Pollen and non-pollen palynomorphs as tools for identifying alder carr deposits: A surface sample study from NE-Germany

Review of Palaeobotany and Palynology 186 (2012) 38–57

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Research paper

Pollen and non-pollen palynomorphs as tools for identifying alder carr deposits: A surface sample study from NE-Germany Anja Prager a,⁎, Martin Theuerkauf a, John Couwenberg a, Alexandra Barthelmes a, André Aptroot b, Hans Joosten a a b

Institute of Botany and Landscape Ecology, Ernst-Moritz-Arndt-University, Grimmer Straße 88, D-17487 Greifswald, Germany Adviesbureau voor Bryologie en Lichenologie - Herbarium (ABL), Gerrit van der Veenstraat 107, 3762 XK Soest, The Netherlands

a r t i c l e

i n f o

Article history: Received 15 January 2012 Received in revised form 30 June 2012 Accepted 9 July 2012 Keywords: alder carr surface samples non-pollen palynomorphs (NPPs) fungal and wood remains wood peat displacement peat

a b s t r a c t Alder wood peat is regularly found in central Europe, but palaeoecological studies of alder wood peat are rare, mainly because of lack of identifiable pollen and macrofossils. One central question is whether alder wood peat accumulates under alder carr vegetation (primary) or as displacement peat with alder roots growing into existing peat formed under open fen vegetation (secondary). Standard palaeoecological methods have proven unsuited to solve this question. This paper presents a surface sample study testing whether non-pollen palynomorphs (NPPs) can be used for identifying various types of fen vegetation, especially alder carrs, and for reconstructing the formation of wood peat. Different substrate types (litter, moss, dead wood, water and exposed peat surface) were sampled in 10 alder carrs, one birch carr and three sites with open fen vegetation in NE-Germany and analysed for pollen and NPPs. Vegetation relevees were recorded and water and soil conditions were measured. We overall recorded 412 NPP types, including 36 known EMA types (recorded in alder carr) and 51 HDV types (recorded in other ecosystems). For present publication we selected the 96 most abundant and characteristic NPPs (including 14 HDV types and 36 known EMA types). 46 NPP types are newly described and illustrated. Our study shows that, opposed to pollen, NPP types clearly reflect the different fen vegetation types making NPPs a valuable tool in palaeoecological studies of fen peat. © 2012 Elsevier B.V. All rights reserved.

1. Introduction In recent years, alder forestry has gained increased attention as an alternative land use practise on drained and abandoned fen peatlands in central Europe (Schäfer and Joosten, 2005). An important question in the implementation is whether alder carrs are able to grow under conditions wet enough to halt peat degradation or even to restore net accumulation. Despite its widespread occurrence, the formation of alder wood peat has rarely been studied palaeoecologically, because of the poor preservation of pollen and macrofossils (Barthelmes et al., 2006, 2012–this issue). Grosse-Brauckmann (2006) distinguishes two possible paths of alder wood peat formation. Primary alder wood peat (‘true’ alder carr peat) results from accumulation of above and below ground biomass in an alder carr. Alder wood is here embedded in a matrix of contemporaneous above-ground remains of Alnus. Secondary alder wood peat results from displacement of existing peat – formed in an open fen – by below ground alder wood (roots). Such displacement peat (cf. Berglund, 1986) accumulates during periods of low water levels when formation of peat at the surface is inhibited or ⁎ Corresponding author. Tel.: +49 3834 864691; fax: +49 3834 864114. E-mail address: [email protected] (A. Prager). 0034-6667/$ – see front matter © 2012 Elsevier B.V. All rights reserved. doi:10.1016/j.revpalbo.2012.07.006

even a net mineralization of surface peat takes place. Alder roots may then penetrate existing underlying peat layers, transforming these into displacement peat, i.e. alder wood embedded in a matrix of older e.g. sedge peat. Above ground remains of Alnus are absent. Secondary alder peat may typically form in fens with a water supply fluctuating over decades to centuries. During wetter phases, open fen vegetation accumulates peat; during drier phases trees colonise the surface and the existing peat is transformed into displacement peat. Differentiating primary and secondary wood peat is strongly hampered by the poor preservation of pollen and macrofossils in alder wood peat (Barthelmes et al., 2006; Barthelmes, 2009; Barthelmes et al., 2012–this issue). Above ground remains such as fruits and leaves of alder and other typical plants from alder carrs that would allow differentiating primary from secondary peat formation are often not preserved. Also pollen is often poorly preserved due to high microbial (Havinga, 1967, 1984; Dilly et al., 1996) and macrofungal activity (Grauwinkel, 1987) in the presence of oxygen and nutrients. Alnus is a strong pollen producer (Janssen, 1959). As ALNUS pollen with its prominent arcs (Moore et al., 1991) is more easily recognised than most other pollen types when corroded, the type will often be overrepresented in palynological studies of highly decomposed fen peat (Janssen, 1959, 1973, 1984; Jackson and Lyford, 1999). Besides ALNUS pollen, only

A. Prager et al. / Review of Palaeobotany and Palynology 186 (2012) 38–57

few pollen types are exclusively produced by taxa from alder carrs, whereas sedges produce little pollen when growing in well established, undisturbed stands (Segerström and Emanuelsson, 2002; Wiebe, 1998). Overall pollen data are thus poorly suited for confirming the local presence of either alder carr or sedge fen vegetation (Waller et al., 2005; Barthelmes, 2009). Non-pollen-palynomorphs (NPPs, i.e. “Types” sensu van Geel, 1972) are often less susceptible to corrosion and might be valuable indicators in highly decomposed peat (van Geel, 2006). Little is, however, known about their dispersal and the ecological conditions they reflect. The few surface sample studies that address this constraint (Mulder and Janssen, 1999; Mulder and de Zwart, 2003; Mulder et al., 2003; Blackford and Innes, 2006; Graf and Chmura, 2006; Yeloff et al., 2007; Cugny et al., 2010; Ejarque et al., 2011; Gelorini et al., 2011) have not included alder carr ecosystems. Prager et al. (2006) presented an initial surface sample study proving the suitability of NPPs in reconstructing past vegetation and environment in alder carrs (Barthelmes et al., 2006, 2012–this issue; Barthelmes, 2009). This paper aims to identify further NPPs that allow to distinguish between primary alder wood peat (by indicating the presence of Alnus during initial peat formation) from secondary alder wood peat = displacement peat (by indicating the presence of open vegetation during initial peat formation) and thus enable better understanding of the natural dynamics of alder carr vegetation and the formation of alder wood peat. 2. Sites and methods This study includes surface samples from 10 alder carrs, one birch carr and three sites with open fen vegetation (see Table 1, Fig. 1) situated in NE-Germany. To cover a wide range of alder carr types, we selected alder carrs with different water regimes (artesian ground

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water, flooding, more or less fluctuating ground water, percolation), water level (dry–very wet) and herb vegetation (Table 1). Water tables were measured in a level tube every second week over a one year period (2003/2004). C/N values (11.3–17.4, Dumas-method), and pH (subneutral: 4.9–6.6) were measured in the upper 5 cm of the peat. Values from open fen sites are similar (C/N = 16.4; pH = 6.2), values from birch carr are different (C/N = 27; pH = 3.1) than values from the alder carrs. In contrast to pollen, it is mostly unknown where and how NPPs are formed and dispersed. At each site, we thus sampled a wide range of microhabitats, including dead wood (birch/alder carr), mosses from hollows (alder carr), mosses from hummocks (birch/alder carr), leaf litter (all), water (alder carr, sedge fen) and surface peat (open fen). Dead wood included branches and other wood collected off the ground, and bark of tree stumps. To standardize the collecting properties of the mosses (cf. Räsänen et al., 2004), we collected only Hypnum cupressiforme except for Sphagnum palustre in site 7. From each site and microhabitat three subsamples were collected randomly and pooled (total volume 250 cm³) in autumn 2003 over an area of 20× 20 m in the carr sites and 5 × 5 m in open fen sites. The vegetation of these sampling areas was recorded. From the resulting 55 bulk samples a volume of 10 cm³ (peat), 50 cm³ (litter, moss, dead wood) or 100 cm³ (water) was used for further sample preparation. Samples were prepared following standard procedures for pollen samples (Fægri and Iversen, 1989) including treatment with HCl and KOH (each 10 min), sieving (mesh width 120 μm, following Moore et al., 1991), treatment with HF, acetolysis (7 min), and mounting in silicone oil (2000 CS). Counting was carried out with 400× magnification (Zeiss Axioskop 40); larger magnifications (600×, 1000×) were used to identify problematic objects. Pollen grains were determined with and named after Moore et al. (1991) or with The Northwest European Pollen Flora

Table 1 Characterisation of the study sites. Classification follows Succow and Joosten (2001): wetness based on median water table: very wet = 20–0 cm above surface, wet = 0–20 cm below surface, moist = 20–45 cm below surface, dry = 45–80 cm. No. Site name

Position

1

Murchin

53° 54′ 58′′ N Alder carr (2.5 ha) within 13° 44′ 28″ E agricultural fields

Artesian groundwater, very wet

2

Lüssow

53° 53′ 28″ N 13° 28′ 41″ E

Alder carr (100 ha) in a large river valley mire

Percolation, flooding, very wet

3

‘Heidenholz’ near Hohenzieritz ‘Mirower Holm’

Alder carr (1.2 ha) in beech and spruce forest Alder carr (7.2 ha), bordered by beech forest and lake Alder carr (7.2 ha), bordered by beech forest and lake Alder carr (20 ha), lake (S), fen meadow/pine forest (N) Birch carr (0.8) in beech forest

Ground water, moist

4 5 6 7 8 9 10 11 12

53° 13° 53° 12° ‘Mirower Holm’ 53° 12° ‘Stolpsee’ near Himmelpfort 53° 13° Southern ‘Haussee’ near 53° Hardenbeck 13° Glewitz 54° 12° ‘Elisenhain’ near Greifswald 54° 13° Drechow 54° 12° Neukloster 53° 11° ‘Katenwiese’ near Greifswald 54° 13°

26′ 05′ 14′ 49′ 14′ 49′ 06′ 07′ 08′ 20′ 02′ 56′ 04′ 27′ 07′ 47′ 50′ 41′ 05′ 19′

05″ N 05″ E 29″ N 04″ E 27″ N 17″ E 49″ N 52″ E 13″ N 16″ E 03″ N 10″ E 34″ N 35″ E 25″ N 38″ E 05″ N 53″ E 38″ N 19″ E

13

‘Katenwiese’ near Greifswald 54° 05′ 38″ N 13° 19′ 21″ E

14

‘Katenwiese’ near Greifswald 54° 05′ 38″ N 13° 19′ 22″ E

Vegetation type and surroundings Water regime/level

Percolation, wet to very wet Percolation, very wet Percolation, wet Percolation, wet to very wet

Alder carr (4.5 ha) in grassland

Ground water, dry

Alder carr (0.05 ha) in mixed deciduous forest Alder carr (2.7 ha) within agricultural fields/meadows Alder carr (5.9 ha) within beech forest large fen 150 m south of mixed deciduous forest (some Alnus at forest margin) large fen 45 m south of mixed deciduous forest (some Alnus at forest margin) large fen 25 m south of mixed deciduous forest (some Alnus at forest margin)

Strongly fluctuating, wet

Ground vegetation Cardamine amara, Caltha palustris, Carex acutiformis, Solanum dulcamara, Urtica dioica Sium latifolium, Carex acutiformis, Berula erecta, Solanum dulcamara, Thelypteris palustris, Urtica dioica Anemone nemorosa, Rubus idaeus, Phragmites australis, Solanum dulcamara, Urtica dioica Carex acutiformis, Solanum dulcamara, Poa trivialis, Thelypteris palustris, Urtica dioica Thelypteris palustris, Carex elata, Solanum dulcamara, Thelypteris palustris, Urtica dioica Valeriana dioica, Valeriana dioica, Menyanthis trifoliata, Solanum dulcamara Molinia caerulea, Carex lasiocarpa, Sphagnum spec. Urtica dioica, Cirsium oleraceum, Geum urbanum

Phalaris arundinacea, Carex acutiformis, Solanum dulcamara, Urtica dioica Percolation, wet Phragmites australis, Geum rivale, Filipendula ulmaria, Urtica dioica, Solanum dulcamara Strongly fluctuating, very wet Iris pseudacorus, Lysimachia thyrsiflora, Galium palustre, Solanum dulcamara Ground water, wet, regular flooding Phalaris arundinacea, Carex disticha, Inula britannica Ground water, very wet, regular flooding

Carex riparia

Ground water, very wet, regular flooding

Phragmites australis

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Fig. 1. Location of the surface sample sites. (a) General map NE Germany with sites. (b–l) Single sites and surrounding areas: (b) 1-Murchin, (c) 2-Lüssow, (d) 3-‘Heidenholz’ near Hohenzieritz, (e) 4 + 5-‘Mirower Holm’, (f) 6-‘Stolpsee’ near Himmelpfort, (g) 7-Southern ‘Haussee’ near Hardenbeck, (h) 8-Glewitz, (i) 9-‘Eldena’ near Greifswald, (j) 10-Drechow, (k) 11-Neukloster, (l) 12 + 13 + 14-‘Katenwiese’ near Greifswald.

(=Punt, 1976; Punt and Clarke, 1980, 1981, 1984; Punt and Blackmore, 1991; Punt et al., 1988, 1995, 2003) indicated by the suffix ‘m’ or ‘p’, respectively. NPPs (‘type’ sensu van Geel, 1972) were identified as follows: HDV-x after van Geel (1972, 1978), van Geel et al. (1980/81, 1982/83, 1989, 2003, 2011), Bakker and van Smeerdijk (1982), Pals et al. (1980), and van der Wiel (1982); UG-x after Gelorini et al. (2011); EMA-x after Barthelmes (2009), Barthelmes et al. (2006, this

volume) and Prager et al. (2006). Newly discovered NPPs were photographed with a Zeiss Axiocam digital camera and consecutively numbered as EMA-types (=Ernst-Moritz-Arndt-University Greifswald). The description of new NPPs is strictly based on the characteristics observed in this study; information from taxonomic literature (e.g. on fungal spores) is not included. In order to differentiate clearly between taxa (written in italics) and palynomorphs (pollen as well as NPP

A. Prager et al. / Review of Palaeobotany and Palynology 186 (2012) 38–57

types), the latter are displayed in the text in SMALL CAPITALS (Joosten and de Klerk, 2002). Excepted from this rule are palynomorphs that represent complete organisms (e.g. algae, testate amoebe). Microfossil frequencies were calculated relative to an upland pollen sum including pollen types that are assumed to originate from upland trees, shrubs, and herbs (AP+NAP). To eliminate (extra) local effects (cf. Janssen, 1959; Janssen and IJzermans-Lutgerhorst, 1973), pollen types produced by (potential) fen taxa (e.g. SALIX, ALNUS, CYPERACEAE, WILD GRASS GROUP) were excluded from the sum. Due to low pollen concentration, namely in dead wood samples, pollen sums are low but mostly above 100 grains. All slides (maximally five per sample) were counted completely. Descriptions of frequency and abundance of NPPs follow Table 2. For illustration, the 55 bulk samples were summarised with respect to sample sites (Figs. 2 and 3) or substrate types (Fig. 4), respectively. Calculation and presentation of the data was made with the software C2 (Juggins, 2003). 3. Results and discussion 3.1. Pollen (Fig. 2) Pollen data from surface sample differ between the major site types. ALNUS reaches highest abundances in alder carr, values vary from 66 to 904% (Fig. 2). The birch carr is characterised by high values of BETULA and SPHAGNUM, ALNUS arrives at 36%. Betula produces a lot of airborn pollen, and even though the highest BETULA value is found in the birch carr, high values are also recorded in the open vegetation. Open fen sites are characterised by high values of WILD GRASS GROUP or CYPERACEAE, ALNUS ranges between 5 and 62%. Yet not in all cases the differentiation is sharp, site 11 (alder carr) and site 13 (open fen) only differ in the somewhat higher abundance of CYPERACEAE. High ALNUS values in open vegetation, as in site 13, have also been described in Waller et al. (2005) and Barthelmes (2009). Differentiating between open fen and alder carr may be further complicated if both vegetation types occur in a fine grained mosaic and not, as in present study, in extended separate stands. Several wetland pollen and spore types, e.g. URTICA DIOICA TYPE and SOLANUM DULCAMARA (group undiff. in Fig. 2) are similarly abundant at all sites, as are the plant species producing these pollen or spore types. SPHAGNUM is clearly restricted to the birch carr site and so is Sphagnum. For some pollen types, plant species that might have produced them were absent from the sampling area. The same applies to pollen types associated with open fen species (e.g. umbelliferae, serratula type respectively). In conclusion, alder carr, birch carr, and open vegetation are not fully distinguishable using modern pollen data. In fossil material the differentiation will be further hampered by poor pollen preservation. 3.2. NPPs (Figs. 3, 4) In the 55 samples from the 14 sites 412 NPP types were recorded, 87 previously described and 325 new. From that, 96 NPPs potentially helpful in distinguishing between vegetation types are presented below. This number includes 14 HDV types and 36 EMA types that

Table 2 Classes of frequency and abundance used in describing the distribution of NPPs. Frequency in % of samples Rare

Occasional

Common

Frequent

>0–25

26–50

51–75

76–100

Abundance in single samples in % of the upland pollen sum Scarce

Recurrent

Numerous

Abundant

0.1–5

6–25

26–75

>75

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have already been found in peat cores (Barthelmes et al., 2006; Barthelmes, 2009; Barthelmes et al., 2012–this issue). We describe and illustrate 46 new EMAs in this paper. The following indicator groups with respect to site (Fig. 3) and substrate types (Fig. 4) may be distinguished. 3.2.1. Indicator group: carr undiff. (N = 13) Thirteen types are associated both with alder and with birch carr and are thus regarded as general indicators of forested fens (carrs). Within this group, some NPPs are related to dead wood and tree leaf litter, including EMA-11, EMA-35, EMA-37, EMA-112, EMA-113, EMA-15 + HDV-114, HDV-36 and HDV-114. EMA-114 is restricted to litter (sedge and birch). HDV-32A is restricted to hummock mosses, i.e. to a microhabitat typical of forested mire. EMA-12, EMA-43 und EMA-49 are not clearly restricted to a single carr substrate type. The origin of most types is unknown. All of them seem to be restricted to the specific conditions prevailing in a carr, including the presence of wood, shadow, no extreme summer temperatures and high humidity. Some types present in displacement peat (cf. Grosse-Brauckmann, 2006) may also form beneath ground level within the peat. 3.2.2. Indicator group: alder carr (N = 31) Among the 31 NPP types associated with alder carr (Fig. 3), EMA-27A, EMA-27B, EMA-49, EMA-52, EMA-124 and EMA-134 concentrate in samples of alder leaf litter (Fig. 4), i.e. these types are clearly related to above ground remnants of alder. In a fossil peat sample these types thus indicate that alder was present during initial peat formation, indicating that the peat was formed as primary alder wood peat. EMA-32 is additionally present in sedge litter, but may still be a reliable indicator for the presence of alder. EMA-3, EMA-4, EMA-7, EMA-8, EMA-9, EMA-24, EMA-25, EMA-26, EMA-30, EMA-31A, EMA-31B, EMA-36, EMA-38, EMA-48, EMA-59, EMA-126A + B, EMA-128, EMA-129 concentrate in dead wood samples. So far we do not know whether these types only result from above ground decay of dead wood; they may also be produced during the decay of alder roots. As such, they are potentially present primary as well as in secondary (displacement) alder wood peat. Below ground formation may primarily apply to NPP types that are products of wood decay (e.g. EMA-6). Fungal spores (e.g. EMA-24) instead may rarely be formed below ground because sporulation of most fungi requires oxygen and is thus hampered in water saturated peat (Carlile et al., 2001). If several NPP types of this group are recorded simultaneously, this may be taken as an indicator for the presence of alder during initial peat formation. EMA-6, EMA-40 and EMA-42 are clearly related to alder carr sites, but occur here in dead wood of alder and (sometimes more abundantly) of birch. Such pattern is characteristic for fungi that depend on specific site conditions (eutrophic/subneutral in alder carr versus mesotrophic/acidic in birch carr) but may use wood from different species (Schwarze et al., 2000). Alternatively, these types may be produced by fungi that generate different decay products (and thus different NPPs) under different conditions (Schwarze et al., 2000). In both cases the formation of EMA-6, EMA-40 and EMA-42 is obviously related to alder carr conditions so that in the fossil record these NPP types indicate the presence of alder carr during primary peat formation. EMA-127 is associated with hummock mosses in alder carrs, indicating the presence of alder carr vegetation during (primary) peat formation. It furthermore points to the presence of drier microhabitats. EMA-123 and EMA-125 are associated with hummock mosses and dead wood. EMA-51 and EMA-130 have no clear substrate preference. EMA-6, EMA-7, EMA-8, EMA-9 are remains of wood that show signs of fungal decay (‘hollows’ in lignin-layers). They show that alder wood is or has been present in a peat layer (which is not always detectable macroscopically). So far it is unknown whether they only form above ground (and would thus indicate primary wood peat

42 A. Prager et al. / Review of Palaeobotany and Palynology 186 (2012) 38–57 Fig. 2. Pollen diagram (following sites). The abundance of pollen types over the sites (manually arranged). Abundances are mean values per site as percentages of an upland pollen sum (ref. Sites and methods); pollen sum presented as absolute numbers of grains.

A. Prager et al. / Review of Palaeobotany and Palynology 186 (2012) 38–57 Fig. 3. NPP diagram (following sites). The abundance of NPP types over the sites (manually arranged). Abundances are mean values per site expressed on a logarithmic percentage scale (log with base 1.072) of an upland pollen sum (ref. Sites and methods); pollen sum presented as absolute numbers of grains; f. = fen, congl. = conglomerated.

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44 A. Prager et al. / Review of Palaeobotany and Palynology 186 (2012) 38–57 Fig. 4. NPP diagram (following substrate types). The abundance of NPP types over the substrate types (manually arranged). Abundances are mean values per substrate type and expressed on a logarithmic percentage scale (log with base 1.072) of an upland pollen sum (ref. Sites and methods); pollen sum presented as absolute numbers of grains; congl. = conglomerated, in substrates: L = litter, h = herbal, b = birch, D = dead wood, w = water, P = peat, M = moss, HM = hummock moss.

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formation) or also below ground (and would then also be present in secondary alder wood peat). At the moment they are thus not suited to distinguish primary and secondary carr peat formation. 3.2.3. Indicator group: birch carr (N = 6) This group includes NPP types that are restricted to or clearly concentrated in birch carr samples. The group is tentative, as only one birch carr is included in this study. EMA-119 and EMA-121 are only registered in above ground remains of birch (leaf litter and mosses) and thus possibly a clear indicator for on site presence of birch during peat formation. EMA-117, EMA-118, EMA-120, EMA-122 also occur in dead wood and may thus be found in displacement peat. If hollows in its lignin layer are recognisable EMA-117 furthermore indicates high decomposition. 3.2.4. Indicator group: open fen (N = 29) Some NPPs types from this group are abundant in open reed and open sedge vegetation: HDV-19, HDV-28, HDV-172, HDV-715, and EMA-86, EMA-90, EMA-91, EMA-92. They possibly reflect the specific conditions (high light availability) of open fen vegetation rather than single plant taxa (e.g. sedges or reed) in the vegetation. HDV-352 and EMA-55, EMA-97, EMA-98, EMA-99, EMA-105, EMA-106, EMA-107, EMA-108, EMA-109, EMA-110, EMA-111 are restricted to or concentrated in (open) sedge fens. Sedges may also be abundant in alder carrs. Yet in contrast to e.g. EMA-39, which is related to grasses and sedges in open as well as in forested habitats, the above NPP types appear to indicate open sedge fen vegetation. Also HDV-368, so far known as a dung indicator (van Geel et al., 2003), was found abundant on sedge litter. Similarly, HDV-58 and HDV-128B, known as indicators for open water in bog pools (van Geel, 1978), are in present data set restricted to sedge fen sites. EMA-92, EMA-93, EMA-94, and EMA-95 are clearly related to Phragmites litter and may thus be regarded as indicators of Phragmites australis. In present study also HDV-172, so far recorded on various other deposits (van Geel et al., 1982/83), occurs in Phragmites litter and surface peat layers. EMA-96, HDV-123 and HDV-351 only occur in surface peat samples from open vegetation. It is unknown where they originate. Overall, NPP types from this open fen group are indicators for open, non-forested vegetation. If present in alder wood peat, they indicate that the peat was initially formed under open vegetation and is thus a displacement peat. 3.2.5. Other indicators (N = 4) EMA-39 is related to sedge and grass litter in open and forested vegetation; it indicates the presence of grass like plants. EMA-1, EMA-45, and EMA-68 are fragments of vessels indicating the presence of cormophytes, which allows distinctions to be made between vegetation types dominated by mosses from those dominated by herbs.

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4. Interpreting NPPs In contrast to pollen, the origin of most NPP types is unclear. Hardly any information is available about where and in which quantities they are produced, how well they are dispersed and where they accumulate. To better understand the origin of NPPs, we assume that a substrate in which a type is five times more abundant than the mean is the host/parent substrate of that particular type. High abundances of NPP types are not necessarily related to the host material, however: EMA-6, a wood remain, is most abundant in the moss samples. We thus additionally used morphologic and, where available, taxonomic information to filter out substrates where a type is possibly produces and where it accumulates and is preserved. Within an alder carr, the majority of fungi grows on dead wood and leaf litter (Table 3). Spores possibly produced in or on dead wood (e.g. EMA-11, EMA-28) are not only abundant in dead wood samples but also in moss and water samples, which indicates that these types are well dispersed within an alder carr and also accumulate in mosses and pools. Similarly, NPPs abundant in leaf litter (e.g. EMA-109, EMA-130) and moss or water samples were possibly produced in leaf litter. The evenness of distribution provides some indication on the dispersal of single NPP types. Types that are abundant in one sample but (nearly) absent in other samples at the same site are obviously poorly dispersed. Single exceptionally high values also indicate production of a NPP on the sampled substrate. Sampling different substrates, implies that samples represent different time periods: while mosses and water from pools contain NPPs accumulated over a rather short period from few weeks to month, dead wood may persist over several years and can contain NPP types accumulated over such longer period. Samples from mosses and pools will thus sharper represent vegetation and site conditions at the time of sampling. Still little is known about the susceptibility of NPPs to decomposition in peat. Similar to pollen, some fungal spores may be more susceptible than others. Continued decay may transform wood remains into other types (Schwarze et al., 2000). Therefore threshold values above which producing taxa are deemed to have been locally present in a (sub)fossil context should be applied with utmost caution (cf. Davis and Shafer, 2006). Similar to pollen, NPPs may easily be transported over greater distance by wind. Also flooding may change NPP composition at a site. In present data set, only site 2 is regularly flooded, which has no marked effect on the overall NPP composition of this site, however (Fig. 3). For all these reasons, the derived indication of NPP types should be used cautiously and interpretations should always be based on a set of indicators. The (mean) percentage values presented here may not be compared with fossil samples unequivocally. Alder wood may mainly include (remnants of) dead wood so that NPPs related to

Table 3 Potential NPP catching, preserving and producing abilities of different substrate types (− = no, + to +++ = less to very high). Substrate type

Catchment and preservation of NPPs

Parent material of NPPs

Resulting abundance of NPPs

Main organism groups of NPPs

Moss cushions

+++

+

+++

Water

++

−

+++

Dead wood

+

+++

+++

Litter

++

+

+++

Soil

++

−

++

Fungi (Döbbeler, 1978), testate amoebae (Hoogenraad and de Groot, 1940; Grospietsch, 1972; Page, 1976), invertebrates Algae (Stevenson et al., 1996), some submerse decaying fungi (Tsui and Hyde, 2003; Czeczuga and Orlowska, 1995; Sati and Belwal, 2005), testate amoebae (ref. mosses), invertebrates Many microfungi (Ellis, 1971, 1976; Ellis and Ellis, 1997) and macrofungi (Grauwinkel, 1987; Breitenbach and Kränzlin, 1984–2000; Schwarze et al., 2000), invertebrates Many fungi: on land plants in general (Ellis and Ellis, 1997, on Common reed (Apinis et al., 1972, 1975; van Ryckegem et al., 2006), invertebrates Many fungi (Barron, 1968; Domsch and Gams, 1970; Domsch et al., 1993), invertebrates, testate amoebae (Page, 1976)

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dead wood may be more abundant than in the summarised values presented here. Furthermore, the abundance of NPPs may be altered during peat formation. Depending on the number in which they are produced, how well they are dispersed and preserved, some may be sharp indicators already in small abundances, others only in high abundances. Finally, our findings may only apply to a geographically limited area. Future surface sample studies need to cover a wider range of ecosystem types (most notably willow stands as the third type of forested fen), and to include various substrates. Sampling should include dead roots to test whether NPPs related to dead wood are produced only above (and then would indicate primary wood peat formation) or also below ground. 5. Conclusions Of the 96 selected NPP types 83 types were identified as indicators for either alder carr (31 types), birch carr (6), alder + birch carr (13), open vegetation (29), or the presence of sedges or grasses and cormophytes (4 types). Clear indicators for above ground remains of alder (NPP types associated with alder litter) and for open vegetation allow to distinguish wood peat formed as true alder carr peat in an alder carr (primary wood peat) from displacement peat (secondary wood peat) formed initially under open vegetation. NPPs have proven a valuable additional source of information in the reconstruction of fen vegetation and site conditions. In highly decomposed peat, where most other proxies are highly decomposed, NPPs may be an important additional source of palaeoecological information. 5.1. New information in earlier described NPPs HDV-19 van Geel 1978: ascospore 1-septate, known from Sphagnum papillosum/imbricatum bog peat. In present study scarcely found in herbal litter and superficial peat in very wet, open fens. Might indicate very wet conditions with open vegetation. HDV-28 van Geel 1978: spermatophore of Copepoda, indicator of (temporary) open water, only known from bog peat. In present study also scarcely recorded in herbal litter and superficial peat in open fen vegetation. Indicates open vegetation with at least temporarily open water. HDV-32A van Geel 1978: Assulina muscorum, theca, observed in bog peat (van Geel, 1978) and in alder carr (Prager et al., 2006). A. muscorum typical in drier mosses (Grospietsch, 1972). In present study recorded at all sites. Frequent and abundant (> 1%) in birch carr and alder carr. Highest values (recurrent to abundant) in hummock moss samples. Indicates drier microhabitats (hummocks) with moss cushions (in alder carr Hypnum cupressiforme, in Birch carr Sphagnum palustre).

HDV-36 van Geel 1978: remains of Acari (Orobatei). In present study common in alder and birch carr, scarce to recurrent (b 10%). Rare in open vegetation. Frequent in moss and dead wood samples. Acari occur in bogs, fens and meadows; valuable indicators only if determined to species level (Schelvis and van Geel, 1989). Thus here no indication. HDV-44 van Geel 1978: ascospores of Ustulina deusta, known from birch carr deposits (van Geel, 1978), associated with Fraxinus and Tilia (van Geel and Andersen, 1988). Ustulina deusta (= Kretzschmaria deusta, Rogers and Ju, 1998) is known from stumps and dead roots of e.g. Fagus, Aesculus, Betula, Taxus and Ulmus (Ellis and Ellis, 1997). Barthelmes et al. (2006) related HDV-44 to alder carr deposits, whereas Prager et al. (2006) could not proof relation to dead wood of alder. In present study occasionally found at all sites, scarce. Most abundant (~ 5%) in sedge litter (sedge fen) and hollow mosses (alder carr). Absent in water, peat and, remarkably, in dead wood. Here no indication. HDV-55A1 van Geel et al. 1980/81, ascospore of Sordaria spec., possibly indicating dung (van Geel et al., 2003). In present study occasional at all sites, scarce. Highest values in sedge fen (~ 5%). Here no indication. HDV-58 van Geel 1978: zygospore of Zygnemataceae. Known from bogs, indicating stagnant, shallow and mesotrophic fresh water (van Geel, 1978). In present study only in wet sedge fen (in sedge litter and open water), scarce. Known indication extended to open fen vegetation. HDV-123 Pals et al. 1980: unknown fungal spore, indicating mesotrophic conditions (van Geel et al., 1980/81), varying moisture in meso-eutrophic conditions (Bakker and Smeerdijk, 1982), possible relation to local Betula growth (Pals et al., 1980). In present study only once recorded in superficial peat sample in common reed under very wet conditions; numerous. Here no indication. HDV-128B Van Geel et al. 1982/83: possibly remnant of algae or other fresh water organism. In present study only in wet sedge fen (in sedge litter and water), scarce. Extending known indication to open, at least wet fen vegetation with temporarily open water. HDV-172 van Geel et al. 1982/83: ascospore of Coniochaeta cf. ligniaria, known from willow carr peat. Coniochaeta ligniaria (Grev.) Massee is known from dead wood (Ellis and Ellis, 1997) and dung (Munk, 1957). In present study occasional in open sites (herbal litter and superficial peat), scarce to highly abundant, highest value in Phragmites australis litter. Rare in alder carr. Indicates open fen vegetation, possibly with Phragmites australis. HDV-351 van Geel et al. 1980/81: fungal cells, indicator of human influence. In present study only in surface peat sample of Common reed fen, scarce. Here no indication. HDV-352 van Geel et al. 1980/81: Arcella sp., theca. In present study determined as Arcella discoides, observed in water plant societies (Grospietsch, 1972). HDV-352 common in open sites, extraordinary high values in wet sedge fen (sedge and Phalaris arundinacea litter). Rare in alder carrs, scarce. Indicates open sedge fen vegetation.

Plate I. (All scale bars are 10 μm): 5. EMA-5 conglomerate: unknown origin. 27B. EMA-27B: unknown fungal spores. 31B.a + b. EMA-31B: conidium fragment prob. of Thielaviopsis state of Ceratocystis moniliformes, a site view, b downward view. 48. EMA-48: unknown conidium. 52. EMA-52: poss. fungal remnant. 59. EMA-59: fungal spore or conidiophore-fragment. 63. EMA-63: unknown fungal spore. 67. EMA-67.a + b: invertebrate remain. 68. EMA-68.a + b: poss. fragment of spiralic tracheid, Cormophyta. 81. EMA-81: prob. chitinoid invertebrate remain. 82.a-c. EMA-82: fungal spore or algal cyst. 83A.,B. EMA-83A + B: poss. conidia of Monodictys paradoxa. 86.a + b. EMA-86: Bactrodesmium-like conidium. 87. EMA-87: elaters of capsulae from a Hepatic, prob. Lophocolea heterophylla. 90. EMA-90: conidium of Arthrinium cf. puccinioides. 91.a + b. EMA-91 ascospore of poss. Ceriophora palustris.

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HDV-368 van Geel et al. 1980/81 (=HDV-386 van Geel et al. 2003): ascospore of Podospora, coprophilous fungus. In present study rare, highest value in Carex acutiformis litter from sedge fen. Besides on dung HDV-368 might be produced on litter. Possibly indicates open vegetation. HDV-715 Bakker and van Smeerdijk 1982: triseptate fungal spore, known from peat layers that formed under eu- to mesotrophic helophyte marsh conditions. In present study only in very wet open sites, scarce to abundant. Highest value in reed fen. Indicates open vegetation under very wet conditions. EMA-1 Prager et al. 2006: tracheids of vascular bundles. Recorded in peat cores (Barthelmes et al., 2006; Barthelmes, 2009; Barthelmes et al., 2012–this issue). In present study frequent, scarce to abundant. Highest values in litter and water. Obviously collected and better preserved in water. Indicates presence of cormophytes, but no distinct vegetation type. EMA-3 Prager et al. 2006: conidium, probably of Melanconis alni. Fungus known from dead twigs and branches of Alder (Ellis and Ellis, 1997). Recorded in peat cores (Barthelmes et al., 2006; Barthelmes, 2009; Barthelmes et al., 2012–this issue). In present study in all samples from alder carr, numerous to abundant, highest values at wetter sites. Also common but only scarce outside alder carr, possibly representing regional deposition of the wind dispersed conidia. Indicates alder (also root wood?). EMA-4 Prager et al. 2006: fungal spore (immature conidium of Melanconis alni?). Recorded in peat cores (Barthelmes et al., 2006; Barthelmes, 2009; Barthelmes et al., 2012–this issue). In present study frequent in alder carr samples, scarce to abundant. Rare outside alder carr. Indicates alder (also roots?). EMA-5 Prager et al. 2006: unknown origin. Recorded in peat cores (Barthelmes et al., 2006; Barthelmes, 2009; Barthelmes et al., 2012–this issue). In present study occasional, usually scarce to recurrent. Abundant in one water sample, may be due to good preservation. Rarely occurring as EMA-5 conglomerate (Plate I, 5) in sedge fen. No clear indication. EMA-6 Prager et al. 2006: unknown origin, possibly corroded wood. Recorded in peat cores (Barthelmes et al., 2006; Barthelmes, 2009). In present study also found without circular to elliptic (corroded-?) areas on the surface (ref. Prager et al., 2006). In present study frequent in alder carrs. Mainly scarce to recurrent; numerous in some litter and hummock moss samples, occurs also in dead wood of birch. Indicates presence of trees (alder, birch?) in eutrophic carr conditions (also roots?), advise on high decomposition. EMA-7 Prager et al. 2006: possibly remnant of highly corroded wood. Recorded in peat cores (Barthelmes et al., 2006; Barthelmes, 2009; Barthelmes et al., 2012–this issue). In present study frequent. Abundant in alder carr samples, outside mostly scarce, possibly reflecting regional deposition. Highest values (100–10,000%) in dead wood (alder and birch), litter of alder and mosses of hummocks. Indicates in situ presence of alder. Indicates presence of trees (alder, birch?) in eutrophic carr conditions (also roots?), hints on high decomposition. EMA-8 Prager et al. 2006: possibly remnant of highly corroded wood. Recorded in peat cores (Barthelmes et al., 2006; Barthelmes, 2009; Barthelmes et al., 2012–this issue). In present study frequent in alder carr, scarce to abundant, >1000% in dead wood of alder. Outside alder carr rare, scarce, possibly reflecting regional deposition. Indicates presence of trees (alder, birch?) in eutrophic carr conditions (also roots?), advise on high decomposition. EMA-9 Prager et al. 2006: possibly highly corroded periderm of hard wood. Recorded in peat cores (Barthelmes et al., 2006; Barthelmes, 2009; Barthelmes et al., 2012–this issue). In recent study frequent in alder carr, abundant in dead wood of alder. Outside alder carr rare, scarce. Indicates presence of alder (also roots?) and high decomposition. EMA-11 Prager et al. 2006: aggregate rays in woody tissue. Recorded in peat cores (Barthelmes et al., 2006; Barthelmes, 2009; Barthelmes et al., 2012–this issue). In present study frequent in

alder carr, scarce to abundant. Highest values in dead wood of alder. Recurrent in dead wood in birch carr. In open vegetation rare, scarce. Indicates presence of alder or birch (also roots?), in high abundances also high decomposition. EMA-12 Prager et al. 2006: possibly thalloconidium of Sporoschisma or Chalara, 1-septate (Prager et al., 2006). In present study rarely also observed with 3 septa and 40 μm long. Recorded in peat cores (Barthelmes et al., 2006; Barthelmes, 2009). In present study only rarely found in alder and birch carr, scarce. Might indicate carr vegetation. EMA-15 Prager et al. 2006+ HDV-114 Pals et al. 1980: both fragments are parts of perforation plates in vessels of wood. If completely preserved, perforation plates can be determined to genus level (Pals et al., 1980; Schweingruber, 1990). Recorded in peat cores (Barthelmes, 2009; Barthelmes et al., 2006, 2012–this issue). In present study both HDV-types common in alder carr, mainly scarce to recurrent. Highest values in dead wood of alder. Recurrent also in dead wood in birch carr. Indicate presence of alder or birch (also roots?). EMA-18 Prager et al. 2006: probably conidium of Oncopodiella trigonella (Sacc.) Rifai, a hyphomycete on Hedera (cf. Ellis and Ellis, 1997), Ulmus and other trees (Ellis, 1971). Recorded in peat cores (Barthelmes, 2009; Barthelmes et al., 2006). In present study rare, scarce to recurrent. Not, as suggested in Prager et al. (2006), a clear indicator for alder. EMA-23 Prager et al. 2006: resembles seta of Phyllactinia spec. (det. A. Aptroot). Recorded in peat core (Barthelmes, 2009). In present study common, usually scarce to recurrent. Higher values in open vegetation, but pattern not distinct. Might indicate open vegetation. EMA-24 Prager et al. 2006: Hypoxylon-like ascospore. Recorded in peat cores (Barthelmes, 2009; Barthelmes et al., 2012–this issue). In present study some individuals larger (18–22 × 6–7 μm) than proposed in Prager et al. (2006). Might represent several species as size ranges of Sphaericaceae often overlap. In present study common in alder carr, mostly scarce, once extraordinary abundant (~ 1,900%) in dead wood of alder. Present also in birch carr, scarce. Might indicate presence of alder (also roots?). EMA-25 Prager et al. 2006: possibly ascospore of Cenangium gradonii Dennis. Recorded in peat cores (Barthelmes, 2009; Barthelmes et al., 2012–this issue). In present study besides 3-septate also 2-septate individuals. C. gradonii known from branches of alder (Ellis and Ellis, 1997). In present study common in alder carr, scarce to abundant, highest values in hummock mosses and dead wood of alder. Rare in open fen. Indicates presence of alder (also roots?). EMA-26 Prager et al. 2006: possibly conidium of dematiaceous hyphomycete, seemingly heavily damaged post mortem. In present study occasional in alder carr, scarce to abundant. Highest values in samples from hummock mosses and dead wood, EMA-26 is thus either produced on or effectively caught by mosses. Possibly indicator for alder carr vegetation. EMA-27A/B: In present study EMA-27 (Prager et al., 2006) has been found in two forms, one fits the previous description; the second is twice as large. Both are similarly distributed, the larger form however is rarer. EMA-27 has thus been divided in EMA-27A (the previously described smaller form) and EMA-27B (the larger form). EMA-27A (former EMA-27 Prager et al. 2006): unknown fungal spore. Recorded in peat cores (Barthelmes, 2009; Barthelmes et al., 2012–this issue). In present study occasional in alder carr, numerous to abundant in all alder litter samples. Clear indicator for above ground alder remains, thus for on-site presence of alder. EMA-27B (Plate I, 27B): unknown fungal spores, seemingly with irregular wall due to post mortem degradation. Single cells globose, 10–14 μm in Ø, wall about 1 μm thick, verrucate with wide but short warts, brown, forms often clusters of two or three cells. In present study rare in alder carr (mainly alder litter), scarce. Clear indicator for on-site alder presence. EMA-30 Prager et al. 2006: unknown fungal spore. In present study rare in alder carr, scarce to recurrent. Not, as suggested in

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Prager et al. (2006), restricted to the driest sites. Might indicate presence of alder. Quite similar to immatured conidia of Exosporium tiliae Link ex Schlechtend. as described and figured in Yurchenko (2001), but postulated distinct black scar missing in EMA-30, recent records only known from Tilia cordata (Yurchenko, 2001). EMA-31A/B: In present study a spore similar but smaller than EMA 31 (Prager et al., 2006) has been found. Both types could be attributed to two different fungi. EMA-31 was thus divided in EMA-31 A (=former EMA-31) and B (smaller form). Recorded in peat cores (EMA-31A and B counted as one in: Barthelmes, 2009; Barthelmes et al., 2012–this issue). EMA 31A: fragment of a conidium probably of Thielaviopsis basicola (Berk. and Br.) Ferraris (det. A. Aptroot), growing on rotten roots, e.g. of Daucus (Ellis and Ellis, 1997). In present study occasional in alder carr samples, mainly scarce, numerous in alder dead wood. Rare in dead wood in birch carr. Probably indicates presence of alder (and birch?), (also roots?). EMA-31B (Plate I, 31B.a + b): conidium fragment probably of Thielaviopsis state of Ceratocystis moniliformes (Hedgc.) C. Moreau (det. A. Aptroot). Hemispherical to bowl-shaped, ø 5–9 μm, 3–5 μm high, wall 1 μm thick, inward folded part (septum?) thinner, bright brown, psilate. C. moniliformes is known from wood of Fagus (Ellis and Ellis, 1997). In present study common in alder carr, scarce to recurrent, in alder dead wood once abundant. Concentrated in dead wood and litter of alder. Indicates presence of alder (also roots?). EMA-32 Prager et al. 2006: unknown fungal spore. Recorded in peat core (Barthelmes et al., 2012–this issue). In present study common in alder carr, missing only on drained site. Mostly scarce, recurrent to abundant in alder litter and once in sedge litter. EMA-32 obviously produced by a fungus on tree (sedge?) litter in alder carrs. Clear indicator for on site presence of alder. EMA-35 Prager et al. 2006: unknown origin, possibly invertebrate remain. Can be smaller than suggested in Prager et al. (2006), new size range: 15–63 × 10–24 μm. Also known from peat cores (Barthelmes, 2009; Barthelmes et al., 2012–this issue). In present study frequent in alder and birch carr, scarce to abundant. Highest values in dead wood and hummock mosses. On open sites occasional, scarce. Indicates presence of trees (alder, birch?) (also roots?). EMA-36 Prager et al. 2006: probably (as EMA-38) conidium of Excipularia fusispora (Berk. & Broome) Sacc. E. fusispora is known from decayed wood and bark of deciduous trees (e.g. Alnus glutinosa, Populus tremula and Salix cinerea; Yurchenko, 2001). Recorded in peat cores (Barthelmes, 2009; Barthelmes et al., 2012–this issue). In present study in all alder carr samples, scarce to abundant; highest values in dead wood and alder litter. Outside alder carr rare, scarce. Indicator for presence of alder (also roots?). EMA-36 and EMA-38 are possibly conidia of the same species. Whether the production of either form is related to specific site conditions, is unknown. EMA-37 Prager et al. 2006: possibly ascospore of Prosthecium auctum (Berk. & Broome) Petrak, observed on dead branches of trees (Ellis and Ellis, 1997) e.g. alder (Dennis, 1981). Recorded from peat cores (Barthelmes et al., 2012–this issue). In present study frequent in birch and alder carr, scarce to recurrent. Rare in open sites. Indicator for presence of birch or alder carr vegetation (also roots?). EMA-38 Prager et al. 2006: probably (as EMA-36) conidium of Excipularia fusispora (Berk. & Broome) Sacc. E. fusispora is known from decayed wood and bark of deciduous trees (e.g. Alnus glutinosa, Populus tremula and Salix cinerea; Yurchenko, 2001). Recorded in peat cores (Barthelmes, 2009; Barthelmes et al., 2012–this issue). In present study frequent in wet and very wet alder carr, outside rare. Mostly scarce to recurrent, one exceptionally high value in dead wood of alder. Indicates presence of alder carr (also roots?). EMA-39 Prager et al. 2006: possibly ascospore of Phaeosphaeria spec. (det. A. Aptroot), highly specialised parasitic fungi on grass-like plants (Shoemaker and Babcock, 1989). Recorded in peat cores (Barthelmes, 2009; Barthelmes et al., 2012–this issue). In present study

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in all vegetation types (except birch carr) occasional, scarce to numerous. Most abundant in sedge and reed litter. Possibly indicates presence of sedges and grasses in forested as well as in open vegetation. Identification to species level could allow sharper indication. EMA-40 Prager et al. 2006: possibly ascospore cf. Atopospora betulina (Fries) Petrak, observed on Betula (Dennis, 1981). In present study common in alder carr, mostly scarce to recurrent. Exceptionally high values in dead wood of alder and birch. Outside alder carr rare, scarce. Indicates presence of trees (alder, birch?) in eutrophic carr conditions (also roots?). EMA-42 Prager et al. 2006: unknown fungal spore. In present study size range extended to 14 × 8 μm. Also found in peat core. In present study common in alder carr, scarce to abundant, highest values in dead wood and hummock mosses. Indicates presence of trees (alder, birch?) in eutrophic carr conditions (also roots?). EMA-43 Prager et al. 2006: possibly spore of Lycogala (Myxomycota). In present study common in alder carr, scarce to abundant. Exceptionally high values in dead wood of alder. Occasional outside alder carr, scarce to recurrent. Indicates presence of trees (alder, birch?) in eutrophic carr conditions (also roots?). EMA-45 Prager et al. 2006: tracheid fragments of vascular bundles of Cormophyta. In present study occasional in alder carr and open vegetation, scarce to recurrent. Slightly more abundant in water and hollow mosses. Obviously collected and better preserved in water and hollow mosses. Indicates presence of cormophytes, but no distinct vegetation type. EMA-48 Prager et al. 2006: unknown conidium. Recorded in peat cores (Barthelmes, 2009). In present study size range extended to 27–29 × 9.5-11 μm, spores with pores in septa (Plate I, 48) probably mature individuals. Wrinkling sometimes restricted to the broader rounded end. In this study occasional in alder carr, scarce to abundant. Highest values in alder dead wood. Indicates presence of alder (also roots?). EMA-49 Prager et al. 2006: possibly: fruit-body appendage (seta) with swollen base. Recorded in peat core (Barthelmes, 2009). In present study occasional in alder carr; scarce to numerous. Highest values in alder litter. Scarce in mosses in birch carr. Clear indicator for above ground remains and thus presence of alder. EMA-51 Prager et al. 2006: unknown origin. Recorded in peat core (Barthelmes, 2009). In this study rare in alder carr. Scarce to recurrent. Might indicate alder carr. EMA-52 Prager et al. 2006: possibly no algal remain (cf. Prager et al., 2006) but rather remnant of litter decaying fungus. Recorded in peat core (Barthelmes et al., 2012–this issue). In present study maximum diameter given by Prager et al. (2006) extended to 60 μm. Now also individuals with net-like structure (Plate I, 52) observed. In present study common in alder carr, mostly scarce, but recurrent to abundant in alder litter. Outside alder carr rare, scarce. Clear indicator for above ground remains and thus presence of alder. EMA-55 Barthelmes et al. 2006: ascospore of Xylariaceae, possibly Helicogermslita, known from willow peat (Barthelmes et al., 2006). In present study only found in very wet sedge fen (peat and litter). Might thus be additionally an indicator for sedge fen.

6. Description, illustration, and distribution of the new non-pollen palynomorphs EMA-59 (Plate I, 59): fungal spore or conidiophore fragment. Cylindric, 3-septate, septa with pores, 22–24 × 5–6 μm, (rarely with 2 septa and only 15–17 μm long), both central cells 6–8 μm long, end cells 5–7 μm, lighter and torn off, often one end slightly conical, light to middle brown, psilate. Resembles thalloconidia from Sporoschismaspecies (det. A. Aptroot); but also the conidiophores (or their fragments) of Arthrinium-species (Ellis and Ellis, 1997). Recorded in peat cores (Barthelmes, 2009; Barthelmes et al., 2012–this issue). In present study

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common in alder carr samples, mostly scarce to recurrent. Might indicate presence of alder carr vegetation. EMA-63 (Plate I, 63.): unknown fungal spore. Ovoid, 5–12 × 2–7 μm, indistinct small round aperture (pore or hilum?) at tapered end, hyaline to very pale brown, psilate. In present study frequent in all vegetation types, scarce to abundant. Highest values in dead wood. Might, if occurring in high values, indicate presence of trees. However, EMA-63 is a simple spore type with few distinct features, which is produced by numerous fungal taxa; also, the size range reported here is extremely large, suggesting that elements of different origin are combined. Hence, the type should be interpreted with caution. EMA-67 (Plate I, 67.a + b): possibly invertebrate remain. Planar fragment with variable outline and size: 10–90 × 10–110 μm, pitted (3.3-5.5 μm in Ø) along lines (= perforated), broken along perforations, dark brown to black. In present study common at most sites, scarce to numerous. No obvious indication. EMA-68 (Plate I, 68.a + b): possibly fragment of spiralic tracheid (Cormophyta), known from roots of e.g. Cyperaceae (D. Michaelis, pers. com.). Spiralic and filiform, 5–67 μm long, Ø of spiral 5–9 μm, Ø of filament 1.0-1.4 μm, hyaline. Similar, but twice as large fragments described as HDV-217 in van Geel et al. (1989). Both remains (illustrated in a, and b) may well have belonged to different species; alternatively, photo 68b may actually be referable to EMA-87 (elaters of a Hepatic). Recorded in peat core (Barthelmes et al., 2012–this issue). In present surface samples common, scarce to recurrent. Indicates presence of cormophytes, but no distinct vegetation type. EMA-81 (Plate I, 81): probably chitinoid invertebrate remnant. Spiralic coiled tape, Ø about 10 μm, 30–40 μm long, distinctly parallel striate, striae in distance of 1 μm, one end tube-like, the other with protruding striae, hyaline to pale brown. In present surface samples occasional, scarce to recurrent. No obvious indication. EMA-82 (Plate I, 82.a-c.): fungal spore or algal cyst, globose, 6–18 μm in Ø, thick wall, hyaline and psilate. EMA-82 resembles spores of numerous fungi (e.g. Chroelosporium state of Peziza ostracoderma Korf; ref. Ellis, 1976) and cysts of algae (e.g. Chloromonas or Chlamydemonas). Recorded in peat core (Barthelmes, 2009). In present study frequent in all vegetation types, mostly scarce to numerous. No obvious indication. EMA-83A (Plate I, 83A): possibly conidium of Monodictys paradoxa (Corda) Hughes, known from bark, trunks and thick branches of Betula (Ellis and Ellis, 1997) and Sorbus aucuparia in Belarus (Yurchenko, 2001). Globose, 13–25 μm in Ø, muriform: composed of single cells of 2–3 μm in Ø, sometimes with hyaline triangular appendage (=remain of conidiophore?), middle to dark brown, psilate. Similar to EMA-83B. Here occasionally recorded in alder carr, birch carr and open fen, scarce to highly abundant. Exceptionally high value (940%) in the one birch wood sample from birch carr, thus clearly a local signal; lacking in birch dead wood of alder carr. However, also in alder dead wood, hummock mosses, but also in bare peat and grass litter of open fen vegetation. EMA-83A indicates here, if highly abundant (a few 100%),

presence of birch, and possibly birch carr conditions; but unclear indication in lower abundances. EMA-83B (Plate I, 83B): (as the similar EMA-83A) also possibly conidium of Monodictys paradoxa (Corda) Hughes, known from bark, trunks and thick branches of Betula (Ellis and Ellis, 1997) and Sorbus aucuparia in Belarus (Yurchenko, 2001). Elliptic to globose, 16–32 × 12.5-29 μm, brown to black, psilate, spore muriform (only visibly in lighter individuals), sometimes with hyaline tri- or quadrangular appendage (=remnant of conidiophore?). Here rarely recorded in alder carr, birch carr and open fen, scarce to highly abundant. Exceptionally high value (440%) in the one birch wood sample from birch carr, thus clearly a local signal; lacking in birch dead wood of alder carr. Abundant also in alder dead wood in alder carr and bare peat in open fen vegetation. EMA-83B indicates here, if highly abundant (a few 100%), presence of birch, and possibly birch carr conditions; unclear indication in lower abundance. EMA-86 (Plate I, 86.a+b): Bactrodesmium-like conidium, cylindrical elongated, ends tapering, often only fragments preserved, 10–32 septa (sometimes with pores), max. 250 μm long, 6-8(−10) μm broad, single cells either cubic (6–8 μm long) or rectangular (11–15 μm long), rectangular cells sometimes with thin pseudoseptum, one end cell hyaline and truncate, the other rounded, end cells often absent, spore brown and psilate. Resembles, but is longer than HDV-125 Pals et al., 1980. Similar spores of unknown origin are presented in Chmura et al. (2006, ‘multi-celled ascospore with end cells missing’), Miola et al. (2006, ‘unidentified fungal spores’, plate III), and as UG-1097 in Gelorini et al., 2011 (as shorter and slightly broader in diameter). Recorded also in peat cores (Barthelmes, 2009; Barthelmes et al., 2012–this issue). This may be identical with Bactrodesmium longisporum M.B.Ellis (suggested by A. Aptroot), known from dead Alnus wood (Ellis and Ellis, 1997), which has conidia that are very variable in length and number of septa, but do agree in width. EMA-29 (Prager et al., 2006) might well refer to the same species. In present study occasional in open fen, mainly in superficial peat, scarce to recurrent. Probably produced by a peat decomposing fungus. Until now no indicative value as no peat samples from carr were analysed. EMA-87 (Plate I, 87): elaters of capsulae from a Hepatic, probably Lophocolea heterophylla (Schrad.) Dumort. (det. A. Aptroot), acicular, fragments filiform, helix-like, sometimes in bundles, 7–12 μm in Ø, 70–165 μm long, light yellowish green. In present study rare, scarce to recurrent, no obvious pattern. No clear indicator value. EMA-90 (Plate I, 90): probably conidium of Arthrinium cf. puccinioides (DC.) Kunze, polygonal, cushion-shaped, 11–13 μm in Ø, one pore (hilum?), brown, psilate. A. puccinioides is observed on dead leaves of e.g. Carex acutiformis (Ellis and Ellis, 1997). In present study occasional in open fen (sedge litter and peat), numerous to abundant. Indicates presence of sedges in open fen vegetation. EMA-91 (Plate I, 91.a + b): possibly ascospore of Ceriophora palustris (Berk. & Broome) Höhn, elliptic to fusiform, often slightly curved (=ventrally straight and dorsally curved), 1 septum (with pore), constricted at septum, slightly constricted at one end (3 μm

Plate II. (All scale bars are 10 μm): 92. EMA-92: poss. fungal seta or plant hair. 93.a + b. EMA-93: ascospore in different views. 94.a + b. EMA-94: poss. fragment of fungal fruit-body (synnema), surface and outline view. 95. EMA-95: fungal tissue. 96. EMA-96: fungal conidium. 97. EMA-97: fungal spore. 98.a + b. EMA-98: fungal fruit-body, site and polar view. 99. EMA-99: fungal spore. 105. EMA-105: poss. immature conidium. 106.a + b. EMA-106: poss. fungal or algal spore, outline and surface view. 107.a + b. EMA-107: poss. fungal or algal spore, surface and outline view. 108. EMA-108: fungal spore. 109. EMA-109: ascospore. 110. EMA-110: possibly fungal seta or plant hair. 111.a-c. EMA-111: unknown origin.

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before tip), 18–22× 7 μm, brown, usually striate (ca. 9 indistinct longitudinal light striae), sometimes rapture along striae. Similar paler spores without striae interpreted as immature spores (Plate I, 91.b). EMA-91 strongly resembles HDV-122 Pals et al., 1980 and HDV-1027 van Geel et al., 2011, ascospores of Munkovalsaria donacina (Niessl) Aptroot, which both are however not described as striate, as well as HDV-1038 van Geel et al., 2011 ascospores of Arecophila sp. (with at least longitudinal structures= slits mentioned). Ceriophora palustris is known from sedge leaves (Ellis and Ellis, 1997) and Iris (Dennis, 1981). Recorded also in peat core (Barthelmes et al., 2012–this issue). In present study only in very wet, open fen, scarce to recurrent. Indicates open vegetation. EMA-92 (Plate II, 92): possibly fungal seta or plant hair, acicular, at base abruptly thickened to 14 μm, 23–96 μm long, dark red, psilate. In present study rare, only in very wet open fen, scarce to numerous (in Phragmites australis litter). Indicates open vegetation (Phragmites australis?). EMA-93 (Plate II, 93.a + b): ascospore, elliptic lenticular, 8–10 × 7–8 × 4–5 μm, wall thinning at both ends, psilate, brown, in side view (foto b) conspicuously lighter band from pole to pole. Most probably a Coniochaeta sp. (det. A. Aptroot). The measurements and the lentiform (not discoidal) shape agree well with C. leucoplaca (Berk. & Rav.) Cain, known worldwide from soil, dung and various plant remains (e.g. Munk, 1957). In present study only in very wet reed fen (Phragmites litter and surface peat); numerous to abundant (> 1000%). Indicator for open vegetation, probably for reed fen. EMA-94 (Plate II, 94.a + b): possibly fragment of fungal fruit-body (synnema = densely packed conidiophores), irregular cylindrical, one end rounded, second detached, 87-95(− 120) × 28-31 μm, brown. Similar synnemata produced by e.g. Doratomyces (Domsch and Gams, 1970; Ellis and Ellis, 1998), and Phaeoisaria clavulata (Grove) Mason & Hughes (Ellis and Ellis, 1997). In present study only once recorded (litter in very wet reed fen), recurrent. Possibly indicates open vegetation with Phragmites. EMA-95 (Plate II, 95): fungal tissue, 100–140 × 50–70 μm, often 1–2 elliptic hollows (8–12 × 12–25 μm), single cells thick-walled and irregularly shaped but always compact, sclerenchymatic, brown. In present study only once recorded (litter in very wet reed fen), recurrent. Possibly indicates open vegetation with Phragmites. EMA-96 (Plate II, 96): fungal conidium, cylindrical with acuminate ends, 5 septa, slightly constricted at septa, 58–60 × 17 μm, both central cells 21–22 μm long, neighbouring cells conical, 4 μm long, end cells obtuse cones, 5 μm long, spore pale brown, end cells hyaline, psilate. In present study only recorded once (superficial peat in very wet Common reed fen), scarce. No indicative value. EMA-97 (Plate II, 97): fungal spore, elongated ovoid (like the sole of a shoe), 1 septum, 18–19 × 5 μm, shorter cell 6–6.5 μm long, constricted near septum, hyaline, rounded at the end, larger cell 12 μm long, slightly tapering at the end, pale brown, spore psilate. In present study restricted to very wet sedge fen; scarce to recurrent, highest value in litter of Carex riparia. Indicator for open vegetation and sedge fen (Carex riparia?). EMA-98 (Plate II, 98.a + b): fungal fruiting body. Hemispherical to frustrum-shaped, 42–51 μm in Ø, 21–25 μm high, ostiole on top

6–7 μm in Ø, brown, irregular cell pattern. In present study only in very wet sedge fen; scarce. Indicates open vegetation and sedge fen. EMA-99 (Plate II, 99): fungal spore, elliptic, 1 septum (with pore), 19–21 × 7–8 μm, pale brown, psilate. In present study only once recorded (in peat of Carex riparia fen), recurrent. Fungus producing EMA-99 might be a peat decomposer, which requires very wet and open conditions. EMA-105 (Plate II, 105): possibly immature conidium, globose, 13 μm in Ø, with large protruding aperture (hilum), 4 cells, cruciform muriform, septa with pore, light brown, psilate. Most resembles immature conidia of Monodictys levis (Wiltshire) Hughes (suggested by A. Aptroot), known from various rotting materials including feathers (Ellis and Ellis, 1998). In present study frequent in wet sedge fen, scarce to recurrent. In carr sites rare, scarce. Indicator for open vegetation (sedge fen). EMA-106 (Plate II, 106.a + b): possibly a fungal or algal spore, globose, 17–20 μm in Ø, light brown, verrucate (verrucae Ø 2–3 μm, 1 μm high). Resembles ascospores of Scutellinia trechispora (Berk. & Br.) Lambotte; a fungus found on mosses and open soil (Ellis and Ellis, 1998). In present study occasional in wet sedge fen, recurrent to abundant (in herbal litter). Outside rare, scarce. Indicates open vegetation and sedge fen. EMA-107 (Plate II, 107.a + b): possibly fungal or algal spore, globose, 42–45 μm in Ø, pale green to hyaline, verrucate (verrucae irregularly distributed and shaped, 5–6 μm in Ø, 3–4 μm high). Similar, but larger than HDV-150 van der Wiel, 1982, and HDV-225 van Geel et al., 1989. In present study only in wet sedge fen, scarce to recurrent. Indicates open vegetation, might be related to Phalaris. EMA-108 (Plate II, 108): fungal spore, 3 septa, elliptic, truncate when end cells (often!) detached, constricted at middle septum, middle septum thick and with pore, 38–50× 15–17 μm, dark brown, psilate. Determination difficult as many fungi produce similar 4-celled ascospores (van Geel, pers. com.). EMA-108 resembles strongly UG-1159 Gelorini et al., 2011 (which is thicker), slightly HDV-16 C van Geel, 1978 (shorter, thinner, septum thinner), and HDV-121 Pals et al., 1980/HDV-970 Garneau, 1993 (not constricted at middle septum). In present study only in wet sedge fen, scarce. Indicates open vegetation, might be related to Phalaris. EMA-109 (Plate II, 109): ascospore, elliptic acuminate obtuse, 2 strongly protruding pores, 33–34 × 16–18 μm, wall thick, dark brown, faint longitudinal furrows. Probably a species of Sphaerodes, e.g. agreeing well with S. episphaeria (Phil. & Plowr.) Clem. (suggested by A. Aptroot), which occurs on other fungi (Ellis and Ellis, 1998). Resembles HDV-55B van Geel, 1978 (psilate). In present study rare, scarce. Might indicate open vegetation and sedge fen. EMA-110 (Plate II, 110): possibly fungal seta or plant hair, acicular, at basis broken, max. 138 μm long, widest (6 μm) at basis, 1–2 septa, dark reddish brown, psilate. In present study only in wet sedge fen, scarce. Might indicate open vegetation and sedge fen. EMA-111 (Plate II, 111.a-c): unknown origin, irregularly elliptic lenticular, polar areas concave, 19–34 × 15–27 μm, brown, psilate. In present study only in wet sedge fen, values of 39.4% (Phalaris litter) and 2973% (Carex litter) indicate local production. Indicates open vegetation and sedge fen.

Plate III. (All scale bars are 10 μm): 112. EMA-112: poss. conidium of Endophragmia collapsa. 113.a + b. EMA-113: fungal spore, 2- and 3septate form. 114.a + b. EMA-114: poss. fruit-body appendage of Phyllactinia sp., single and grouped individuals. 117.a + b. EMA-117: poss. wood remain (Betula?), single and grouped individuals. 118.a + b. EMA-118: poss. fungal spore, single individual and short chain. 119.a + b. EMA-119: fungal spore (conidium), two development stages? 120.a + b. EMA-120: prob. conidium of Brachysporium britannicum, 2- and 1septate form. 121. EMA-121: ascospore, prob. of Sordariaceae. 122.a + b. EMA-122: unknown origin, different development stages?. 123. EMA-123: conidium.

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EMA-112 (Plate-III, 112): possibly conidium of Endophragmia collapsa B. Sutton. Ovoid to ovoid elongated, 2–3 septa, constricted at septa, one end cell rounded, opposite end cell slightly tapering, thinner walled and mostly collapsed or at least crumpled, hyaline, whole spore 12–15 × 7–8 μm, round end cell and central cell(s) ca. 4 μm long, pale brown, spore psilate. E. collapsa is observed on dead Alnus (Ellis, 1976). In present study occasional in alder carr; scarce to recurrent. Scarce in litter in birch carr. Indicates presence of trees (alder, birch?) (also roots?). EMA-113 (Plate III, 113.a+b): fungal spore, ovoid to ovoid elongated, 26–28×10–12 μm, 2–3 septa (2 septa partition spore in unequally large cells; 3 septa in equally large cells), slightly constricted at septa (with pore), one end cell rounded, opposite end cell conical, hyaline, spore psilate, pale greyish brown (except one end cell). In present study occasional in alder carr; scarce to abundant. Highest values in dead wood of birch (in alder and in birch carr) and in mosses. Indicator for presence of trees (birch, alder?) (also roots?). EMA-114 (Plate III, 114.a + b): possibly fruit body appendage (seta) of Phyllactinia sp. (det A. Aptroot). Acicular to elongated club-shaped, base (10 × 8–9 μm) constricted and kneeled, main part tapering from 7 to 3 μm Ø to rounded or truncated tip, whole remain 52–116 × 8–9 μm, pale brown, psilate, sometimes groups of EMA-114 with detached tissue remain observed (photo b). P. guttata (Fries) Léveillé is known from leaves (Dennis, 1981). In present study rare in alder carr, mostly scarce. Present in birch carr (litter and mosses). Highest values in birch and sedge litter. Indicates carr vegetation. EMA-117 (Plate III, 117.a + b): possibly wood remain (Betula?), lenticular, outline irregular, circular to polyangular, single ‘cells’ 13–22 μm in Ø, brown, psilate, often pitted (fungal decay?), sometimes EMA-117 embedded in a hyaline matrix (photo b). In present study rare, mostly scarce. One exceptionally high value (4,061%) in dead wood of birch (in birch carr) however indicates that EMA-117 is formed in/from birch wood. Indicator for birch in birch carr (also roots?) and high decomposition. EMA-118 (Plate III, 118.a + b): possibly fungal spore. Globose, 9–14 μm in Ø, separate (foto .a) or in chains of up to 3 spores (foto .b), one aperture, light yellow, verrucate/baculate. Resembles HDV-225 van Geel et al., 1989 (which is hyaline and larger). Resembles conidia of Periconia byssoides Pers. ex Mèrat, cosmopolitan found on blackened areas, on dead herbaceaous stems and leaf spots, always associated with other fungi (Ellis, 1971; Ellis and Ellis, 1997). The observed distribution does not support this identification. In present study only in birch carr. High value in dead wood. Indicator for presence of birch in birch carr (also roots?). EMA-119 (Plate III, 119.a + b): fungal spore (conidium), fusiform to club-shaped, slightly curved, 62–71× 12–14 μm, with up to 7 septa (pseudosepta or post mortem dissolving?), one end tapering to a blunt tip, opposite end with a hyaline appendage with pore (hilum) or truncated, non-septate forms probably immaturate, pale brown to greyish brown, psilate. Very similar to HDV-1037 van Geel et al., 2011 (cf. Helminthosporium sp., but with 70–260 μm far to large), resembles also HDV-339 van Geel et al., 1980/81 (shorter, thicker, not curved), HDV-147 van der Wiel, 1982 (thicker, not curved, septa only at one end), as well as HDV-1034 van Geel et al., 2011 (shorter and with additional longitudinal septa, verrucate, and long narrow cervical above attachment place). In present study restricted to birch carr (birch litter and mosses); scarce. Indicates in situ presence of birch/birch carr. EMA-120 (Plate III, 120a + b): probably conidium of Brachysporium britannicum Hughes, ovoid, 2 (rarely 1) septa (with pore), 17–19× 10–13 μm, proximal cell hyaline, with aperture and often hyaline remnant of conidiophore, other cells brown, spore psilate. B. britannicum is observed on wood and bark of Betula and other deciduous trees (Ellis, 1971; Ellis and Ellis, 1997). Similar to HDV-1036 van Geel et al., 2011, also attributed to genus Brachysporium (but with 22–28 μm to long). In present study only in birch carr. High value (529%) in dead wood. Indicates presence of birch (also roots?) in birch carr.

EMA-121 (Plate III, 121): ascospore, probably of Sordariaceae, most likely a Sordaria sp. Elliptic, slightly compressed, 9–14 × 7–9 μm, 1 pore, dark brown, psilate. Resembles HDV-55A1 van Geel et al., 1980/81 (larger). In present study only twice recorded, high value (53%) in a moss sample from birch carr. Might indicating dung. EMA-122 (Plate III, 122,a + b): unknown origin. Discus with elevated ring on both sides, 11-12 μm Ø, ring Ø variable (possibly depending on development stage), light greenish yellow to hyaline, psilate. In present study only twice recorded, high value (40%) in dead wood of birch in birch carr. Indicates presence of birch (also roots?) in birch carr. EMA-123 (Plate III, 123): conidium. Club-shaped, distal end rounded, proximal end tapering, sometimes with long hyaline appendage (conidiophore or hyphae?), sometimes slightly curved, 7–12 septa, 24–37 μm long, max. 2.5-3.5 μm thick, single cells 2–5 μm long, longest cells at tapering end, conidium hyaline, psilate. In present study occasional in alder carr, scarce to numerous. Highest values in hummock mosses but also abundant in (alder and birch) dead wood. Indicates presence of alder carr, possibly drier habitats. EMA-124 (Plate IV, 124.a + b): possibly conidium. Ovoid to elliptic, 1 septum (with pore), strongly constricted at septum, one end with large aperture (3.3 μm Ø), second rounded, spore 27–28 × 8–10 μm, hyaline to pale brown, longitudinal rugulate to striate. In present study rare in alder litter, scarce. Clear indicator for above ground remains and thus presence of alder. EMA-125 (Plate IV, 125): probably conidium of Corynesporopsis quercicola (Borowska) P.M.Kirk (det. W. Gams). Elliptic to cylindrical, usually 2 septa, 15–18 × 6-7 μm, (1 in 100 3-septate, 20–22 × 6–7 μm), septa with pore, central cell(s) bright brown, longer (6–8 μm) than hyaline end cells, spore psilate, small aperture in one or both end cells (hilum?). C. quercicola is observed on dead wood of Quercus (Ellis and Ellis, 1997) and other trees (Kirk, 1981). In present study occasional in alder carr, scarce to numerous. Clearly concentrated (collected or produced?) in hummock mosses. One high value (24%) in alder wood. However, high values of EMA-125 also correspond to the distribution of Quercus seedlings. Might indicate presence of trees, including alder (also roots?). EMA-126 (=EMA-126A + B)(Plate IV, 126A + B): fragments of the same unknown (Helicoon/Helicodendron) fungal spore: coiled filamentous conidium forming a convex cylinder (EMA-126A) with lid and base (EMA-126B). Filament 10 μm Ø, brown, scabrate, septate, cells 5–9 μm long, cylinder 46–55 μm Ø and 43–75 μm long. EMA-126A: fragment of the cylinder, 4–7 windings/whorls with usually 5 visible septa (full winding ~ 15-20 septa). EMA-126B: fragment of the convex base/lid, filament tapering towards centre. Similar to conidia of Helicoon fuscosporum Lindner (Ellis and Ellis, 1997), which has the filaments = coils only 3–4 μm, whole spore-body only 25–30 μm in Ø, and fewer septa per coil plane; known from wet stems of Rubus idaeus. Also similar to conidia of Helicodendron conglomeratum Glenn-Bott (spore-body only 50 × 30 μm), on fallen leaves of deciduous trees decaying submerged in more or less static water in GB (Ellis, 1976). In present study EMA-126A and B are similarly distributed: occasional in alder carr, scarce. Indicator for alder carr (also roots?). EMA-127 (Plate IV, 127.a + b): possibly spore of Diderma (Myxomycota, det. M. Schnittler). Globose, 11 μm Ø, pale brown, clavate to pilate, ‘pilae’ max. 1 μm long. In present study occasional in alder carr, mostly scarce, one exceptionally high value (30%) in hummock mosses. Might indicate alder carr. EMA-128 (Plate IV, 128.a+b): probably conidium of Actinocladium rhodosporum Ehrenb. ex Pers. (det. A. Aptroot). Tripod-like with conidiophore (‘tripod head’), total length 46–78 μm, centre muriform, ‘legs’ 5–6 μm wide at base, tapering towards ends, 21–54 μm long, tips torn off, proximal septated (2–3 septa in the first 15 μm), ‘legs’ in angle of 40-70° to conidiophore axis, conidiophore 3–4 μm wide at base, greyish brown, psilate. A. rhodosporum observed on wood and bark of numerous

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Plate IV. (All scale bars are 10 μm): 124.a + b. EMA-124: possibly conidium, surface and outline view. 125. EMA-125: prob. conidium of Corynesporopsis quercicola. 126A,B. EMA-126A + B: fragments of the same unknown (Helicoon/Helicodendron?) fungal spore: of convex cylinder (EMA-126A), of lid or base (EMA-126B). 127.a + b. EMA-127: poss. spore of Diderma (Myxomycota), outline and surface view. 128.a + b. EMA-128: prob. conidium of Actinocladium rhodosporum, less or more torn off. 129.a + b. EMA-129: fungal spore (conidium). 130. EMA-130: fungal conidium. 134. EMA-134: fungal spore.

trees (Ellis, 1971), also alder (Yurchenko, 2001). In present study rare in alder carr, scarce to recurrent. Highest value (27%) in dead wood of alder. One observation in sedge fen, scarce. Indicates presence of alder (also roots?). EMA-129 (Plate IV, 129.a+ b): fungal spore (conidium). Ovoid to pyriform, 1 septum (with pore), slightly constricted at septum, 13–19× 6–9 μm, apical cell rounded and usually larger, basal cell

conical, with protruding aperture (hilum), whole spore hyaline to pale brown, psilate, often dented. Resembles conidia of numerous fungi, e.g. Cordana, Arthrobotrys, Hypomyces, Drepanopeziza, Endophragmia and Endophragmiella and thus can include spores of many taxa. In present study rare in alder carr, clearly concentrated in alder dead wood. One observation in sedge fen, scarce. Indicates presence of alder (also roots?).

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EMA-130 (Plate IV, 130): fungal conidium, ovoid, 1 septum (with pore), 11–13 × 9–10 μm, smaller cell 4–5 μm long, aperture (hilum), larger cell rounded, brown (darker than smaller cell), spore psilate. In present study rare in alder carr, scarce. Might indicate alder carr vegetation. EMA-134 (Plate IV, 134): fungal spore. Chain of 4–8 cells, 14–20 μm long, single cells globose, flattened towards neighbouring cells, 3–7 μm Ø, chain curved or kneeled, pale brown to hyaline, psilate. EMA-134 has few distinct features and might thus represent spores of species from, e.g. Sporormia, Sporormiella (which are however often dark) and Gliomastix murorum. Recorded also in peat core (Barthelmes et al., 2012–this issue). In present study occasional, scarce to abundant. Highest values in alder litter but recurrent also on other substrates in alder carr and in sedge fen. Clearly indicates, at least if abundant, presence of alder carr vegetation. Acknowledgements We thank Dr. Bas van Geel for helping in identifying non-pollen palynomorphs and inspiration, Dr. Walter Gams (Utrecht, The Netherlands), Prof. Martin Schnittler, Dr. Dierk Michaelis for their help in determining new NPP types, Dr. Ingo Koska, Andreas Kaffke, Martin Schumann, Alexander Seuffert, Annette Köber for measuring site conditions, and Erika Retzlaff (all Greifswald) for sample preparation. Two anonymous referees are acknowledged for helpful comments on the manuscript. The study was supported by DBU scholarship programme number 2002/396. References Apinis, A.E., Chesters, C.G.C., Taligoola, H.K., 1972. Colonisation of Phragmites communis leaves by fungi. Nova Hedwigia 23, 113–124. Apinis, A.E., Chesters, C.G.C., Taligoola, H.K., 1975. Microfungi colonizing nodes and internodes of aerial standing culms of Phragmites communis Trin. Nova Hedwigia 26, 495–507. Bakker, M., van Smeerdijk, D.G., 1982. A Paleoecological study of a late Holocene section from “Het Ilperveld”, Western Netherlands. Review of Palaeobotany and Palynology 36, 95–163. Barron, G.L., 1968. The Genera of Hyphomycetes from soil. The Williams & Wilkins Company, Baltimore. Barthelmes A., 2009. Vegetation dynamics and carbon sequestration of Holocene alder (Alnus glutinosa) carrs in NE Germany. PhD Thesis, Greifswald University, Greifswald. Barthelmes, A., Prager, A., Joosten, H., 2006. Palaeoecological analysis of Alnus wood peats with special attention to non-pollen palynomorphs. Review of Palaeobotany and Palynology 141, 33–51. Barthelmes, A., de Klerk, P., Prager, A., Unterseher, M., Joosten, M., 2012. Expanding the approach of NPP analysis to eutrophic and forested sites - Part II: Occurrence and significance of (surface sample) NPPs in a Holocene wood peat section. Review of Palaeobotany and Palynology. 186, 22–37 (this issue). Berglund, B.E. (Ed.), 1986. Handbook of Holocene Palaeoecology and Palaeohydrology. John Wiley and Sons, Chichester. Blackford, J.J., Innes, J.B., 2006. Linking current environments and processes to fungal spore assemblages: surface NPM data from woodland environments. Review of Palaeobotany and Palynology 141, 179–187. Breitenbach, J., Kränzlin, F., 1984–2000. Pilze der Schweiz, Bd. 1–6. Mycologia, Luzern (in German). Carlile, M.J., Watkinson, S.C., Gooday, G.W., 2001. The Fungi, 2nd ed. Academic Press, London. Chmura, G.L., Stone, P.A., Ross, M.S., 2006. Non-pollen microfossils in Everglades sediments. Review of Palaeobotany and Palynology 141, 103–119. Cugny, C., Mazier, F., Galop, D., 2010. Modern and fossil non-pollen palynomorphs from the Basque mountains (western Pyrenees, France): the use of coprophilous fungi to reconstruct pastoral activity. Vegetation History and Archaeobotany 19, 391–408. Czeczuga, B., Orlowska, M., 1995. Attractiveness of leaves of twenty-five tree species for aquatic Hyphomycetes representatives. Roczniki Akademii Medycznej w Bialymstoku 40/2, 233–242. Davis, O.K., Shafer, D.S., 2006. Sporormiella fungal spores, a palynological means of detecting herbivore density. Palaeogeography Palaeoclimatology Palaeoecology 237, 40–50. Dennis, R.W.G., 1981. British Ascomycetes, Revised ed. Cramer Vaduz. Dilly, O., Wachendorf, C., Irmler, U., Blume, H.-P., Munch, C., 1996. Changes of abiotic and biotic parameters in the course of decomposition of leaf litter in a Black Alder (Alnus glutinosa (L.) Gaertn.) forest of a moranic landscape of Northern Germany. Ecosystems 6, 31–42. Döbbeler, P., 1978. Moosbewohnende Ascomyceten. I. Die Pyrenocarpen, den Gametophyten besiedelnden Arten. Mitteilungen der Botanischen Staatssammlung München 14, 1–360 (in German).

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