NON-POLLEN PALYNOMORPHS

Major groups of the non-pollen palynomorphs

ACRITARCHS
ALGAE	Chlorococcales (coccal green algae) Cyanobacteria Desmidiales Dinoflagellate cysts Zygnemataceae
ANIMALIA	Helminths Mites Neorhabdocoela Rotifera Tardigrada Testate amoebae
BRYOPHYTA
FUNGI	Ascospores Conidiospores Spores of Glomeromycota Spores of rusts Spores of smuts
PLANTA
PROTOZOA
PTERIDOPHYTA
SPERMATOPHYTA
UNKNOWN	Unknown taxonomical position
UNORGANIC

Pediastrum and Botryococcus are very well known objects in palynology. Beside them Coelastrum (UG-1233), Tetraedron (HdV-66, HdV-371) and Scenedesmus (HdV-770, UG-1239) occur in pollen slides. Different species of Pediastrum demonstrate different responses to environmental parameters such as turbidity, water chemistry, nutrient status, pH and geography, making Pediastrum a good indicator for recording changes in the trophic status of a lake (e.g. Nielsen and Sørensen 1992, Jankovská and Komárek 2000). Identification and palaeoecological interpretation of the fossil green algae can be done using overview works of Komarek and Marvan (1992), Jankovská and Komárek (2000) and Komárek and Jankovská (2001).

References:
Komárek J, Jankovská V (2001) Review of green algal genus Pediastrum: implication for pollenanalytical research. Berlin, Stuttgart: Cramer.
Komárek J, Marvan P (1992) Morphological differences in natural populations of the genus Botryococcus (Chlorophyceae). Archiv für Protistenkunde 141: 65-100.
Jankovská V, Komárek J (2000) Indicative value of Pediastrum and other coccal green algae in palaeoecology. Folia Geobotanica 35: 59-82.
Nielsen H, Sørensen I (1992) Taxonomy and stratigraphy of late-glacial Pediastrum taxa from Lysmosen, Denmark – a preliminary study. Review of Palaeobotany and Palynology 74: 55-75.

Cyanobacteria

Cyanobacteria (formerly blue green algae) are photosynthetic prokaryotes able to produce oxygen and to fix nitrogen. They can be found in terrestrial and aquatic habitats and are known for causing blooms, which can be toxic. In fossil material, cyanobacteria, especially akinetes of Aphanizomenon (HdV-600) and Anabaena (HdV-601), showed phosphate-eutrophication of a Medieval lake as a consequence of intensification of farming and fertilization of the area around the lake (van Geel et al. 1994, 1996). There is no identification key for fossil Cyanobacteria and they can be identified with help of the descriptions in Ralska-Jasiewiczowa and van Geel (1992), van der Wiel (1982), van Geel et al. (1983b, 1994, 1996).

References:
Ralska-Jasiewiczowa M, van Geel B (1992) Early human disturbance of the natural environment recorded in annually laminated sediments of Lake Gosciaz, Central Poland. Vegetation History and Archaeobotany 1: 33-42.
van der Wiel AM (1982) A palaeoecological study of a section fromthe foot of the Hazendonk (Zuid-Holland), based on the analysis of pollen, spores and macroscopic remains. Review of Palaeobotany and Palynology 38: 35-90.
van Geel B, Hallewas DP, Pals JP (1983b) A Late Holocene deposit under the Westfriese Zeedijknear Enkhuizen (Prov of Noord-Holland, The Netherlands): palaeoecological and archaeological aspects. Review of Palaeobotany and Palynology 38: 269-335.
van Geel B, Mur LR, Ralska-Jasiewiczowa M, Goslar T (1994) Fossil akinetes of Aphanizomenon and Anabaena as indicators for medieval phosphate-eutrophication of Lake Gościąż (Central Poland). Review of Palaeobotany and Palynology 83: 97–105.
van Geel B, Odgaard BV, Ralska-Jasiewiczowa M (1996) Cyanobacteria as indicators of phosphate-eutrophication of lakes and pools in the past. Pact 50: 399-415.

Desmidiales

Desmidiales are the unicellular green algae of Conjugatophyceae. They inhabit mostly freshwater and may be found between Sphagnum mosses in peat bogs. Their cells are divided in two symmetrical parts separated by an isthmus. However, in the fossil stage they are represented by a half of the cell: Closterium idiosporum-type (HdV 60), Closterium cf. rostratum (HdV 372), Staurastrum (HdV 983). Together with other algal palynomorphs, desmids can be used as proxies of human impact on lacustrine ecosystem (McCarthy et al. 2018). Identification of the fossil remains to genus or species level is possible by using algological literature (Coesel and Meesters 2007, 2013 and Šťastný, 2010, 2013).

References:
Coesel PFM, Meesters KJ (2007) Desmids of The Lowlands - Mesotaeniaceae and Desmidiaceae - Identification key. Zeist: KNNV Publishing.
Coesel PFM, Meesters KJ (2013) European Flora of the desmid genera Staurastrum and Staurodesmus - Identification key for Desmidiaceae. Zeist: KNNV Publishing.
McCarthy FMG, Riddick NL, Volik O, Danesh DC (2018) Algal palynomorphs as proxies of human impact on freshwater resources in the Great Lakes region. Anthropocene 21: 16-31.
Šťastný J (2010) Desmids (Conjugatophyceae, Viridiplantae) from the Czech Republic; new and rare taxa, distribution, ecology. Fottea 10: 1-74.
Šťastný J (2013) Unveiling hidden species diversity in desmids (Desmidiales, Viridiplantae). PhD Thesis. Prague: Charles University in Prague.

Dinoflagellate cysts

Cysts of dinoflagellates (dinocysts) are found mostly in marine sediments but also in freshwater. They represent resting stages in the life cycle of dinoflagellates that are microscopic unicellular algae belonging to the phylum Dinoflagellata. Two systems exist for identification of the dinoflagellates: (1) for biologists based on motile cells and (2) for palaeontologists based on resting stages. Since only 10-20% of dinoflagellate species produce cysts composed of highly resistant dinosporin, dinocyst assemblages provide only limited information about the former ecological community. Nevertheless, numerous studies on the distribution of dinocysts highlight their usefulness for reconstruction of salinity, temperature, ice-cover and productivity or eutrophication (de Vernal and Marret 2007, Marret and Zonneveld 2003, Zonneveld et al. 2013). There are several NPP types from dinocysts: Lingulodinium machaerophorum (HdV 704A), HdV 703A, HdV 703B, HdV 703C, HdV 704B, HdV 704C. Dinocyst studies belong to a special branch of palynology and several common identification keys for Quaternary dinocysts exist as papers or on-line resources (Rochon et al. 1999; Mudie et al. 2017; https://www.marum.de/Karin-Zonneveld/dinocystkey.html ).

References:
De Vernal A, Marret F (2007) Organic-walled dinoflagellate cysts: tracers of sea-surface conditions. Developments in Marine Geology 1: 371-408.
Marret F, Zonneveld KAF (2003) Atlas of modern organic-walled dinoflagellate cyst distribution. Review of Palaeobotany and Palynology 125: 1-200.
Mudie PJ, Marret F, Mertens KN, Shumilovskikh L, Leroy SAG (2017) Atlas of modern dinoflagellate cyst distributions in the Black Sea Corridor: from Aegean to Aral Seas, including Marmara, Black, Azov and Caspian Seas. Marine Micropaleontology 134: 1-152.
Rochon A, de Vernal A, Turon J-L, Matthiessen J, Head MJ (1999) Distribution of recent dinoflagellate cysts in surface sediments from the North Atlantic Ocean and adjacent seas in relation to sea-surface parameters. AASP Contribution Series, 35. Dallas: American Association of Stratigraphic Palynologists Foundation.
Zonneveld KAF, Marret F, Versteegh GJM, Bogus K, Bonnet S, Bouimetarhan I, Crouch E, de Vernal A, Elshanawany R, Edwards L, Esper O, Forke S, Grøsfjeld K, Henry M, Holzwarth U, Kielt J-F, Kim S-Y, Ladouceur S, Ledu D, Chen L, Limoges A, Londeix L, Lu S-H, Mahmoud MS, Marino G, Matsouka K, Matthiessen J, Mildenhal DC, Mudie P, Neil HL, Pospelova V, Qi Y, Radi T, Richerol T, Rochon A, Sangiorgi F, Solignac S, Turon J-L, Verleye T, Wang Y, Wang Z, Young M (2013). Atlas of modern dinoflagellate cyst distribution based on 2405 data points. Review of Palaeobotany and Palynology 191: 1-197.

Zygnemataceae

Zygnemataceae are unbranched filamentous green algae of Conjugatophyceae, inhabiting shallow, stagnant, oxygen-rich freshwater lakes, ponds, small pools or wet soils. A few species inhabit salt water, but there are no marine representatives. During the sexual reproduction, Zygnemataceae produce thick-walled zygospores, which can fossilize and be found in the sediment archives. Apart from zygospores, asexual resting spores (aplanospores) can also be formed. The spores allow the algae to overcome unfavourable conditions such as drying of the sediment surface during summer or freezing during winter. In most cases the morphological characteristics of zygospores and aplanospores are necessary for identification to the species level. For information on extant Zygnemataceae, reference is made to Transeau (1951), Randhawa (1959), Kadlubowska (1984), and Hoshaw and McCourt (1988). Fossil spores recorded in pollen slides are: Mougeotia (HdV 61, HdV 133, HdV 134, HdV 135, HdV 136, HdV 141, HdV 313, HdV 373, HdV 711, HdV 772, HdV A, HdV B, t micro 418), Zygnema-type (HdV 213, HdV 314, t micro 419), Spirogyra (HdV 130, HdV 131, HdV 132, HdV 210, HdV 211, HdV 212, HdV 315, HdV 342, HdV 773, HdV D1, t micro 417, Tmic 19, UG 1240), Debarya (HdV 214). They have been described and their use as palaeoenvironmetal indicators was started by van Geel (1976). However, until now there is no formal identification key for spores of Zygnemataceae and the reference for fossil spores is made to a couple of papers with original descriptions and photographs of the types such as Ellis-Adam & van Geel (1978), van Geel and van der Hammen (1978), van Geel and Grenfell (1996), van Geel (1976, 1979, 2001), van Geel et al. (1989, 1994).

References:
Ellis-Adam AC, van Geel B (1978) Fossil zygospores of Debarya glyptosperma (De Bary)Wittr. (Zygnemataceae) in Holocene sandy soils. Acta Botanica Neerlandica 27: 389–396.
Hoshaw RW, McCourt RW (1988) The Zygnemataceae (Chlorophyta): a twenty-year update of research. Phycologia 27: 511–548.
Kadlubowska JZ (1984) Conjugatophyceae. I. Zygnemales. Süsswasserflora von Mitteleuropa. Band 16. NewYork: Fischer.
Randhawa MS (1959) Zygnemaceae. Indian Council of Agricultural Research, New Delhi.
Transeau EN (1951) The Zygnemataceae. Columbus Graduate School Monographs, Contributions in Botany, 1. Columbus, Ohio, 327 pp.
van Geel B (1976) Fossil spores of Zygnemataceae in ditches of a prehistoric settlement in Hoogkarspel (The Netherlands). Review of Palaeobotany and Palynology 22: 337–344.
van Geel B (1979) Preliminary report on the history of Zygnemataceae and the use of their spores as ecological markers. Proc. IV int. palynol. conf. Lucknow (1976-1977) 1: 467–469.
van Geel B (2001) Non-pollen palynomorphs. In: Smol JP, Birks HJB, Last WM (Eds.) Tracking environmental change using lake sediments. Vol. 3: Terrestrial, Algal, and Siliceous indicators. Dordrecht: Kluwer Academic Publishers, pp. 1-17.
van Geel B, van der Hammen T (1978) Zygnemataceae in Quaternary Colombian sediments. Review of Palaeobotany and Palynology 25: 377–392.
van Geel B, Coope GR, van der Hammen T (1989) Palaeoecology and stratigraphy of the Lateglacial type section at Usselo (The Netherlands). Review of Palaeobotany and Palynology 60: 25–129.
van Geel B, Mur LR, Ralska-Jasiewiczowa M, Goslar T (1994) Fossil akinetes of Aphanizomenon and Anabaena as indicators for medieval phosphate-eutrophication of Lake Gościąż (Central Poland). Review of Palaeobotany and Palynology 83: 97–105.
van Geel B, Grenfell HR (1996) Spores of Zygnemataceae. In J. Jansonius, D.C. McGregor (Eds.), Palynology: principles and applications. Vol. 1: Principles. Dallas: American Association of Stratigraphic Palynologists Foundation, pp. 173–179.

ANIMALIA

Helminths

Helminth eggs are often found in sediments, indicating presence of parasitic worms, their host and the disease in the area. In palynological studies, especially very resistant eggs such as of Trichuris (HdV-531), Ascaris (Brinkkemper and van Haaster 2012), Capillaria (Shumilovskikh et al. 2016b), Dicrocoelium (Shumilovskikh et al. 2016a) occur. While harsh palynological laboratory treatments might change the morphology of the eggs, identification of resistant helminth eggs is still possible (e.g. Lardín and Pacheco 2015). Studies on helminths eggs from sediments and latrines has developed to a special field of palaeoparasitology, where gentle methods of sample preparation are used.

References:
Brinkkemper O, van Haaster H (2012) Eggs of intestinal parasites whipworm (Trichuris) and mawworm (Ascaris): Non-pollen palynomorphs in archaeological samples. Review of Palaeobotany and Palynology 186: 16-21.
Lardín C, Pacheco S (2015) Helminths. Handbook for identification and counting of parasitic helminths eggs in urban wastewater. London: IWA Publishing.
Shumilovskikh LS, Hopper K, Djamali M, Ponel P, Demory F, Rostek F, Tachikawa K, Bittmann F, Golyeva A, Guibal F, Talon B, Wang L-C, Nezamabadi M, Bard E, Lahijani H, Nokandeh J, Omrani Rekavandi H, de Beaulieu J-L, Sauer E, Andrieu-Ponel V (2016a) Landscape evolution and agro-sylvo-pastoral activities on the Gorgan Plain (NE Iran) in the last 6000 years. The Holocene 26 (10): 1676-1691.
Shumilovskikh, LS, Seeliger M, Feuser S, Novenko E, Schlütz F, Pint A, Pirson F, Brückner H (2016b) The harbour of Elaia: A palynological archive for human environmental interactions during the last 7500 years. Quaternary Science Reviews 149: 167-187.

Testate amoebae

Testate amoebae are a group of eukaryotic microorganisms with a decay-resistant shell, which can get fossilized. While studies on testate amoebae are a subject of specialized analysis (Charman et al. 2000; Mitchell et al. 2008), the shells can be identified from pollen slides and used for reconstructions of hydrological changes, eutrophication or pollution (Grospietsch 1952; Mazei and Tsyganov 2006; Payne et al. 2012; Swindles and Roe 2007). Several NPP types were assigned to testate amoeba such as Amphitrema flavum (HdV-31A), A. wrightianum (HdV 31B, HdV 186, t 22), Assulina muscorum (HdV 32A) and A. seminulum (HdV-32B).

References:
Charman DJ, Hendon D, Woodland WA, 2000. The identification of testate amoebae (Protozoa: Rhizopoda) in peats. QRA Technical guide no. 9, Quaternary Research Association, London, 147 p.
Grospietsch T (1952) Die Rhizopodenanalyse als Hilfsmittel der Moorforschung. Die Naturwissenschaften 14: 318-323.
Mazei YA, Tsyganov AN (2006) Presnovodnye rakovinnye ameby (Freshwater Testate Amoebae) . Moscow: KMK Sci. Press.
Mitchell EAD, Charman DJ, Warner BG (2008) Testate amoebae analysis in ecological and paleoecological studies in wetlands: past, present and future. Biodiversity and Conservation 17: 2115-2137.
Payne RJ, Lamentowicz M, van der Knaap WO, van Leeuwen JFN, Mitchell EAD, Mazei Yu (2012) Testate amoebae in pollen slides. Review of Palaeobotany and Palynology 173: 68-79.
Swindles GT, Roe HM (2007) Examining the dissolution characteristics of testate amoebae (Protozoa: Rhizopoda) in low pH conditions: Implications for peatland palaeoclimate studies. Palaeogeography, Palaeoclimatology, Palaeoecology 252: 486-496.

BRYOPHYTA

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FUNGI

Fungal remains often occur in the pollen samples in form of thick-walled normally dark-coloured spores or dark-coloured hyphae. While hyphae cannot be identified morphologically, the variety of the fungal spores have distinct morphological characteristics. Fossil fungi are known since the Proterozoic (Taylor et al. 2015) and they are actively studied in geology (e.g. Elsik 1976, Graham 1962, Taylor and Osborn 1996). A special system has been developed for the description of fossil fungal remains (Elsik 1983) summarized by Kalgutkar and Jansonius (2000), presented in the on-line database of fossil fungi: https://advance.science.sfu.ca/fungi/fossils/Kalgutkar_and_Jansonius/.
While working with deep-time geology it is difficult to establish the connection of fossil finds to the modern mycoflora, but this is possible to do with Quaternary material. Taxonomically the majority of fungal spores recorded in palaeoecological studies belong to Ascomycetes, Basidiomycetes and Hyphomycetes (van Geel 2001), in rare cases to Zygomycetes (e.g. zygospores of Mucor sp. described by Shumilovskikh et al. 2015a). However, for interpretation purposes they are usually grouped by their nutritional strategies rather than by taxonomy: (1) saprotrophs, general or specialized on dung (coprophilous fungi), wood (lignicolous fungi), charred material (carbonicolous fungi) or soil, (2) parasites such as rusts and smuts, and 3) mycorrhizal fungi (Webster and Weber 2007).

References:
Elsik WC (1976) Microscopic fungal remains and cenozoic palynostratigraphy. Geoscience and Man 15: 115-120.
Elsik WC (1983) Annotated glossary of fungal palynomorphs. AASP Contribution Series 11: 1-35.
Graham A (1962) The role of fungal spores in palynology. Journal of Paleontology 36: 60-68.
Kalgutkar RM, Jansonius J (2000) Synopsis of fossil fungal spores, mycelia and frutifications. American Association of Stratigraphic Palynologists Foundation Contribution Series 39: 1-423.
Shumilovskikh LS, Schlütz F, Achterberg I, Bauerochse A, Leuschner HH (2015a) Non-pollen palynomorphs from mid-Holocene peat of the raised bog Borsteler Moor (Lower Saxony, Germany). Studia Quaternaria 32: 5-18.
Taylor TN, Osborn JM (1996) The importance of fungi in shaping the paleoecosystem. Review of Palaeobotany and Palynology 90: 249-262.
Taylor TN, Krings M, Taylor EL (2015) Fossil fungi. Amsterdam: Elsevier.
Van Geel B (2001) Non-pollen palynomorphs. In: Smol JP, Birks HJB, Last WM (Eds.) Tracking environmental change using lake sediments. Vol. 3: Terrestrial, Algal, and Siliceous indicators. Dordrecht: Kluwer Academic Publishers, pp. 1-17.
Webster J, Weber RWS (2007) Introduction to fungi. Cambridge: Cambridge University Press.