A BM-2 (96 cm), B HdV 83 (14 cm), C Delitschia sp. (84 cm), D Brachysporium bloxami (84 cm), E Brachysporium pendulisporum (96 cm), F Puccinia -type (38 cm), G HdV 10 (4 cm), H Meliola ellisii (4 cm), I HdV 20d (4 cm), J BM-3 (32 cm), K HdV 463 (108 cm), L Helicoon pluriseptatum (52 cm), M Dictiosporium australiense (BM-4) (112 cm), N Geoglossum sphagnophilum (40 cm). Bar scale 10 μm. The in di ca tion in cm re fers to the po si tion in the core. 

A BM-2 (96 cm), B HdV 83 (14 cm), C Delitschia sp. (84 cm), D Brachysporium bloxami (84 cm), E Brachysporium pendulisporum (96 cm), F Puccinia -type (38 cm), G HdV 10 (4 cm), H Meliola ellisii (4 cm), I HdV 20d (4 cm), J BM-3 (32 cm), K HdV 463 (108 cm), L Helicoon pluriseptatum (52 cm), M Dictiosporium australiense (BM-4) (112 cm), N Geoglossum sphagnophilum (40 cm). Bar scale 10 μm. The in di ca tion in cm re fers to the po si tion in the core. 

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In order to reconstruct regional vegetation changes and local conditions during the fen-bog transition in the Borsteler Moor (northwestern Germany), a sediment core covering the period be tween 7.1 and 4.5 cal kyrs BP was palynologically investigated. The pollen diagram demonstrates the dominance of oak forests and a gradual replacement of trees by...

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Context 1
... swamp-bog tran si tion as sem blage in cludes as co - spores of Cercophora sp., Coniochaeta lignaria, Podospora sp. (Fig. 5M); Delitschia sp. (Fig. 6C) and Sordaria sp., which be long to mostly fac ul ta tive coprophilous groups, in - clud ing some spe cies that also grow on other kinds of de cay - ing or ganic ma te rial (Lundquist 1972, Rich ard son 2001, Krug et al. ...
Context 2
... (Fig. 6A) are 3-celled rhom boid fun gal spores, 27-28 × 10-12 µm. The up per two brown-col oured cells are di vided by a sep tum with a dark-brown rim of 6-7 µm breadth and the lower cell is hyaline with a pore of 1 µm. As BM-2 co in cides with max i mum of wood-de cay ing fun gal spores, the type might rep re sent conidiospores of saprotro- ...
Context 3
... Geel (1978) cor re lated find ings of HdV 83 ( Fig. 6B) with those of type HdV 106, a rep re sen ta tive of rel a tively wet, oligotrophic bog con di tions later iden ti fied as eggs of the oribatid mite Rhysotritia ardua (C.L. Koch) (Bakker, van Smeerdijk 1982). In the Borsteler Moor, HdV 83 oc curs more of ten es pe cially in wet phases of the Sphag num peat ...
Context 4
... 360 was also at trib uted to the ge nus Brachyspo- rium (van Geel et al. 1981) and de scribed as a fun gal spore of 19-22 × 14-17.5 µm ex clud ing the hyaline cells (1-3) sit u - ated at ei ther end. The spore body is strongly pig mented, with a gran u lated in ner wall side, and some times a hyaline "epi- sporium" is seen. This type (Fig. 6E) is very sim i lar in mor - phol ogy and size to conidiospores of Brachysporium pendu- lisporum S. Hughes, which are fusoid to limoniform, over all 30-42.5 × 15-17.5 µm, 4-5-septate, with a brown to dark- brown cen tral cell that is sep a rated by thick septa from the small hyaline po lar cells (Réblova, Seifert 2004;Markovskaja, ...
Context 5
... swamp-bog tran si tion as sem blage in cludes uredo - spores and teleutospores of the Puccinia-type (Fig. 6F;van Geel et al. 1981: HdV 357). A strong cor re la tion of HdV 10 ( Fig. 6G) with Calluna vulgaris pol len, seeds, leaves and pos si bly roots has been dem on strated in sev eral re cords (van Geel 1978). HdV 10 is there fore a good in di ca tor of lo cally dry con di tions in raised bog peats (e.g. van Geel 1978, Kuhry 1985. HdV 10 can also cor re late with Erica tetralix re mains (Bakker, van Smeerdijk 1982). ...
Context 6
... other fun gal spore as so ci ated with Calluna vulgaris is the ascomycete Meliola ellisii Roum (Fig. 6H), which was sug gested to have been a com mon par a site on C. vulgaris in peat bogs dur ing the At lan tic to Subatlantic pe riod (van Geel 1978: HdV 14). In the Borsteler Moor, spores of Meliola ellisii are char ac ter is tic for Sphag num peat, con firm ing that Calluna vulgaris grow ing on min eral soils is not in fected (van Geel ...
Context 7
... 20d (Fig. 6I) is char ac ter ized by ta per ing api cal ends, in con trast to the three-septate as co spores of HdV 20 (van Geel 1978). Van Geel (1978) sug gested Empetrum as pos si ble host of HdV 20. In the Borsteler Moor, HdV 20d oc - curs in low fre quen cies dur ing the oligotrophic ...
Context 8
... ( Fig. 6J) is a 3-4-septate, pale, slightly curved fun gal spore of 51-52 × 10 µm, with rounded ends, each end with an api cal pore of 1.5 µm. The septa are very thin, some - times dis placed or dis solved. In the Borsteler Moor, BM-3 oc curred in just one sam ple (34 cm) dur ing a wet phase of the oligotrophic ...
Context 9
... also found HdV 463 (Fig. 6K) de scribed from the Amtsven sec tion (Kuhry 1985) and as sumed to be a Stuar- tella spe cies. Fol low ing the fig ures in Müller (1962), this iden ti fi ca tion might be doubt ...
Context 10
... pluriseptatum van Beverwijk (Fig. 6L) is known from peat bogs and marshy places on birch leaves, pine nee dles, pine cones, leaves of red oak, and grass blades (Van Beverwijk 1954in van Geel 1978. In the Borsteler Moor, H. pluriseptatum is re stricted to the first wet Sphag num ...
Context 11
... BM-4 ( Fig. 6M) has pale brown conidia of 38-40 × 10-12 µm, con sist ing of one trun cate cell with 3 ver ti cal straight or slightly curved cy lin dri cal 8-septate arms of more or less sim i lar length, ar ranged close to each other. Mor pho - log i cally, the type is very sim i lar to Dictyosporium austra- liense Sutton 1985(Sutton 1985, Goh et al. ...
Context 12
... sphagnophilum Ehrenb. (Fig. 6N) is re - ported to grow among Sphag num and its spores were found with Sphag num re mains (van Geel 1978: Type 77A). The spores were en coun tered from the At lan tic and Subboreal pe - ri ods, es pe cially in the up per parts of hum mocks, just be fore the wet Scheuchzeria palustris over grow ing phases and in a layer char ac ter ized ...

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... In each sample about 700 pollen grains of arboreal pollen (AP), along with non-arboreal pollen (NAP) and spores of cryptogamous plants, as well as NPPs were counted, determined, and identified with specific keys (e.g. van Geel, 2001;Beug, 2004;Miola, 2012;Shumilovskikh et al., 2015) and with the use of Non-Pollen Palynomorph Image Database (NPP-ID) (Shumilovskikh et al., 2022). ...
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This paper reports on new, previously undescribed non-pollen palynomorphs (NPPs) of fungal origin preserved in moss polsters in a forest environment of a temperate zone. Modern NPPs were recovered from moss samples from 114 sites located in different types of forest ecosystems of Białowieża Forest, north-eastern Poland – a forest complex with a high degree of naturalness, similar to forests that covered large areas of Europe in the past. 98 types of fungal NPPs were identified, of which 20 morphotypes presented here are considered new, not previously described. Half of the new remains could be identified to species or genus level, thereby conferring ecological indicator value to them. Each new, determined NPP can be a useful tool in environmental reconstructions. This study shows that moss polsters in forested areas contain many undescribed NPPs, the description and analysis of which can deliver additional ecological information about ecosystems.
... Non-pollen palynomorphs (NPP) range in size about 10-250 μm, which can be observed during routine pollen counting (Shumilovskikh and van Geel, 2020;Shumilovskikh et al., 2015a). To analyze NPP, their percentages were calculated based on the total pollen counting. ...
... Pollen was identified using a Zeiss Axio Lab A1 microscope at 400x magnification following Reille (1992) and Beug (2004). Non-pollen palynomorphs (NPPs) identification is based on Van Geel (1978), Shumilovskikh et al. (2015) and NPP Image Database at http:// nonpollenpalynomorphs.tsu.ru (Shumilovskikh et al., 2022). ...
... Conspicuous charcoal peaks -which concentration reached 300-4500 particles cm −3 in the subzone 1a, and 100 particles cm −3 in subzones 1b and 1c -suggest a high fire frequency during this period. The appearance of Onagraceae pollen and Gelasinospora (HdV-1), occurring mainly in peat layers containing charcoal (Shumilovskikh et al., 2015;Van Geel, 1978), imply the presence of burned area within the vicinity of the sampling site. Fungal spores of Brachysporium sp. ...
... Fungal spores of Brachysporium sp. (HdV-359) are abundant in the subzone 1b, belonging to Brachysporium bloxami, a fungus that lives on decaying wood (Shumilovskikh et al., 2015). ...
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This paper presents a reconstruction of natural environmental dynamics, wildfires and vegetation change in northwest Putorana Plateau during the last 1300 years. The study area is a remote and poorly investigated region of subarctic Siberia, relatively untouched by human impacts, which offers a unique opportunity to examine natural environmental dynamics and climate-vegetation-fire relationships. The paleoenvironmental reconstructions are based on multi-proxy analysis of the Gervi peatland including pollen, plant macrofossil, testate amoebae and charcoal analysis, loss on ignition measurements and AMS radiocarbon dating. The results revealed the main phases of regional paleoenvironmental change: a warm period between 680 and 1200 C.E. (Common Era) corresponding to the Mediaeval Climate Anomaly (MCA), followed by climate cooling during the Little Ice Age (LIA, 1200–1850 C.E.) and subsequent centuries. Climate amelioration during the MCA led to afforestation of northwestern Putorana Plateau and an expansion of spruce extending 70 km northeast of its present geographical range. A quantitative water-table reconstruction was generated using a testate amoebae transfer function and suggested that relatively dry climate conditions during the MCA triggered high fire frequencies. The LIA appeared in the study area as a period of pronounced climate cooling and moderate moistening, which caused an extension of tundra vegetation and a dramatic decline of fire activity. Distinct environmental changes at the study site were detected since 1990 C.E., characterized by a high peat accumulation rate and rising water table. Since 1990 C.E., the macroscopic charcoal accumulation rate in the peat core increased abruptly, suggesting a recent increase in the fire frequency in the study region.
... The helicosporous conidia at Drzewce, similar to the modern genera Helicoon and Helicodendron, are barrel-shaped, enclosing the air that allowed buoyancy and the dispersal of the conidia (Zhao et al., 2007). Fossil helicosporous fungi from swampy environments previously were reported from Eocene to Holocene deposits (van Geel, 1978;Kalgutkar and McIntyre, 1991;Kalgutkar and Sigler, 1995;Shumilovskikh et al., 2015;Romero et al., 2021;Saxena et al., 2021). ...
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Palynological analysis of the 1st mid-Polish lignite seam (MPLS-1) of the Drzewce deposit (Konin region, central Poland) was used as the data source for palaeoenvironmental and palaeoclimatic interpretations. Lignites of the 1st group developed in the middle Miocene, during and shortly after the last peak of the Mid-Miocene Climatic Optimum, over a large area of Poland, and they are the youngest of the main Neogene lignite seams in Poland. In the Konin region, these lignites have a relatively significant thickness (up to 20 m) and therefore they are (or were in the past) exploited in several open-pit mines. A total of 36 palynological samples from the 6.3-m-thick seam of the Drzewce opencast mine was studied in detail. Palynological analysis of the lignite seam indicates that the area was overgrown by palustrine wetland communities, similar in composition to modern pocosins. The most characteristic elements of them were shrubs in the Ericaceae family. The climate at that time was warm temperate and humid. The estimated mean annual temperature (MAT) for the lignite seam at Drzewce is 15.7–17.8 °C. Comparison with other palynofloras from the MPLS-1 shows that the climate during the formation of the group of seams was more or less homogenous across the entire Polish Lowlands. Sedimentological data and results of palynological studies (including NPPs) at Drzewce indicate that the palaeomires were relatively distant from the channels of the river system in the Konin Basin. The fossil fungal assemblage indicates dense vegetation on damp, swampy soils and the presence of small, shallow-water bodies, with a variable water level or even periodic reservoirs, existing only during the wet season or after floods. In small, flooded depressions, such as the pools in bogs, filamentous green algae occurred. The presence of zygospores of the desmids Desmidiaceaesporites cosmarioformis most probably indicates relatively nutrient-poor (ombrotrophic) conditions. Fluctuations in the frequency of individual plant taxa (including Sequoia and Sciadopitys) are likely to reflect changes in water level and trophic conditions.
... The identification of pollens and spores followed Reille (1992Reille ( , 1995. The identification of non-Pollen Palynomorphs followed the main NPP atlas and publications describing NPP types (Cugny, 2011;Gelorini et al., 2011;Shumilovskikh et al., 2015;van Geel et al., 2003van Geel et al., , 2011. The identification of A minimum of 300 pollen grains were identified and counted as a pollen sum, according to the results obtained by the experimental approach developed in Sector D for samples coming from less organic sediments (Revelles, 2017d). ...
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The Early Neolithic lacustrine site of La Draga (Banyoles, Spain), where one of the first agri-cultural communities of the Iberian Peninsula settled between 7270 and 6750 cal. years BP, rep-resents an important school for bioarchaeological studies aiming to investigate the dialectic between human societies and vegetal environment in the past. In Lake Banyoles, the first evidences of human impact on the regional deciduous oak formations during the Mid-Holocene have been documented, thanks to a multi-scale approach that included pollen data from the archaeological site and from the associated lake sequence. In sector D located under the phreatic level, new methodologies of spatial analysis were applied in palynology, integrating non-pollen palino-morphs (NPPs) as good paleoecological indicators at a very local scale. The success of such methodologies allowed to describe the formation processes of the site, and evidenced patterns of social use of space by the Neolithic community in the past. In this work, a spatial approach in archaeopalynology is applied to Sector A of La Draga: in this sector, located at the highest topographical point of the site, the subaerial terrigenous sedimentation conditions have not allowed the conservation of organic matter and wooden elements, and a palimpsest of archaeological structures corresponding to different phases of occupation is observed. A total of 25 samples were analysed, from the horizontal sampling of three archaeological structures and two stratigraphic levels of the northern subsector, excavated between 2018 and 2021. The results of 51 horizontal samples from the southern subsector were also integrated to the analysis. The application of multivariate statistical analyses and spatial interpolation algorithms (IDW) enabled to identify the main climatic, taphonomical and anthropic factors associated to the formation of the horizontal pollen record in Sector A. In the pollen spectrum of Sector A, herbaceous plants dominate, as a result of direct and indirect human influence on the landscape in the past (deforestation, crops and use of plants in the deposit). The results show that the great spatial heterogeneity observed in the record is largely conditioned by taphonomical factors linked to terrigenous sedimentation in Sector A: subaerial conditions and soil erosion processes favoured the expansion of ferns in the area, and did not allow the development of coprophilous fungal organisms as observed in Sector D. However, it has been possible to identify some patterns linked to the incidence of human activity. Three structures appear associated with the use of fire, indicated by the accumulation of spores of carbonicolous and lignicolous fungi, and an accumulation of anthropic herbaceous taxa was identified to the Northeast of the sector in structure E263, indicating a possible area of storing and processing plants of socio-economic use. Finally, 14 new types of NPPs were described in Sector A. This new contribution to the understanding of the formation processes of the site and the use of space and plant resources by the first Neolithic societies in the north-eastern Iberian Peninsula demonstrates the validity of the spatial approach in archaeopalynology.
... Spores of Gelasinospora sp. (HdV-1) fungi -which are mainly carbonicolous and often grow on charred wood -and charcoal particles were common in PONE, providing additional independent evidence that fires occurred in the area adjacent to the lake and mountain slopes (Shumilovskikh et al., 2015;Van Geel, 1978). At the same time, pollen assemblages from PTHE demonstrate the highest peak in micro-charcoal values compared to other part of the sediment core, which reflects the high input of charcoal particles not only at the local, but also at the regional level. ...
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We present a reconstruction of vegetation dynamics and fire history from the western part of the Putorana Plateau during the Late-Holocene. The study area is located in the remote and poorly investigated region of Central Siberia, which represents an important area for understanding climate and environmental changes in the Russian Subarctic. Pollen and macroscopic charcoal data from three closely located lakes along an altitudinal transect in the Khantaika River basin show no major changes in vegetation in the study area during the last 3.9 ka BP. However, a detailed analysis of the data reveals an extension of forest coverage in lake catchments at about 3.1 ka BP followed by a gradual degradation of woodlands, and an expansion of shrubs and tundra vegetation at around 2.7–2.5 ka BP. Fire activity was relatively high between 3.0 and 2.0 ka BP compared to all but the most modern part of the record, while macroscopic charcoal accumulation reaches a maximum in the most recent surface sediments. This suggests an increase in the frequency and area of fires in the region since the end of the 19th century, which has no analog during the Late-Holocene.
... The zone is divided into four subzones. In this part of the profile, high proportion of Sphagnum and Tilletia sphagni spores suggest wet, oligotrophic conditions and presence of pools ( van Geel, 1978;Shumilovskikh et al., 2015). 5d Hyalosphenia-Enthophlyctis-Copepoda 25-10 Maxima of Hyalosphenia subflava and Empetrum/Ledum; continuous presence of Enthophlyctis lobata, Copepoda spermatophores and Gelasinospora; disappearance of Habrotrocha. ...
... Furthermore, abundance of mosses which colonised a hydrophobic peat surface following a fire event (Turner and Swindles, 2012 van Geel, 1978;Yeloff et al., 2007), and Assulina a high water level in the peat-bog (Lamentowicz and Mitchell, 2005). Also Habrotrocha points to the wet phase ( Shumilovskikh et al., 2015). Copepoda spermatophores indicates at least seasonally flooded area ( van Geel, 1978; van Geel and Middeldorp, 1988;Shumilovskikh et al., 2015). ...
... Also Habrotrocha points to the wet phase ( Shumilovskikh et al., 2015). Copepoda spermatophores indicates at least seasonally flooded area ( van Geel, 1978; van Geel and Middeldorp, 1988;Shumilovskikh et al., 2015). 5b Gelasinospora 145-105 Peak of Gelasinospora; presence of Gaeumannomyces but with percentage lower than previous subzone. ...
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Results of dendrochronological analysis of subfossil pine (Pinus sylvestris) buried in the Imszar raised bog deposits (NE Poland, Podlasie Region), were compared with multiproxy analysis of peat deposits (pollen and Non Pollen Palynomorphs, plant macrofossils, Cladocera, and geochemical analyses). The peat (3.3 m thick) accumulation started in the Allerød interstadial. The oldest pine tree population grew in this peatland in ca 6790–6530 cal yrs. BP: floating chronology (determined with the wiggle–matching method) covers this period is 255 years long. The growth of trees on the fen was related to the drying out of the fen at that time, which caused the break in peat accumulation (hiatus). Apart from numerous fallen trees excavated during the peat extraction, a horizon of vertical in situ tree trunks ingrown to the ground, were exposed during peat exploitation. On the basis of a dozen or so trees, a second, “dead forest” floating chronology 197 years long, was compiled with a time range of ca 4255-4060 cal yrs. BP. The youngest, third pine floating chronology (310 years long), elaborated on the basis of ex situ tree trunks, covers the time frame of ca 3130-2820 cal yrs. BP. Multiproxy analyzes of fen deposits indicate that the three stage colonization of the peatland by bog pine was associated with the periods of warming and overdrying in the climate of the Holocene, during which the groundwater table in the Imszar peatland significantly lowered. In turn, the dying off stages of the pine trees occurred in the phases of climate humidity growth, during which the groundwater level in the fen raised. They are well correlated with analogous bog pine germination and dying-off phases in the other European peatlands.
... Gill et al. 2009;van Geel et al. 2011b;Basumatary and McDonald 2017;van Asperen et al. 2021). Algal palynomorphs, zoological microfossils, and many fungal spores provide information on former local processes like natural and human-induced fire, hydrological conditions and erosion processes (Kuhry 1997;van Geel and Aptroot 2006;Montoya et al. 2010;van Geel et al. 2011a;Kołaczek et al. 2013;Schlütz and Shumilovskikh 2013;Shumilovskikh et al. 2015aShumilovskikh et al. , b, 2017McCarthy et al. 2021;Mudie et al. 2021). In geoarchaeology, NPPs contribute to a more holistic view of the past (Brinkkemper and van Haaster 2012;Revelles et al. 2016;Shumilovskikh and van Geel 2020;Shumilovskikh and Schlütz 2021). ...
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Full-text available
Non-pollen palynomorphs (NPPs) form a large group of biological objects found in palynological slides besides pollen grains. This includes various remains of algae and fungi, shells, resting stages and eggs of invertebrates, among others. Publications of NPP-types started in the 1970s with studies of BvG and colleagues, and large numbers of new types continue to be published every year. For an overview of this diverse world of “extra fossils”, we created the Non-Pollen Palynomorph Image Database (NPP-ID) to gather NPP knowledge, structured by acronyms and known taxonomy to assist identification and palaeoecological interpretation ( https://nonpollenpalynomorphs.tsu.ru/ ). An integral part is a database of illustrations, descriptions and ecological background of NPPs. While numerical data are routinely stored in open access repositories, the NPP-ID enables the definitions, identification and interpretation of the NPP taxa to be shared. The NPP-ID operates as an open research project aiming to provide open access to descriptions and illustrations of NPPs. However, due to publication rights, access to some original images is restricted and registration by users is required. We encourage palynologists to contribute to the further growth of the database by uploading their own microphotographs or drawings under an open access license. Contributors will be acknowledged by co-authorship in publications on updates of the NPP-ID.
... In contrast, copepod spermatophores (HdV-28) are well known from peat sediments (e.g. Rudolph 1917;Hesmer 1929;Frey 1964;van Geel 1978;Prager et al. 2012;Shumilovskikh et al. 2015a). Hesmer (1929) found the copepod Canthocamptus with spermatophores inside and provided the first description of this palynomorph. ...
... Its conidiospores are known from peat bogs in the Netherlands (HdV-30, van Geel 1978) and other peat soils of Europe and Canada (e.g. Kuhry 1997;Shumilovskikh et al. 2015a;Fig. 19b). ...
... The best-known representative of this family in palaeoecology is Ustulina (Kretzschmaria) deusta ( Fig. 21a-b) It is a weak parasite causing a soft rot of broadleaved trees, such as beech, oak, lime, horse chestnut and maple (Hartmann et al. 2007), and common on stumps of dead deciduous trees (Ellis and Ellis 1997). Their ascospores, as HdV-44, are known from a number of palaeoecological studies (van Geel 1978;Pals et al. 1980;van Geel et al. 1986;Barthelmes et al. 2012;Prager et al. 2012;Shumilovskikh et al. 2015a;Doyen and Etienne 2017). Amphisphaerella dispersella (HdV-310; Fig. 21g), another representative of Xylariaceae, is saprobic on Populus. ...
Article
Non-pollen palynomorphs (NPPs) are ‘extra’ microfossils often found in palynology slides. These include remains of organisms within the size range of pollen grains (∼10-250 microns), resistant to laboratory treatments used for the preparation of palynological samples. NPPs are a large and taxonomically heterogeneous group of remains of organisms living in diverse environments. Taxonomically, they belong to a wide variety of groups such as cyanobacteria, algae, vascular plants, invertebrates, and fungi. The aim of this chapter is to provide a general overview of NPP groups observed in palynology slides. It includes more than 40 of the most common groups starting with acritarcha, cyanobacteria and algae, moving through transitional groups to animals and fungi and finishing with human-made objects such as textile fibres. Although far from complete, it provides an updated overview of taxonomical diversity of NPPs and their indicator values. Further works on NPP identifications are of great importance to improve of our current knowledge. Since NPPs occur in all kinds of sediments, their analysis is a powerful tool for reconstructing environmental changes over time. Further detailed studies of specific NPP groups and their indicator values will open the way for new fields of study.
... reported the rare occurrence of 3septate conidia. The records from Germany and the Netherlands based on subfossil palynomorphs from peat samples(Shumilovskikh et al. 2015, van Geel et al. 1981 appear convincing according to the presented conditions of the conidia and are supported by collection of more extant material from the German Alps. Intercalary (subterminal) conidiogenous cells were not re-Brachysporium pendulisporum from Germany (R. Kirschner 263). A. Conidiophore with apically attached conidium. ...