Coprophilous Fungal Spores Research Articles

Combined pollen and macrofossil data as a source for reconstructing mosaic patterns of the early medieval urban habitats – a case study from Gdańsk, N. Poland

Plant macrofossils, pollen, fungal spores and other non-pollen palynomorphs (NPPs), were analyzed in three archaeological exposures established in a small area in the oldest part of the historic city of Gdańsk. One of the aims was to investigate how precisely the combined pollen and macrofossil data may reflect the diversity of local habitats within an urban archaeological site and to compare the range of information obtained using each of the methods separately. The other aim of the study was to reconstruct the environmental conditions in the earliest occupied area of the historic centre of Gdańsk. The repeatable composition and, in many points, consistency among pollen and macrofossil data in all three profiles strongly suggest that the apparently biased structure of these fossil remains in fact reflects a mosaic pattern of the vegetation in the area of the site. The combined pollen and macrofossil data enabled separation of local elements from those transported to the site by different means. The results show that already at the beginning of the 12th c., the area was almost totally deforested, but up to the mid-13th c. the settlement organization was still relatively loose, resembling a rather rural structure with well-developed ruderal plant communities and patches of semi-natural vegetation. At the coring area, strongly eutrophic riparian vegetation was bordering patches of dry grassland on a sandy holm. The large proportion of pollen of various entomophilous plants mostly from meadows and pastures is explained as a result of the presence of dung, as confirmed by the abundances of coprophilous fungal spores. The study indicates that parallel pollen and macrofossil analysis is essential for correct interpretation of fossil assemblages. Application of both methods enables broadening of the range of remains representing various organisms and obtaining more precise tools for environmental reconstruction and better understanding of the taphonomic processes at the archaeological site.

RECONSTRUCTING RECENT LAND DEGRADATION IN THE SEMI‐ARID KAROO OF SOUTH AFRICA: A PALAEOECOLOGICAL STUDY AT COMPASSBERG, EASTERN CAPE

ABSTRACTThis paper forms part of an ongoing, multidisciplinary study of land degradation in the Eastern Cape, South Africa. A series of studies of erosion, runoff and the development of gullies, badlands and sediment delivery to small farm reservoirs in the Sneeuberg uplands have shown that the semi‐arid Karoo is vulnerable to soil erosion. Cereal cultivation and increased livestock numbers are thought to have played a major role in triggering erosion over the last 200 years. To test this assumption, we analysed pollen, non‐pollen palynomorphs and microscopic charcoal from sediments that have infilled one farm dam since the 1930s and which is located in the Sneeuberg uplands of the Eastern Cape. The results, supported by sediment source tracing, indicate that soil erosion was closely associated with agricultural activity. The reconstructions appear to be in agreement with historical documentary records. However, records of livestock stocking densities were found to be insufficiently detailed to enable direct matching against microfossil proxies for grazing, especially the coprophilous fungal spore, Sporormiella type. This study demonstrates the importance of combining sediment tracing techniques with palynological analyses to provide a more informed interpretation of land use changes as recorded by lake and reservoir sediments. Copyright © 2012 John Wiley & Sons, Ltd.

114 Coprophilous fungal spores and selected non-pollen palynomorphs (NPPs) in lake and peat deposits from Greenland

114 Coprophilous fungal spores and selected non-pollen palynomorphs (NPPs) in lake and peat deposits from Greenland

Holocene history of Lac des Lauzons (2180 m a.s.l.), reconstructed from multiproxy analyses of Coleoptera, plant macroremains and pollen (Hautes-Alpes, France)

A 1.70 m core extracted from the Lac des Lauzons, Haut Champsaur, French Alps, at 2180 m altitude, provided a detailed Holocene record of beetles, pollen and plant macrofossils, enabling the reconstruction of local palaeoenvironmental changes during the last 10 000 years. After an early phase of colonization by plants and insects, corresponding to the Lateglacial interstadial, a long phase of relative stability of the ecosystems (at least in the vicinity of the lake) is recorded. Strikingly, there is no evidence from beetle and plant macrofossils that the treeline reached the altitude of Lauzons during the Holocene climate optimum, although this period is characterized by major forest expansion in many high-altitude sites in the southern French Alps. The uppermost part of the record is blurred by the infilling of the lake, progressively turning into a peat bog. This sequence also provides an opportunity to compare the records of coprophilous fungal spores and coprophilous beetles and to improve the interpretation of these proxies in terms of their significance as proxies for pastoralism.

Grazing impacts and woodland management in Eriksfjord: Betula, coprophilous fungi and the Norse settlement of Greenland

This paper focuses on the impact of Norse settlement on vegetation and landscape around the head of Tunulliarfik (Eriksfjord) in southern Greenland. Pollen, radiocarbon, microscopic charcoal and fungal spore data are presented from a peat monolith which was collected close to the ruins of a large Norse farm complex (group O39 at Qinngua in the former Eastern Settlement). Landnam is identified at ca. cal. a.d. 1020 by a small decrease in pollen from Betula, a slight increase in Poaceae, and the appearance of pollen from Norse apophytes (native plants favoured and spread by human activity) and anthropochores (not native and unintentionally introduced by people). Increases in microscopic charcoal and palynological richness are also apparent. This pattern is broadly consistent with that seen in other pollen diagrams from this region. The sequence is unusual for Greenland, however, in that relatively high Betula pollen percentages (average 20% TLP) are recorded throughout the period of settlement, up to the end of the 14th century a.d. before the profile becomes truncated. If these data are primarily representative of the dry land vegetation, then they suggest that birch woodland and scrub may well have persisted close to the farm throughout the Norse period. Given the potential resource value of woodland to the settlers, this may imply that birch was being managed sustainably, as was the case in Iceland during the medieval period. Coprophilous fungal spores typically found on animal dung are abundant during the early phase of settlement, yet subsequently decline in abundance. This may indicate a likely decrease in grazing intensity or livestock numbers over time, possibly in response to climatic deterioration and/or soil erosion that is expected to have placed increased stress on the pastoral farming system.

Dung fungi as indicators of past herbivore abundance, Kruger and Limpopo National Park

Effective wildlife management needs historical data on herbivore abundance and its interactions with vegetation, climate and disturbance over longer time periods that is available through observational and archive data. Spores specific to herbivore dung provide a potential source of information on past herbivore abundances. This paper sets out to evaluate the potential of fungal spores as environmental indicators and in particular the use of coprophilous fungi in understanding past herbivore densities and their impact on the savanna landscape of Kruger and Limpopo National Parks (South Africa and Mozambique). Spore assemblages from six sedimentary cores are analysed and compared with the pollen data. Spores of coprophilous fungi, Coniochaeta cf ligniaria, and Sordariaceae in particular provide a valuable source of information about past herbivore densities. The spore assemblages of investigated localities show historical fluctuations in herbivore abundance. Peaks in wild/domestic herbivore densities can be seen, on a local scale from 800 to 900 AD and another at 1400 AD, however, these cannot be linked with any significant changes in vegetation. The last 200–300 years have seen an increased abundance of herbivores in the Limpopo floodplain sites, particularly domestic cattle. There is no clear correspondence between changes in herbivore abundance and local vegetation in this period or the 20th century. However, domestic cattle, together with wild herbivores, probably contributed to creating a mosaic type of landscape with heterogeneous tree cover.

Late Holocene land-use and vegetation dynamics in an upland karst region based on pollen and coprophilous fungal spore analyses: an example from the Burren, western Ireland

The late Holocene environmental history of two karstic uplands in the Burren, western Ireland is reconstructed. The palaeoecological investigations focus on species-rich, upland plant communities of high biogeographic interest that include Sesleria-dominated grasslands and heath communities with Dryas octopetala, Arctostaphylos uva-ursi and Empetrum nigrum. Short monoliths taken from shallow peats were pollen analytically investigated. Particular attention was paid to non-pollen palynomorphs, and especially coprophilous fungal spores as indicators of environmental change and pastoral activity at local level. The exposed north-west Burren uplands carried Pinussylvestris-dominated woodland during the mid and late Bronze Age. The demise of pine on these uplands at ca. 600 B.C. is ascribed to human impact. Evidence is presented for increased pastoral activity in the uplands from early medieval times (ca. sixth century A.D.) onwards. Farming, involving intensive grazing of the uplands, attained greatest intensity during the late 18th and early 19th century, and resulted in more or less total clearance of Corylus scrub which, prior to that, was common in both the upland and lowland Burren. The potential of non-pollen palynomorphs and especially the coprophilous fungal spore record for elucidating traditional Burren farming practices, including winterage (a type of transhumance), is highlighted.

Fresh insights into long‐term changes in flora, vegetation, land use and soil erosion in the karstic environment of the Burren, western Ireland

Summary 1. The study focuses on species‐rich, upland, heathy vegetation with arctic‐alpine floristic affinities and Sesleria grasslands in the karstic Burren region, western Ireland. The investigations aimed at reconstructing the long‐term development of these high conservation‐value communities and the role of farming in their formation and long‐term survival. 2. The methods used included pollen analysis and 14C‐dating of short monoliths and investigation of grykes (fissures in karstic limestone) for evidence of soil erosion. Special attention was paid to fossil, coprophilous fungal spores as indicators of local grazing. The strong local character of the pollen records facilitated identification of inter‐site differences as well as regional patterns. It is shown that open pine woodland characterized the Cappanawalla uplands between c. 1500 BC and 500 BC. It is proposed that such woodlands, with floristic affinities to Scandinavian open pine woodlands on calcareous soils, provided a suitable environment for the present‐day, open heath vegetation with species such as Dryas octopetala, Arctostaphylos uva‐ursi, Geranium sanguineum and Empetrum nigrum. 3. Burning of vegetation as a management tool was important in the uplands over most of the last two millennia. Firing seems to have ceased with the onset of more intensive grazing regimes in the 18th century. 4. Synthesis. Upland palaeoarchives, derived from shallow peaty deposits, show that the upland Burren supported mainly plagioclimax Corylus‐dominated woody vegetation and grasslands from c. 1500 BC (beginning of present record), until possibly as late as the 17th century AD. In the uplands of the north‐western Burren, open, species‐rich pinewoods with hazel dominated. The northern‐arctic elements in the present‐day upland flora survived clearances, involving initially Pinus sylvestris (c. 500 BC) and subsequently Corylus avellana (c. AD 1600). Colluvial material retrieved from grykes supports the idea of considerable soil loss occurring as late as the first and early 2nd millennium AD. The investigations highlight the potential of upland palaeoarchives, consisting of short sequences, for elucidating vegetation and land‐use dynamics in karstic environments such as the Burren.

Spores of coprophilous fungi from under the Dawson tephra (25,300 14C years BP), Yukon Territory, northwestern Canada

Based on the analysis of pollen and macrofossils Zazula et al. [Zazula, G.D., Froesne, D.G., Elias, S.A., Kuzmina, S., la Farge, C., Reyes, A.V., Sanborn, P.T., Schweger, C.E., Smith, C.A.S., Mathewes, R.W., 2006. Vegetation buried under Dawson tephra (25,300 14C yr BP) and locally diverse late Pleistocene paleoenvironments of Goldbottom Creek, Yukon, Canada. Palaeogeography, Palaeoclimatology, Palaeoecology 242, 253–286.] reported an in situ riparian grassy meadow ecosystem, and regional well-drained steppe-tundra, buried by Dawson tephra (ca 25,300 BP) at Goldbottom Creek in the Canadian Yukon Territory. An additional study of fungal spores in pollen slides from that site was completed. Ascospores of the coprophilous Sordaria type, Sporormiella type and Podospora type, appeared to be of common occurrence, pointing to the former presence of herbivores. The analysis of the various spore types of coprophilous fungi in pollen slides is useful and may lead to further knowledge of the role of different herbivore taxa in past ecosystems.

The impact of tourism and reindeer herding on forest vegetation at Saariselkä, Finnish Lapland: a pollen analytical study of a high-resolution peat profile

A high-resolution peat profile from the vicinity of Saariselkä (68°25.24'N, 27°25.19'E), a large tourist centre in northern Finnish Lapland, was sampled continuously at near annual resolution to a depth of 22 cm in order to interpret the changes in pollen assemblages and vegetation resulting from human interference and reindeer herding. Fifteen AMS dates were wiggle-matched with the atmospheric 14C calibration curve. The resulting age—depth model allows the calculation of pollen accumulation rates (PAR grains/cm2 per yr), which, in turn, enables the fine-scale human impact to be interpreted on much better grounds than would be possible with the classical pollen percentages approach. Results indicate that increasing tourist numbers since the building of the Saariselkä resort in 1950 are not seen in the pollen diagrams. Only a slight, although clear, increase in the pollen taxa of disturbed sites (mainly Gramineae with some herb pollen taxa) can be detected from the PAR diagram after the 1950s, along with a decrease of Pinus pollen following the clearance of the ski slopes in the 1970s. Principal components analysis (PCA) of fossil pollen spectra and modern pollen analogues obtained from both natural and human-disturbed sites confirms that human interference that has affected the vegetation around Saariselkä is not visible in the pollen record to the degree that might have been expected. This is interpreted as resulting from the considerably different nature of human impact at Saariselkä, where the infrastructure is fully developed and weed species are efficiently removed. The remarkable growth of reindeer population in the area during the1980s, however, can be observed in the peat profile through an increase in the number of coprophilous fungal spores ( Podospora, Sporormiella). This increase in fungal spores occurs simultaneously with a decrease in Betula pollen, which is interpreted to be a consequence of reindeer foraging the young Betula trees and seedlings.

Development of modern analogues for natural, mowed and grazed grasslands using pollen assemblages and coprophilous fungi

Tens of thousands of hectares of salt marsh have formed along eastern Canada's macro-tidal Bay of Fundy. Most of these marshes have been diked and converted to agricultural land since the arrival of Acadian settlers in the 17th century. Although there are currently no controlled salt marsh restoration efforts underway in the Bay of Fundy, there are isolated examples of salt marshes that were restored as a result of unplanned dike breaches. In an effort to develop pollen modern analogues of local dikeland and salt marsh land uses, surface soil samples were collected from John Lusby Marsh, a dikeland to which tidal flooding was restored in the late 1940s, and two nearby diked marshes, one left fallow and one under active agricultural management. Using discriminant analysis (DA) and principal components analysis (PCA) we successfully separated pollen assemblages representing three grassland land use types: salt marsh, farmed dikeland and fallow dikeland. The farmed dikeland land use was further divided into grazed or manured land use and hayed or unmanured land use based on the presence and relative proportions of coprophilous fungal spores such as Sordaria-type, Tripterospora-type, Cercophora-type, Chaetomium-type, Podospora-type and Sporormiella-type. This work marks the first attempt at incorporating dung fungal spores into a palynological analysis using controlled conditions and demonstrates how the inclusion of dung fungal spore abundances can relay information on past land uses. This brings a new interpretive dimension to palynological analyses without requiring any additional sample collection or preparation.

Linking current environments and processes to fungal spore assemblages: Surface NPM data from woodland environments

While the basic premise of palaeoecological research is that the past and present can be linked, qualitatively or quantitatively, this has not been the case with much of the work in fungal spores and other non-pollen palynomorphs (NPPs). This relates partly to the shortage of ecological data, especially in terms of microfungi on plants or in soils. To address this problem, modern samples have been analysed from moss and from subsurface soil from Quercus woodland in SW England. This paper reports modern fungal and pollen assemblage data from grazed and ungrazed sites in order to test the use of fungal NPMs as indicators of grazing in prehistory. Preliminary conclusions, based here on a small number of samples, suggest that spores of coprophilous fungi are, as expected, more prevalent in soil samples from grazed woodland, but that a zero count does not mean absence of animals. Other fungal types also respond to grazing through indirect mechanisms, probably due to changes in the abundance of host plants and ground conditions.

Environmental history of the dry forest biome of Guerrero, Mexico, and human impact during the last c. 2700 years

Two lake sediment cores from Madre del Sur mountain range,Guerrero State, west-central Mexico were studied to examine the pastdynamics of the dry forest biome. Pollen, spores of coprophilousfungi, cyanobacteria and lithological changes are presented. The390-cm Tixtla core (1730'N, 99°24W, 1400 maltitude) represents the last 2700 cal. yr; the 340-cm Huitziltepeccore (17°45N, 99°28W, 1430 m altitude) representsthe last 2050 cal. yr. Pollen shows climate-dependent competitionbetween dry deciduous forest (with Asteraceae, Chamaesyce-type,Bursera, Euphorbia, Myrica and Lysiloma-Pithecellobium-type as main components) and moister(mesophyllous) forest (with Pinus, Quercus, Alnus and Juglans). Human impact on the environment is apparent duringthe entire record, as evidenced by crop cultivation (Zea mays,Amaranthus, Capsicum, Chenopodium), coprophilous fungi (Podospora-type and Sporormiella-type), deforestation (at Tixtla in particular) and lake eutrophication, ie, cyanobacteria blooms. Environmental change at both sites is summarized as follows. Period 1 (2700-2450 cal. yr BP): abundant dry deciduous forest and climate conditions drier than today. Period 2 (2450-1950 cal. yr BP): little change in proportions of dry deciduous and mesophyllous forest but floral composition of the forests differed between sites. Human occupation was low. Period 3 (1950-1550 cal. yr BP): dry deciduous forest decreased while mesophyllous forest increased, indicating more humid climatic conditions. Populations expanded significantly around 1950 cal. yr BP. Period 4 (1550-1070 cal. yr BP): maximum expansion of mesophyllous forest. Climate was more humid than today. Pre-Hispanic populations at Tixtla reached high numbers leading to deforestation and eutrophication. Period 5 (1070-c. 750 cal. yr BP): shows abundant mesophyllous forest and lowest proportions of dry deciduous forest. Climate continued to be humid. There is evidence of forest harvesting, crop cultivation and dense human population. Period 6 (c. 750-225 cal. yr BP): dry deciduous forest expanded and mesophyllous forest decreased. There was significant deforestation by pre-Hispanic people around Lake Tixtla. High population densities and frequent fires at both sites caused eutrophication of the lakes. Domesticated herbivores were introduced ∼ad 1500 and the presence of dung is reflected in the record of coprophilous fungi. Period 7 (the last 225 calendar years): the basins were used intensively by local people and their cattle. Dry deciduous forest continued to increase while mesophyllous forest decreased.

Interactions of climate and land use documented in the varved sediments of Seebergsee in the Swiss Alps

This paper presents a multiproxy high-resolution study of the past 2600 years for Seebergsee, a small Swiss lake with varved sediments at the present tree-line ecotone. The laminae were identified as varves by a numerical analysis of diatom counts in the thin-sections. The hypothesis of two diatom blooms per year was corroborated by the 210Pb and 137Cs chronology. A period of intensive pasturing during the ‘Little Ice Age’ between ad 1346 and ad 1595 is suggested by coprophilous fungal spores, as well as by pollen indicators of grazing, by the diatom-inferred total phosphorus, by geochemistry and by documentary data. The subsequent re-oligotrophication of the lake took about 88 years, as determined by the timelag between the decline of coprophile fungal spores and the restoration of pre-eutrophic nutrient conditions. According to previous studies of latewood densities from the same region, cold summers around ad 1600 limited the pasturing at this altitude. This demonstrated the socio-economic impact of a single climatic event. However, the variance partitioning between the effects of land use and climate, which was applied for the whole core, revealed that climate independent of land use and time explained only 1.32% of the diatom data, while land use independent of climate and time explained 15.7%. Clearly land use in‘ uenced the lake, but land use was not always driven by climate. Other factors beside climate, such as politics or the introduction of fertilizers in the seventeenth and eighteenth centuries also in‘ uenced the development of Alpine pasturing.

Palaeoecological perspectives on pattern and process in plant diversity and distribution adjustments: a comment on recent developments*

Abstract.Two developments in the understanding of the relationship between sedimentary pollen assemblages, vegetation and plant diversity are discussed. The Prentice model of vegetation–pollen relationships has improved our understanding of how and at what scale sedimentary pollen records vegetation. The modelling framework allows improved palaeoecological study designs that may, potentially, give important new insights into processes of plant migrations in response to climate change. Also, reconstructions of plant communities and landscape openness may be improved. Competing hypotheses for the relationship between vegetation and palynological diversity are discussed and it is concluded that more attention should be focused on evenness aspects of palynological diversity and on hypothesis testing.