetosa-type and Cichoriaceae pollen grains in- Moreover, based on data from the topmost part of the PG6 dicates that some open plant commu ni ties spread, proba bly pollen dia gram (Fig. 2), where frequen cies of Kretzschmaria as a result of grazing. Moreover, presence of spores of the deusta and Quercus change simul ta neously, it could be con- coprophilous Cercophora-type, Sporormiella-type and Sor- cluded that this fungus might infected oaks more often than daria-type may confirm this type of ac tiv ity in the close vi - other trees. However, it should be taken into account that cinity of the studied site. However, it remains an open ques- from the begin ning of the LPAZ PG6-2 the local envi ron - tion whether grazed plots were lo cated in a for est or in areas ment has changed, and the es tablish ment of a local ripar ian outside the forest. and de cid u ous for est oc curred, which per mit ted the pres ence The third phase (LPAZ PG6-3) corre sponds to the pe - of ani mals and, subse quently, spores of coprophilous fungi, riod of in ten si fi ca tion of set tle ment pro cesses that prob a bly as well as Kretzschmaria deusta. started from the late 13th century AD onwards. Pollen data in- As co spores of Kretzschmaria deusta were previ ously dicate changes in the wood land com po sition and show the iden ti fied as Ustulina deusta or as non-pollen microfossil fre quent oc cur rence of hu man im pact in di ca tors. To some ‘Type HdV-44’ by van Geel (1978), who also rec ognized extent, the anthropogenic forest distur bances were proba bly that those spores were produced by a repre sen ta tive of the related to the pas turing of livestock and firewood col lecting. Xylariaceae. The fun gus Kretzschmaria deusta (Hoffm.) The increase of coprophilous fungi sug gests that the live- P.M.D. Mar tin (Ustulina vulgaris Tul. and C. Tul; Ustulina stock frequented the wetlands and fens located close to the deusta Hoffm.) is a tree patho gen that is also called brit tle sam pling site, proba bly more often than before. It should be cinder, and it is found in tem perate regions of the north ern ANTHROPOGENIC TRANS FOR MA TION OF THE VEG E TA TION 27 hemisphere (Rogers and Ju, 1998; van Geel et al., 2013). It REF ER ENCES causes soft-rot in wood, breaking down both cel lulose and Ahmed, S.E., Cain, R.F., 1972. Re vi sion of the gen era Sporormia lignin, and it decays the trunk and roots of living trees. The and Sporormiella. Ca na dian Jour nal of Bot any 50, 419–477. fungus contin ues to decay wood after the host tree has died, Beckherrn, C., 1880. Rastenburg historisch-topographisch darge- mak ing Kretzschmaria deusta a fac ul ta tive par a site (Rog ers stellt, Verlag Schlemm, Kêtrzyn. and Ju, 1998). This fungus has a wide host range among Beckherrn, C., 1885. Verzeichnis der die Stadt Rastenburg betre- broadleaved trees in Europe, includ ing Acer, Aesculus, Al- ffenden Urkunden, Altpreußische Monatsschrift 22, 505–605. nus, Betula, Carpinus, Castanea, Fagus, Fraxinus, Populus, Beckherrn, C., 1889. Schaffer’s Chronik von Rastenburg. Druck Quercus, Salix, Tilia and Ulmus. It has been rarely noted on und Verlag von W. Kowalski, Rastenburg. Abies and Taxus (Wilkins, 1934). Al though there is no evi - Behre, K.E., 1981. The in ter pre ta tion of anthropogenic in di ca tors in dence that Kretzschmaria oc curs on Picea abies, po ten tially pol len di a grams. Pollen and Spores 23, 2, 225–245. it can invade this tree (J. Rogers, personal comm.). More- Behre, K.E., 1988. The role of man in Eu ro pean veg e ta tion his tory. over, Schwarze (2007) showed ex peri men tally the abil ity of In: Hunt ley B., Webb T. (eds), Veg e ta tion his tory, 633–672. Kluwer Ac a demic Pub lish ers, Dordrecht. Kretzschmaria deusta to cause de cay in liv ing spruce trees. Bell, A., 1983. Dung fungi. An il lus trated guide to coprophilous Chlebicki (2008) re ported that some xylariaceous fungi oc- fungi in New Zealand Victo ria Univer sity Press, Wellington. cur on Picea abies in Poland. Wilkins (1934) summa rized Bell, A., 2005. An il lustrated guide to coprophilous fungi Ascomy- many papers dealing with the patho genic ity of Kretzsch- cetes of Aus tra lia. CBS Biodiversity Series No. 3, Centraal- maria. It is an oppor tunis tic para site that takes advan tage of bu reau voor Schimmelcultures, Utrecht. already signif i cantly dam aged trees (Rogers and Ju, 1998). Berglund, B.E., Ralska-Jasiewiczowa, M., 1986. Pol len analy sis and Open wounds on tree trunks are needed to account for major pol len di a grams. In: Berglund, B.E., Ralska-Jasiewiczowa M. expan sions of the fun gus. Moreover, attacks by Kretzsch- (eds), Hand book of Ho lo cene Palaeo ec ol ogy and Palaeohy- maria are usually asso ci ated with infec tions by some other drology, 455–484.Wiley and Sons Ltd., Chichester-To ronto. fungi (Wilkins, 1934). Beug, H.J., 2004. Leitfaden der Pollenbestimmung für Mittel- europa und angrenzende Gebiete, Verlag Dr. Friedrich Pfeil, München. CON CLU SIONS Birks, H.J.B., 2003. Quan ti ta tive palaeoenvironmental re con struc - tions from Holo cene bi o log i cal data. In: Mackay A., Battarbee Inte gra tion of data from pollen, non-pollen palyno- R.W., Birks H.J.B., Oldfield, F. (eds), Global Change in the morph and microcharcoal parti cle anal yses proved to be a Holo cene, 342–357. Verlag Hodder Ar nold, Lon don. useful tool for assess ing human occu pa tion of the Poganowo Blackford, J.J., 2000, Char coal frag ments in surface sam ples fol - area. It is possi ble to distin guish three phases of vege ta tion low ing a fire and the im pli ca tions for in ter pre ta tion of subfossil dynam ics connected with local human impact dated to the char coal data. Palaeo ge ogra phy, Palaeoclimatology, Palaeo- Mid dle Ages (ca 10th–16th c.). The old est, first phase, corre - ec ol ogy 164, 33–42. sponds to very inten sive human ac tiv ity within the Poga- Blackford, J.J., Innes, J.B., 2006. Link ing cur rent en vi ronments and nowo set tlement complex and re lates to rel atively large open processes to fun gal spore assem blages: surface NPM data from areas created by regu lar human-caused burning. This region woodland en vi ron ments. Re view of Palaeobotany and Palyno- logy 141, 1, 179–187. was, to a con sid er able ex tent, cov ered by mead ows and Bobiec, A., 2012. Bia³owie¿a Pri meval For est as a rem nant of cul - ruderal com mu nities, and only small plots were used as fields tur ally mod i fied an cient for est. Eu ro pean Jour nal of For est Re - and pastures. The second phase was a period when human ac - search 131, 5, 1269–1285. tiv ity de creased and wood land re gen er a tion took place. Fi- Burchardt, L., Pawlik-Skowroñska, B., 2005. Zakwity sinic – kon- nally, the third phase reflects response of a local vege ta tion to kurencja miêdzygatunkowa i œrodowiskowe zagro¿enie. Wia- hu man ac tiv ity dated to the 13th–16th cen tu ries AD. In creas - domoœci Botaniczne 49,1–2, 39–49. ing ruderalisation and intensification of human impact were Chlebicki, A., 2008. Some over looked and rare xylariaceous fungi mani fested by indi ca tor species of trampled and grazed vege - from Poland. Pol ish Botan i cal Jour nal 53, 1, 69–70. ta tion. The me di eval man age ment most prob a bly pro moted Connor, S.E., Araújo, J., van der Knaap, W.O., van Leeuwen, oaks and pastured oak woods. During the youngest part of J.F.N., 2012. A long-term per spec tive on bio mass burning in this phase, a signif i cant spread of Quercus was ac com pa nied the Serra da Estrela, Portu gal. Quater nary Science Reviews 55, 114–124. by an increase in spores of coprophilous fungi and the para - Cugny, C., Mazier, F., Ga lop, D., 2010. Modern and fos sil non-pol - sitic fungus Kretzschmaria deusta. We suppose that grazing len palynomorphs from the Basque mountains (west ern Pyr e - ani mals caused damage to the roots and trunks of oak trees, nees, France): the use of coprophilous fungi to re con struct thus trig ger ing in fec tions with Kretzschmaria deusta. pas to ral ac tiv ity. Veg e ta tion His tory and Archaeobotany 19, 391–408. Ac knowl edge ments Currás, A., Zamora, L., Reed, J.M., García-Soto, E., Ferrero, S., Armangol, X., Mezquita-Joanes, F., Marqués, M.A., Riera, S., The study was part-fi nanced by the Pol ish Na tional Re search Julià, R., 2012. Cli mate change and human impact in cen tral Coun cil (NCN), through pro ject no. DEC-2011/01/B/HS3/04167. Spain dur ing Ro man times: high-res olu tion multi-proxy anal y- We would like to ex press our grat i tude to Bas van Geel and Lionello sis of a tufa lake re cord (Somolinos, 1,280 m asl). Catena 89, Morandi for pro vid ing ac cess to some of the spe cial ized lit er a ture. 31–53. We would like to thank Jack D. Rog ers for valu able sug ges tions and Ejarque, A., Miras, Y., Riera, S., 2011. Pol len and non-pol len comments on host-par a site as soci a tion. We are grate ful to Bas van palynomorph in di ca tors of veg e ta tion and high land graz ing ac - Geel and an anon y mous reviewer for their sugges tions in im prov ing tiv i ties ob tained from mod ern sur face and dung datasets in the this manu script. east ern Pyr e nees. Re view of Palaeobotany and Palynology 28 M. SZAL et al.

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