Vegetation History and Archaeobotany

, Volume 14, Issue 4, pp 518–527 | Cite as

The first subfossil records of Urtica kioviensis Rogow. and their consequences for palaeoecological interpretations

Original Article

Abstract

Among plant remains from Mesolithic layers dating from 9249 to 7779 B.C. at the excavation site of Friesack IV in north-eastern Germany, nutlets of Urtica kioviensis were identified. Morphological studies have shown that they clearly differed from all other European Urtica species investigated. In contrast, pollen morphological investigations revealed only slight differences between the central European Urtica species, which could hardly have been noticed during routine or normal pollen analyses. The records of U. kioviensis nutlets are the first subfossil finds reported and prove the indigenous status of this taxon in north-eastern Germany. The records are discussed in the context of the overall species spectrum of the Mesolithic layers and consequences for the interpretation of pollen analytical studies concerning human impact are pointed out.

Keywords

Biogeography Mesolithic Macro remains Pollen morphology North-eastern Germany 

Introduction

Extensive archaeological excavations have been carried out at the Mesolithic-Neolithic settlement site near Friesack (Havelland, north-eastern Germany) by the Museum für Ur- und Frühgeschichte at Potsdam, between 1977 and 1989 (Fig. 1). Although the site has been known since the beginning of the 20th century (Schneider 1932), the first palaeobotanical investigations there were made by Stoller (1927) and later by Bertsch (1953), to estimate the age of the cultural layers long before the radiocarbon method came into existence.
Fig. 1

Location of the excavation site in the Rhin-Luch near Friesack, Brandenburg, Germany. The mire landscape of the Rhin-Luch is marked with dashed signs (inserted map after Gramsch 2000)

Today the age of the layers is known in detail and based on numerous radiocarbon dates. Accordingly, the many layered sequence of sediments, consisting of sands, organic sands and peat muds indicating many Mesolithic occupations, dates from about 9250 to 5550 B.C., thus from the Pre-boreal to the Atlantic (Gramsch 1979, 1981, 2000; Görsdorf and Gramsch 2004). In the course of the excavations, samples for pollen and macrofossil analysis were taken. Whereas the results of pollen analysis have already partly been published (Kloss 1987a,b), macrofossil analysis did not start until 2002.

The investigated material originates from the early Mesolithic layers of an excavation section through the bank of the small river Friesacker Rhin or its early Holocene predecessor. Due to favourable sedimentation conditions in the littoral of the Rhin the preservation of the majority of the samples is excellent, so even string-, cord-, and net fragments made of Salix bast were preserved under waterlogged conditions (Kernchen and Gramsch 1989; Körber-Grohne 1995). In the course of macrofossil analysis, some Urtica nutlets were recorded which in their morphology differed considerably from those of U. dioica and U. urens, the two species quite common in the region today. A morphological comparison of recent nutlets of European Urtica species showed that the subfossil Friesack nutlets only matched those of U. kioviensis. As well as the morphology of fruits and pollen, the distribution area and ecology, and results of macro remains analyses are described and some implications for palaeoecological interpretations discussed below.

Study area

The Mesolithic-Neolithic site of Friesack IV is situated in the Unteres Rhinluch, some 60 km northwest of Berlin on part of the north German lowland (Fig. 1). It is a (formerly) swampy area with extended fens, developed after the last glaciation (Weichselian) at the junction of the ‘Warsaw-Berlin’ and the ‘Torun-Eberswalde’ glacial valleys which led the meltwater through to the North Sea via the river Elbe. Bordered by the somewhat higher land of ground moraines and terminal moraines of the last glaciation, it has been a landscape widely covered by shallow lakes just after the deglaciation (Gramsch 2000). During the Late-glacial, inland dunes were formed on and of fluvial glacial and ground moraine sediments which are like islands in the lake landscape. They have been suitable and favoured as settlement places since Mesolithic times, which was also the case at Friesack IV. Minerogenic material also containing the Mesolithic finds was washed from these sandy mounds downhill into the shallow lakes and covered by muds and peat during phases without human activity. Mire development itself started at the beginning of the Holocene by means of lake infilling, although large scale peat growth did not start until about 6000 B.C. The peat layer finally reached 2–3 m on average, but in depressions up to 15 m and more (Mundel et al. 1983).

This trackless landscape existed until the 18th century, when peat exploration and exploitation started. Drainage ditches, dams, sluices, bridges and channels were built to lower the groundwater level, to excavate peat and to transport the excavated peat mainly to Berlin for heating. The last improvement programme was conducted from 1970–1982 by massively lowering the groundwater level with the aim of gaining large areas for farmland, mainly for pastures and meadows, partly for arable farming. Therefore archaeological sites such as Friesack IV and their preservation were heavily endangered, so the decision for a systematic excavation was made, starting in 1978 after prospection in 1977 (Gramsch 2000).

Morphology and recent distribution of Urtica kioviensis

Urtica kioviensis (Fig. 2) is monoecious and therefore easily distinguished from U. dioica when flowering. However, there are other features which allow easy identification in the field. The whole plant, especially the leaves, is characterised by a shining light green appearance, and the basal part of the stem is creeping and rooting at the lower nodes. The upper part of the plant is ascending (Fig. 2). The upper stipules of Urtica kioviensis leaves are joined at the base or up to the middle part of it. Even at 10× magnification the differing hairiness of U. kioviensis is striking, consisting almost completely of stinging hairs (Fig. 3). Bristle hairs, which are common in U. dioica, are rare.
Fig. 2

Urtica kioviensis (from Jäger et al. 1988). The scale is 1 cm

Fig. 3

Stinging hairs of Urtica kioviensis. The scale is 1 mm

U. kioviensis was recognised as an autonomous taxon by means of specimens collected in the vicinity of Kiev (hence the epithet!) about 160 years ago by the Russian botanist A.S. Rogowitsch (1843). Detailed information on its present distribution and phytosociological status are provided by Konczak et al. (1968) and most recently by Wollert et al. (2003). Accordingly, U. kioviensis is an element of the thermophilic floodplain and lowland landscapes, and shows a sub-continental, pontic-pannonic distribution (Fig. 4). Its disjunctive distribution spans a sickle-shaped area from eastern Europe westwards to the main distribution area in Hungary and the northwestern adjacent countries (Pannonic lowlands), and further northwards to northeast Germany (mainly the Havelland area in the State of Brandenburg and a few records in Mecklenburg-Vorpommern, Fig. 5). In 1994 Faurholdt and Schou reported the first finds from Denmark, which are the northernmost records so far. The southernmost and very isolated occurrence on the Hula plain in Israel is remarkable (Feinbrun-Dothan and Danin 1991). However, the indigenous nature of this latter population is doubted by Wollert et al. (2003).
Fig. 4

Total range of Urtica kioviensis (from Wollert et al. 2003). The only record outside Europe (Hula plain, Israel) is not displayed in the map

Fig. 5

Distribution of Urtica kioviensis in north-eastern Germany (after Benkert et al. 1996 and Wollert et al. 2003; supplemented). +  extinct, F  subfossil finds near Friesack

In the German flora, U. kioviensis was first mentioned by Zólyomi (1936) after the revision of Urtica herbarium samples from north-eastern Germany. Subsequent observations (for example, Scholz and Sukopp 1960; Konczak et al. 1968; Benkert et al. 1996) have shown that U. kioviensis occurs regularly in the Berlin-Potsdam area as well as in the Havel valley between Berlin and the river Elbe. Outside of the Havel region there are only a few other records in north-eastern Germany (Fig. 5). Because of the concentration of records along the river Havel, Müller-Stoll et al. (1962) classified U. kioviensis as a river corridor plant. It grows mainly in wet facies of Phragmites-, Phalaris- and Carex riparia reed swamps as well as in wet willow scrubs (Alno-Salicetum cinereae) and different Alnetum facies (Gutte et al. 1973; Wollert et al. 2003, in prep.). Sometimes U. kioviensis is also found in floodplain woods of the alliance Alno-Ulmion (Emrović et al. 1964; Schoknecht 1988; Oberdorfer 2001).

Material and methods

Approximately 500 samples from all excavation trenches and layers were systematically collected in the field during the excavations in the years 1978–1989. The first 25 samples for macrofossil analysis were selected on the basis of certain identification and linkage to a distinct excavation layer. They proved sufficient to answer the geobotanical questions discussed in this study. However, more samples are due to be analysed for a comprehensive investigation of the botanical macrofossil remains from the Mesolithic site. The majority of the first samples (21) has already been sieved using mesh sizes of 2, 1, 0.5 and 0.2 mm, before being transported to Wilhelmshaven. The other four samples were treated just prior to the analysis (mesh sizes: 3.15, 0.71 and 0.25 mm) and volumes of the not previously treated samples were between ca. 2.5 and 5 l. Whereas the 21 pre-processed samples were analysed in total, the other four samples were subsampled. Here the coarse fraction was analysed in total, the middle and fine fractions partially.

The remains were sorted and identified using a stereo microscope with 10–50× magnification. In some cases a compound microscope was used (500x). The results were stored in an archaeobotanical database (Kreuz and Schäfer 2002). Photos and measurements of the subfossil seeds were made both on wet material and after conservation with polyethyleneglycol according to Kučan (1991). Following the identification of the conspicuous nutlets, a pollen morphological investigation was carried out in order to detect possible morphological differences between U. dioica/urens and U. kioviensis.

Investigations of recent U. kioviensis nutlets were carried out with specimens from the herbarium of the University of Göttingen and with plants collected in Sacrow near Berlin. Nutlets of other Urtica species were taken from the reference collection of the NIhK at Wilhelmshaven and from herbarium specimens provided by the Botanic Garden and Botanical Museum of Berlin-Dahlem. Pollen material was obtained from dried herbarium specimens of U. kioviensis collected by the author V. Kummer and originating from three different locations in north-eastern Germany: Sacrow, 22.07.1992, Grid Ref 3544/4; Berlin-Kohlhasenbrück, 17.07.1991, Grid Ref 3644/2; Prietzen, 17.09.1992, Grid Ref 3239/4.

The pollen material was acetolyzed and mounted in glycerine jelly. Pollen measurements were carried out at a total magnification of 1250x using a light microscope. At least 100 measurements of pollen size per specimen were made to give a reasonable degree of statistical reliability. Terminology follows Beug (2004).

Samples for radiocarbon dating were taken during the excavations in the 1980s and were dated at the former Zentralinstitut für Alte Geschichte und Archäologie, Academy of Science of the German Democratic Republic, Berlin. The dating material was charcoal or charred wood. The conventional14C data (Gramsch 2000; Görsdorf and Gramsch 2004) were calibrated with Calib 5.0.0 in conjunction with Stuiver and Reimer (1993), based on IntCal04 (Reimer et al. 2004).

Results and discussion

Dating the subfossil finds

Out of the 25 samples analysed, 12 could be dated by means of correlation to radiocarbon dated excavation horizons. In total 15 14C dates were available (Table 1). They range from 9249 to 7749 B.C., thus from the middle/late Pre-boreal (Firbas IV) to the middle of the later Boreal (Firbas Vb) .
Table 1

List of radiocarbon data (compiled from Görsdorf and Gramsch 2004)

Lab. Nr

14C date uncal B.P.

Calibrated age (2 σ) B.C.

Bln-3019

9640 ± 70

9249 – 8813

Bln-3020

9640 ± 60

9244 – 8826

Bln-3025

9340 ± 70

8775 – 8348

Bln-3009

9240 ± 70

8625 – 8297

Bln-3024

9180 ± 70

8593 – 8276

Bln-3000

9220 ± 60

8603 – 8296

Bln-3008

9040 ± 70

8443 – 7969

Bln-3027

9040 ± 70

8443 – 7969

Bln-3023

9040 ± 60

8424 – 7976

Bln-3017

9010 ± 70

8334 – 7957

Bln-3006

9000 ± 70

8319 – 7955

Bln-3013

8980 ± 60

8291 – 7965

Bln-3014

8980 ± 60

8291 – 7965

Bln-3003

8940 ± 60

8284 – 7879

Bln-3011

8840 ± 60

8220 – 7749

Fossil fruits of Urtica kioviensis

The fruits of U. kioviensis are nutlets, whose pericarps closely adhere to the seeds and are therefore often referred to as achenes. The nutlets appear pointed-elliptic in outline with their greatest width at the equator (Figs. 67). Since the nutlets are more or less compressed, their thickness averages only 0.45 mm. Further size measurements of recent and subfossil material are given in Table 2.
Table 2

Size measurements of recent and fossil nutlets of Urtica kioviensis and U. dioica (n = number of nutlets measured)

Specimen

Length × breadth (in mm)

L/B ratio

U. kioviensis, recent

herbarium, Göttingen

n=13

1.50–1.62–1.80 × 0.80–0.87–1.00

1.67–1.88–2.25

U. kioviensis, recent

Sacrow

n=40

1.50–1.66–1.90 × 0.70–0.87–1.00

1.68–1.91–2.31

U. kioviensis, subfossil

Friesack

n=50

1.40–1.68–2.00 × 0.70–0.86–1.00

1.60–1.96–2.43

U. kioviensis, subfossil

Friesack (PEG)

n=24

1.40–1.69–1.80 × 0.70–0.88–1.00

1.56–1.93–2.25

U. dioica, recent

ref. collection NIhK

n=40

1.00–1.16–1.40 × 0.70–0.81–0.90

1.22–1.45–1.75

U. dioica, subfossil

Friesack

n=50

0.95–1.08–1.20 × 0.60–0.71–0.80

1.31–1.53–1.75

Fig. 6

Subfossil nutlets of Urtica kioviensis and U. dioica recorded in the samples from Friesack. The morphological separation of subfossil nutlets of the two species is clearly visible

Fig. 7

Comparison of recent nutlets of selected European Urtica species. A subfossil nutlet of Urtica kioviensis is included. From left to right: Urtica atrovirens, U. membranacea, U. dioica, U. kioviensis, U. kioviensis (subfossil), U. morifolia, U. urens, U. cannabina, U. pilulifera

At first glance, nutlets of U. kioviensis seem to be rather similar to those of U. dioica. However, they can be separated clearly. The most conspicuous distinguishing feature apart from the pointed-elliptic outline is the noticeably greater length which averages about 1.65 mm, in contrast to the much shorter and slightly ovate nutlets of U. dioica which were also recorded in the fossil samples (Table 2). The size measurements that were made correspond well to the specifications cited by Chrtek (1979a), Gel’tman (1988) and Faurholdt and Schou (1994). Although U. kioviensis had been described in several publications since 1843, in fact Chrtek was the first author to adopt fruit measurements for identification of the central European Urtica species (Chrtek 1979a, 2002).

A comparison of recent nutlets of selected European Urtica species including the subfossil find of U. kioviensis is displayed in Fig. 7. It shows that the pointed-elliptic subfossil nutlets found in the Friesack samples only match those of U. kioviensis. The distinct identification corresponds with its isolated taxonomic status within the section Urtica (Table 3). The same is true for the other isolated nettle species U. cannabina, U. urens, U. pilulifera and U. morifolia, whose nutlets are also easily recognised by means of their size and shape. Hardly distinguishable from each other by means of nutlets are the taxa within the subsect. Urtica, which all are like those of U. dioica, apart from those of U. membranacea.
Table 3

Taxonomy and distribution of European Urtica species (after Chrtek 1979b; Gel’tman 1988, 1993; Paiva 1993)

Taxon names

Important synonyms

Distribution

Subgen. Urtica

  

Sect. Urtica

  

Subsect. Kiovienses Geltm.

  

   U. kioviensis Rogow.

U. radicans Bolla

U. bolla Kanitz

central and eastern Europe

Subsect. Urtica

  

   Urtica dioica L.

 

throughout Europe

   U. sondenii (Simm.) Avror. ex Geltm.

U. dioica subsp. sondenii (Simm.) Hyl

northern and eastern Europe

   U. pubescens Ledeb.

U. dioica subsp. pubescens (Ledeb.) Domin

Russia

   U. galeopsifolia Wierzb. ex Opiz

U. dioica subsp. galeopsifolia (Wierzb. ex Opiz) Chrtek

western, central and eastern Europe

   U. atrovirens Req. ex Loisel.

 

central Mediterranean

   U. bianorii (Knoche) Paiva

U. atrovirens subsp. bianorii (Knoche) Font Quer & Garcias Font

Balearic Islands

   U. membranacea Poir. in Lam.

U. dubia Forskal

   U. caudata Vahl

Mediterranean and western Europe

Subsect. Cannabinae Geltm.

  

   U. cannabina L.

 

Russia and Ukraine

Sect. Urentes Geltm.

  

   U. urens L.

 

throughout Europe

Subgen. Sarcourtica Chrtek

  

   U. pilulifera L.

U. dodartii L

Mediterranean

Subgen. Dendrourtica Chrtek

  

   U. rupestris Guss.

 

Sicily

   U. morifolia Poir.

 

Canary Islands, Azores

Pollen morphology

Pollen grains of U. kioviensis are (2-) 3 (-4)-zonoporate, rarely mono- or 5-porate. The circular pores (diameter ca. 1.5–2.0 μm) slightly protrude and a more or less distinct annulus is visible. The exine is thin, and a differentiation of exine layers is not distinguishable in light microscopy. The sculpturing is scabrate, although this is only visible with phase contrast lighting (Fig. 8). The distribution and size of the scabrae show a great variety between different specimens and even within a specimen, although a blotchy pattern was often observed. The pollen grains are spheroidal with a P/E ratio of about 0.9. Measurements range from 14–18 μm (polar axis) and 14–22 μm (equatorial axis) respectively. Details of pollen morphological studies are presented in Table 4.
Table 4

Size measurements and variation of pore numbers in different specimen of Urtica kioviensis (P  polar axis; E  equatorial axis; n  number of pollen grains examined)

Specimen

Size (in μm)

P/E ratio

Number of pores

U. kioviensis, Sacrow n=250

P 14.1–16.0–18.2

E 14.0–17.5–22.0

0.92

1 pore:

2 pores:

3 pores:

1

30

219

U. kioviensis, Berlin n=100

P 14.0–16.0–18.0

E 16.0–18.1–21.0

0.88

2 pores:

3 pores:

4 pores:

1

95

4

U. kioviensis, Prietzen n=150

P 13.7–15.5–18.2

E 14.6–17.3–21.8

0.89

2 pores:

3 pores:

4 pores:

5 pores:

3

119

27

1

Fig. 8

Recent pollen grains of Urtica kioviensis (1250×, phase contrast). Polar view, high focus with scabrae clearly visible (left), polar view, cross section (middle), equatorial view, high focus with porus and distinct annulus (right)

U. kioviensis has been investigated in the context of pollen morphological studies only once before (Tarnavaschi et al. 1967), although literature on European Urticaceae pollen morphology is plentiful (e.g. Punt and Malotaux 1984; Beug 2004). The most thorough work has been done by Sorsa and Huttunen (1975), which covers not only most European taxa but also species from other continents. However, in contrast to the nutlets of Urtica, the relationships between taxonomy and morphology of Urtica pollen grains are less pronounced, apart from the large periporate pollen grains of U. pilulifera, which separate it off as a member of the subgenus Sarcourtica.

The microscopic observations and measurements of U. kioviensis pollen grains made during this investigation have shown that its main features, number of pores, size and sculpturing, correspond to the features of most of the taxa belonging to the subgenus Urtica. Therefore a pollen analytical separation of U. kioviensis pollen grains by means of light microscopy is not suggested to be possible. This is particularly true for the case of U. dioica pollen grains which share a distinct anulus with U. kioviensis grains. Some authors separate U. urens from U. dioica pollen grains on the basis of different distribution of scabrae (for example Punt and Malotaux 1984; Fægri and Iversen 1989; Chester and Raine 2001). However, because of considerable variations from distant positioned and regularly distributed scabrae to blotchy patches of scabrae in U. kioviensis grains, a pollen analytical differentiation from U. urens is also not suggested.

Noteworthy is the regular occurrence of diporate pollen grains in Urtica kioviensis which amounts up to 12% in one specimen. Diporate pollen grains as an expression of a phylogenetic trend of apertural evolution are common in Urticaceae (Thanikaimoni 1986), for example in Pilea spp., but rarely observed in Urtica. The records of diporate U. kioviensis pollen grains possibly mirror the special taxonomic status within the section Urtica but, again, cannot be employed for pollen analytical identification due to the low statistical reliability of this feature.

Details of the overall species spectrum

Out of 25 samples from the Friesack site, a total of about 18200 botanical macrofossil remains were recorded and have been assigned to about 110 different taxa.

The dominating species range found was typical of floating-leaved communities and bank reed swamps (Nymphaeion, Phragmition and Bolboschoenion) with taxa such as Nuphar, Nymphaea, Typha spp., Sparganium spp. and Schoenoplectus spp. (Table 5). Most of the taxa of the reed communities are also typical of the present-day environment of U. kioviensis in north-eastern Germany where it is often found in Phalaridetum arundinaceae (reed canary-grass) or Phragmitetum australis (common reed) associations along the river Havel or at the fringes of its lakes (Konczak et al. 1968; Burkart 1998). Other typical habitats are sedge or bulrush communities, such as Cicuto-Caricetum pseudocyperi, Caricetum ripariae or Typhetum latifoliae (Hilbig and Reichhoff 1974; Wollert et al. 2003).
Table 5

Selected taxa of the overall species spectrum of the Friesack site. Charred remains were also recorded from taxa marked with an asterisk

 

Number

Presence (%)

Aquatic species

Nuphar lutea*

811

88

Nymphaea cf. alba*

263

84

Hydrosere

Carex pseudocyperus

211

40

Cladium mariscus

842

72

Lycopus europaeus

214

68

Lysimachia thyrsiflora/vulgaris

58

60

Menyanthes trifoliata

220

84

Schoenoplectus lacustris

489

84

Sparganium spp.

4

16

Stachys palustris

120

64

Typha angustifolia

116

48

Typha latifolia*

1555

76

Urtica kioviensis

321

84

Pioneers of shores and banks

Myosoton aquaticum

1255

72

Polygonum lapathifolium agg.

745

72

Woodland; collected species

Corylus avellana*

192

72

Fragaria vesca

9

20

Ruderals

Atriplex spp.

24

16

Capsella bursa-pastoris*

95

40

Chenopodium album agg.

2090

72

Chenopodium ficifolium

14

12

Chenopodium glaucum/rubrum

3809

64

Chenopodium polyspermum

2

8

Erodium cicutarium

28

28

Eupatorium cannabinum

146

56

Mentha aquatica/arvensis

133

68

Plantago major

31

12

Potentilla argentea-type*

408

69

Urtica dioica

338

88

Valerianella locusta

49

48

In addition to these hydrosere taxa, some plant remains were found which might indicate human activity at the Mesolithic site. These are first and foremost collected taxa such as Fragaria vesca (strawberry) and Corylus avellana (hazelnut), the latter often found as charred nutshell fragments. As well as these, there are records of possible anthropogenic indicators suggesting ruderal habitats such as Chenopodium album agg., Erodium cicutarium, Atriplex spp., Valerianella locusta, Plantago major and Urtica dioica (Table 5).

However, classification of the latter proves problematic, since U. dioica is also a natural component of reed communities where it can grow next to U. kioviensis. The same holds true for some Chenopodiaceae taxa. In addition to Chenopodium album, C. glaucum/rubrum, C. ficifolium and C. polyspermum were also recorded — species with their natural habitats on river banks. For comparison, see also investigations of Mesolithic sites in river valleys, such as Behling and Street (1999), Bos and Urz (2003) and Knörzer and Meurers-Balke (1999).

Whereas the majority of the macrofossil remains was preserved uncharred, several records of charred seeds of Nuphar lutea (9249 to 7749 B.C.) and Nymphaea alba (9249 to 8813 B.C.), which are possibly linked with human activity at this site, are noteworthy. Charred seeds of N. lutea were also found for example at the early Neolithic site of Hoge Vaart in the Netherlands (Brinkkemper et al. 1999) and seeds of the related species N. pumilum at the Mesolithic site of Halsskov in Denmark (Kubiak-Martens 2002). The use of Nymphaeaceae seeds for food is well known especially by indigenous peoples in North America and Australia for whom they serve as an important starch source (for more details, see discussion in Kubiak-Martens 2002, p. 29).

A detailed and comprehensive presentation of vegetation development and plant use of the Mesolithic site of Friesack will be given after completion of macrofossil analyses and archaeological evaluation of the excavations.

Apart from macrofossil remains connected with Mesolithic activity there are also notable finds from the Friesack site from a floral historical point of view. These are cuticle fragments from Viscum leaves (8220 to 7749 B.C.) and seeds of Cornus sanguinea (8319 to 7955 B.C.), as well as Viburnum opulus (8220 to 7749 B.C.), which provide valuable information on the early Holocene immigration of shrubs in north-eastern Germany (cf. Brande 1993). In addition the following, in fossil samples some normally rare taxa were also recorded for the Boreal at Friesack IV: Arctostaphylos uva-ursi, Selaginella selaginoides, Holosteum umbellatum and Potentilla supina.

Conclusions

The morphological studies have shown that fruits of U. kioviensis are clearly different not only from the other central European indigenous taxa such as U. urens and U. dioica, but also from all other European Urtica species distributed today. Therefore U. kioviensis can be identified to species level by means of its nutlets. It is suggested that this easily recognisable feature should be added to current floras and identification keys. In contrast, pollen grains of U. kioviensis and U. dioica cannot be separated exactly, and it is also not suggested that U. kioviensis from U. urens should be separated by means of pollen analysis. Vegetation historians and archaeobotanists therefore have to rely on the recovery of fruits.

Urtica kioviensis was recorded with a high presence (84% of the samples) covering the whole range of time represented by them, 9249–7749 B.C. Thus Urtica kioviensis has been indigenous in north-eastern Germany since the late Pre-boreal at the latest and was present in the Rhinluch at least during the major part of the early Holocene. In contrast to records in the Havelland region, there is no current evidence for U. kioviensis growing in the Rhinluch today, mainly due to the present landscape, which has been drained and totally changed by human activity.

The disjunctive distribution area of U. kioviensis leaves some questions unresolved so far, such as the status of the record in Israel (Wollert et al. 2003). Above all, nothing is known about the past distribution area for which now Friesack IV is to our knowledge the first subfossil evidence. With the identification details to hand, U. kioviensis nutlets can clearly be identified in future studies as well as in revisions of material already investigated. This will provide further information about the past distribution of U. kioviensis and today’s distribution gaps.

The occurrence of U. kioviensis also has consequences for the evaluation of human impact in pollen diagrams. In contrast to U. dioica, U. kioviensis is an oligo- to mesohemerobic species, that is, it does not occur in habitats strongly influenced by humans. Since pollen analytical separation of the two nettle species is not possible, care has to be taken with the interpretation of pollen diagrams from the distribution area of U. kioviensis. If sampling took place in the vicinity of lakes or river systems, increased percentages of Urtica do not necessarily indicate human impact. This is particularly true for riverine sampling sites in the Pannonic lowlands which is the main distribution area, but applies also to the whole distribution range.

When investigating Friesack vegetation development by pollen analysis, Kloss (1987a) took this carefully into consideration without having the evidence of macro remains. Now it can be shown that his considerations were right.

Notes

Acknowledgement

The authors wish to thank B. Gramsch (Potsdam), M. Burkart, M. Ristow, K. Tielbörger (all Potsdam University), B. Seitz (TU Berlin), E. Welk (Halle University) and H. Wollert (Teterow) for providing of literature, record data and herbarium material. Many thanks also to H. Nowak-Krawietz (BGBM Berlin-Dahlem) for arranging the loan of Urtica herbarium specimens. A special thank to R. Kiepe and T. Reiser (NIhK Wilhelmshaven) for extensive technical assistance.

References

  1. Behling, H., Street, M. (1999). Palaeoecological studies at the Mesolithic site at Bedburg-Königshoven near Cologne, Germany. Vegetation History and Archaeobotany, 8, 273–285CrossRefGoogle Scholar
  2. Benkert, D., Fukarek, F., Korsch, H. (1996). Verbreitungsatlas der Farn- und Blütenpflanzen Ostdeutschlands. Fischer, JenaGoogle Scholar
  3. Bertsch, K. (1953). Geschichte des deutschen Waldes, 4th ed. Fischer, JenaGoogle Scholar
  4. Beug, H.-J. (2004). Leitfaden der Pollenbestimmung. Pfeil, MünchenGoogle Scholar
  5. Bos, J.A.A., Urz, R. (2003). Late Glacial and early Holocene environment in the middle Lahn river valley (Hessen, central-west Germany) and the local impact of early Mesolithic people - pollen and macrofossil evidence. Vegetation History and Archaeobotany, 12, 19–36CrossRefGoogle Scholar
  6. Brande, A. (1993). Die Entwicklung der Dendroflora in Brandenburg seit der Eiszeit. Beiträge zur Gehölzkunde, 1993, 77–84Google Scholar
  7. Brinkkemper, O., Hogestijn, W.-J., Peeters, H., Visser, D., Whitton, C. (1999). The early Neolithic site at Hoge Vaart, Almere, the Netherlands, with particular reference to non-diffusion of crop plants, and the significance of site function and sample location. In: Behre, K., Willcox, G. (eds) Proceedings of the 11th IWGP Symposium, Toulouse, 1998. Vegetation History and Archaeobotany, 8, 79–86CrossRefGoogle Scholar
  8. Burkart, M. (1998). Die Grünlandvegetation der unteren Havelaue in synökologischer und syntaxonomischer Sicht. Archiv naturwissenschaftlicher Dissertationen, 7, 1–260Google Scholar
  9. Chester, P.I., Raine, J.I. (2001). Pollen and spore keys for Quaternary deposits in the northern Pindos Mountains, Greece. Grana, 40, 299–387CrossRefGoogle Scholar
  10. Chrtek, J. (1979a). Klič k určení zástupcú rodu Urtica L. v ČSR (Key to the identification of Urtica L. species in the ČSR). Zprávy Československé botanické společnosti, 14, 1–7Google Scholar
  11. Chrtek, J. (1979b). Bemerkungen zur Gliederung der Gattung Urtica L. Folia Geobotanica et Phytotaxonomica, 14, 265–266Google Scholar
  12. Chrtek, J. (2002). Urticaceae. In: Kubat, K., Hrouda, L., Chrtek, J., Kaplan, Z., Kirschner, J., Štepánek,, J. (eds). Klič ke květeně České republiky. Academia, Praha, pp 139–140Google Scholar
  13. Emrović, B., Glavač, V., Pranjić, A. (1964). Über die Stammform der spitzblättrigen Esche (Fraxinus angustifolius Vahl) in verschiedenen Auenwaldgesellschaften des Savagebietes in Kroatien (Jugoslawien). Schweizerische Zeitschrift für Forstwesen, 3, 143–162 Google Scholar
  14. Fægri, K., Iversen, J. (1989). Textbook of pollen analysis, 4th edn. Wiley, ChichesterGoogle Scholar
  15. Faurholdt, N., Schou, J.C. (1994). Sump-Nælde (Urtica kioviensis Rogow.) – ny art for Norden. Urt, 1994/3, 67–73Google Scholar
  16. Feinbrun-Dothan, N., Danin, A. (1991). Analytical flora of Eretz-Israel. CANA, Jerusalem Gel’tman, D.V. (1988). Genus Urtica L. (Urticaceae) in URSS. Novitates Systematicae Plantarum Vascularium, 25, 68–80 Google Scholar
  17. Gel’tman, D.V. (1988). Genus Urtica L. (Urticaceae) in URSS. Novitates Systematicae Plantarum Vascularium, 25, 68–80Google Scholar
  18. Gel’tman, D.V. (1993). Urtica L. In: Tutin, T.G. et al. Flora Europaea, vol 1, 2nd ed. University Press, Cambridge, pp 79–80Google Scholar
  19. Görsdorf, J., Gramsch, B. (2004). Interpretations of 14C-datings of the Mesolithic site Friesack, Germany. In: Higham, T.F.G., Bronk Ramsey, C., Owen, D.C. (eds.). Radiocarbon and Archaeology: Proceedings of the 4th Symposium Oxford 2002. Oxbow, Oxford, pp 303–311Google Scholar
  20. Gramsch, B. (1979). Neue Ausgrabungen auf dem mesolithisch-neolithischen Fundplatz Friesack, Kr. Nauen. Ausgrabungen und Funde, 24, 56–61Google Scholar
  21. Gramsch, B. (1981). Der mesolithisch-neolithische Moorfundplatz bei Friesack, Kr. Nauen. 2. Vorbericht. Ausgrabungen und Funde, 26, 65–72Google Scholar
  22. Gramsch, B. (2000). Friesack: Letzte Jäger und Sammler in Brandenburg. Jahrbuch des Römisch-Germanischen Zentralmuseums Mainz, 47, 51–96Google Scholar
  23. Gutte, P., Jage, H., Jage, J. (1973). Urtica kioviensis Rogow im Elbe-Havel-Winkel (Bezirk Magdeburg, DDR). Gleditschia, 1, 95–97Google Scholar
  24. Hilbig, W., Reichhoff, L. (1974). Zur Vegetation und Flora des Naturschutzgebietes „Schollener See“, Kreis Havelberg. Hercynia N.F., 11, 215–232Google Scholar
  25. Jäger, E., Schubert, R., Werner, K. (1988). Werner Rothmaler Exkursionsflora, vol. 3, 7th edn. Atlas der Gefäßpflanzen. Volk und Wissen, BerlinGoogle Scholar
  26. Kernchen, I., Gramsch, B. (1989). Mesolithische Netz- und Seilreste von Friesack, Bezirk Potsdam, und ihre Konservierung. Veröffentlichungen des Museums für Ur- und Frühgeschichte Potsdam, 23, 23–27Google Scholar
  27. Kloss, K. (1987a). Pollenanalysen zur Vegetationsgeschichte, Moorentwicklung und mesolithisch-neolithischen Besiedlung im Unteren Rhinluch bei Friesack, Bezirk Potsdam. Veröffentlichungen des Museums für Ur- und Frühgeschichte Potsdam, 21, 101–120Google Scholar
  28. Kloss, K. (1987b). Zur Umwelt mesolithischer Jäger und Sammler im Unteren Rhinluch bei Friesack. Veröffentlichungen des Museums für Ur- und Frühgeschichte Potsdam, 21, 121–130Google Scholar
  29. Knörzer, K.-H., Meurers-Balke, J. (1999). Die frühholozäne Flora des Rheintales bei Neuss und der Erftaue bei Hombroich. Decheniana, Beiheft, 38, 1–181Google Scholar
  30. Konczak, P., Sukopp, H., Weinert, E. (1968). Zur Verbreitung und Vergesellschaftung von Urtica kioviensis Rogowitsch in Brandenburg. Verhandlungen des Botanischen Vereins der Provinz Brandenburg, 105, 108–116Google Scholar
  31. Körber-Grohne, U. (1995). Bericht über die botanisch-mikroskopische Bestimmung des Rohmaterials von einigen Schnüren, Seilen und Netzen von Friesack, Landkreis Havelland. Veröffentlichungen des Brandenburgischen Landesmuseums für Ur- und Frühgeschichte, 29, 7–12Google Scholar
  32. Kreuz, A. Schäfer, E. (2002). A new archaeobotanical database program. In: Jacomet, S., Jones, G., Charles, M., Bittmann, F. (eds) Archaeology of Plants. Current Research in Archaeobotany. Vegetation History and Archaeobotany, 11, 177–179CrossRefGoogle Scholar
  33. Kubiak-Martens, L. (2002). New evidence for the use of root foods in pre-agrarian subsistence recovered from the Late Mesolithic site at Halsskov in Denmark. In: Jacomet, S., Jones, G., Charles, M., Bittmann, F. (eds) Archaeology of Plants. Current Research in Archaeobotany. Vegetation History and Archaeobotany, 11, 23–31CrossRefGoogle Scholar
  34. Kučan, D. (1991). Eine neue Methode zur Fixierung fossiler unverkohlter Pflanzenreste, dargestellt an Beispielen laufender Untersuchungen aus Samos und Oberaden. In: Renfrew, J.M. (ed) New light on early farming. Recent developments in Palaeoethnobotany. Edinburgh University Press, pp 15–38Google Scholar
  35. Müller-Stoll, W.R., Fischer, W., Krausch, H.-D. (1962). Verbreitungskarten brandenburgischer Leitpflanzen. Vierte Reihe. Wissenschaftliche Zeitschrift der PH Potsdam, Mathematisch-Naturwissnenschaftliche Reihe, 7, 95–150Google Scholar
  36. Mundel, G., Trettin, R., Hiller, R. (1983). Zur Moorentwicklung und Landschaftsgeschichte des Havelländischen Luches. Archiv für Naturschutz und Landschaftsforschung, 23, 251–264Google Scholar
  37. Oberdorfer, E. (2001): Pflanzensoziologische Exkursionsflora, 8th edn. Ulmer, StuttgartGoogle Scholar
  38. Paiva, J. (1993). Urtica L. In: Castroviejo et al. (eds) Flora Iberica, vol 3. CSIC, Madrid, pp 263–268Google Scholar
  39. Punt, W., Malotaux, M. (1984). The Northwest European Pollen Flora, 31. Cannabaceae, Moraceae and Urticaceae. Review of Palaeobotany and Palynology, 42, 23–44CrossRefGoogle Scholar
  40. Reimer, P.J., Baillie, M.G.L., Bard, E., Bayliss, A., Beck, J.W., Bertrand, C., Blackwell, P.G., Buck, C.E., Burr, G., Cutler, K.B., Damon, P.E., Edwards, R.L., Fairbanks, R.G., Friedrich, M.,Guilderson, T.P., Hughen, K.A., Kromer, B., McCormac, F.G., Manning, S., Bronk Ramsey, C., Reimer, R.W., Remmele, S., Southon, J.R., Stuiver, M., Talamo, S., Taylor, F.W., Plicht, J. van der, Weyhenmeyer, C.E. (2004). IntCal04 Terrestrial radiocarbon age calibration, 26 - 0 ka BP. Radiocarbon 46, 1029–1058Google Scholar
  41. Rogowitsch, A. (1843). Urtica kioviensis. Species nova plantarum. Bulletin de la Société Impériale des Naturalistes de Moscou, 16, 324–326Google Scholar
  42. Schneider, M. (1932). Die Urkeramiker. LeipzigGoogle Scholar
  43. Schoknecht, T. (1988): Ein Vorkommen von Urtica kioviensis ROGOW. in Mittelmecklenburg.- Botanischer Rundbrief für den Bezirk Neubrandenburg 20, 49–51Google Scholar
  44. Scholz, H., Sukopp, H. (1960). Zweites Verzeichnis von Neufunden höherer Pflanzen aus der Mark Brandenburg und angrenzenden Gebieten. Verhandlungen des Botanischen Vereins der Provinz Brandenburg, 98–100, 123–149Google Scholar
  45. Sorsa, P., Huttunen, P. (1975). On the pollen morphology of the Urticaceae. Annales Botanici Fennici, 12, 165–182Google Scholar
  46. Stoller, J. (1927). Moorgeologische Untersuchung im Havelländischen Luche nordwestlich von Friesack zur Feststellung des Alters einer mesolithischen Kulturschicht an der III. Rhinbrücke. Jahrbuch der Preußischen Geologischen Landesanstalt, 48, 748–764Google Scholar
  47. Stuiver, M., Reimer, P.J. (1993). Extended 14C data base, a revised Calib 3.0 14C age calibration program. Radiocarbon, 35, 215–230Google Scholar
  48. Tarnavaschi, I.T., Serbănescus-Jitariu, G., Mitroiu, N., Rădulescu, D. (1967). Zur Pollenmorphologie der Urticales aus der Flora Rumäniens. Revue roumaine de biologie - Série de botanique, 12, 251–262Google Scholar
  49. Thanikaimoni, G. (1986). Pollen apertures: form and function. In: Blackmore, S., Ferguson, I.K. (eds) Pollen and spores. Form and function. Academic Press, London, pp 119–136Google Scholar
  50. Wollert, H., Bolbrinker, P., Welk, E. (2003). Zum Vorkommen und soziologischen Verhalten von Urtica kioviensis Rogowitsch in der Mecklenburgischen Schweiz (Ostmecklenburg) sowie zur gegenwärtigen Verbreitung der Art. Botanischer Rundbrief für Mecklenburg-Vorpommern, 38, 9–20Google Scholar
  51. Wollert, H., Bolbrinker, P., Funk, B. (2005): Zu einem weiteren Vorkommen von Urtica kioviensis im Streitholz westlich Teterow (Ostmecklenburg). Botanischer Rundbrief für Mecklenburg-Vorpommern 40 (in prep.)Google Scholar
  52. Zólyomi, B. (1936). Urtica kioviensis Rogowitsch neu für die deutsche Flora. Verhandlungen des Botanischen Vereins der Provinz Brandenburg, 76, 152–156Google Scholar

Copyright information

© Springer-Verlag 2005

Authors and Affiliations

  • Steffen Wolters
    • 1
  • Felix Bittmann
    • 1
  • Volker Kummer
    • 2
  1. 1.Niedersächsisches Institut für historische KüstenforschungWilhelmshavenGermany
  2. 2.Institut für Biochemie und BiologieUniversität PotsdamPotsdamGermany

Personalised recommendations