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Coprophilous fungal spores are an emerging and potential proxies for indicating the long-term pastoral history. However, the indication of coprophilous fungal spores for dung and grazing intensity remains uncertain. In this study, in Horqin Sandy Land, 27 livestock dung samples and 33 surface soil samples were collected, and the indication of coprophilous fungal spores was discussed for the first time. The results showed that a total of 38 fungal spore types were recognized in the modern samples from the Horqin Sandy Land, including seven coprophilous fungal spore types. The coprophilous fungi assemblages vary in different dung types and may enable the diagnosis of specific livestock taxa. Coprophilous fungal spores, especially Sporormiella and Podospora are the most reliable dung indicators and can indicate the presence of herbivores. Moreover, some fungal spores, including Helminthosporium, Apiosordaria, HdV-104, and Glomus A were also potential indicators of grazing activities in the Horqin Sandy Land. On this basis, the concentration of coprophilous fungal spores may have the potential to be a quantitative indicator of grazing intensity. This study clarifies the indication of coprophilous fungal spores on dung and grazing intensity in the Horqin Sandy Land and provides a theoretical reference for further research on pastoral history in the region. [ABSTRACT FROM AUTHOR]

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The main aim of this study is to produce a modern analog for pollen and non-pollen palynomorphs (NPPs) preserved in soil in relation to the different vegetation types and herbivore impact in the Kaziranga National Park (KNP). The pollen data obtained reflects both the extant vegetation types in each habitat as well as landuse, but some site-by-site variation was observed with respect to the coprophilous fungal spores present in the palynoassemblage. Canonical correspondence analysis (CCA) analysis of pollen data reveals the presence of five significantly different vegetation types, while the non-pollen palynomorphs are relatively similar in relation to the different vegetation types. The long-term association of the wildlife and their impact on different vegetation types is one of the main reasons for the variation seen in the depositional pattern in the assemblage. The openland area is one of the most important areas for wildlife in the KNP as indicated by the presence of marker pollen and coprophilous fungal spores in the palynoassemblages. Coprophilous fungal spores were most abundant in this vegetation type reflecting the higher density of herbivores. The representation of pollen and coprophilous fungal spores from the swamp samples reflected the overall composition of all vegetation types existing in the KNP. This data can be utilized as a baseline for the interpretation of paleoecological and paleoherbivory studies in other parts of the Indian subcontinent as well as its potential application at a global level. [ABSTRACT FROM AUTHOR]

Upland moorlands are important landscapes, but many are considered degraded as a result of human activities. Consequently, their protection and restoration are of substantial concern. In Europe, restoration activities are often aimed at reversing the effects of 19th and 20th century "agricultural improvements," which often involved major drainage schemes. However, the ecological effects and long‐term ecological context of "agricultural improvement" are not yet fully understood. To develop this understanding, we analyze paleoecological data (pollen, coprophilous fungal spores, microcharcoal) from five upland peatland sites using a range of analytical approaches: cluster analysis, principal component analysis, rate‐of‐change analysis, and regression analyses incorporating documentary historical data. The sites are located on Exmoor (South West England, UK), a landscape that typifies historic upland degradation. We demonstrate that in this landscape, 19th century drainage is associated with declines in Sphagnum and non‐arboreal taxon richness; over longer timescales burning is associated with enhanced graminoid monocot abundance and grazing with lower taxon richness. We also show that rate‐of‐change in moorland vegetation communities during the 19th century is not distinctive in a long‐term context: change has been a constant in this landscape, rather than an exception during the 19th century. Our findings indicate that the aims of "restoration" interventions intended to increase Sphagnum abundances, increase taxon richness and reduce graminoid dominance are consistent with the long‐term dynamics of peatland systems, such as those on Exmoor. "Restoration" deemed successful in these terms may or may not resemble pre‐drainage conditions, which were themselves a function of millennia of successive moorland management regimes. [ABSTRACT FROM AUTHOR]

Abstract Upland moorlands are important landscapes, but many are considered degraded as a result of human activities. Consequently, their protection and restoration are of substantial concern. In Europe, restoration activities are often aimed at reversing the effects of 19th and 20th century “agricultural improvements,” which often involved major drainage schemes. However, the ecological effects and long‐term ecological context of “agricultural improvement” are not yet fully understood. To develop this understanding, we analyze paleoecological data (pollen, coprophilous fungal spores, microcharcoal) from five upland peatland sites using a range of analytical approaches: cluster analysis, principal component analysis, rate‐of‐change analysis, and regression analyses incorporating documentary historical data. The sites are located on Exmoor (South West England, UK), a landscape that typifies historic upland degradation. We demonstrate that in this landscape, 19th century drainage is associated with declines in Sphagnum and non‐arboreal taxon richness; over longer timescales burning is associated with enhanced graminoid monocot abundance and grazing with lower taxon richness. We also show that rate‐of‐change in moorland vegetation communities during the 19th century is not distinctive in a long‐term context: change has been a constant in this landscape, rather than an exception during the 19th century. Our findings indicate that the aims of “restoration” interventions intended to increase Sphagnum abundances, increase taxon richness and reduce graminoid dominance are consistent with the long‐term dynamics of peatland systems, such as those on Exmoor. “Restoration” deemed successful in these terms may or may not resemble pre‐drainage conditions, which were themselves a function of millennia of successive moorland management regimes.

The abundance of coprophilous (dung-inhabiting) fungal spores (CFS) in sedimentary records is an increasingly popular proxy for past megaherbivore abundance that is used to study megaherbivore-vegetation interactions, timing of megaherbivore population declines and extinctions, and the introduction of domesticated herbivores. This method often relies on counting CFS alongside pollen and tracers of known concentration such as exotic pollen or synthetic microspherules. Prior work has encouraged reporting CFS abundances as accumulation rates (spores/unit2/year) or concentration (spores/unit3) instead of percentages relative to the total pollen abundance, because CFS percentages can be sensitive to fluctuations in pollen influx. In this work, we quantify the uncertainty associated with estimating concentration values at different total counts and find that high uncertainty is associated with concentration estimates using low to moderate total counts (n = 20 to 200) of individual fungal spore types and tracers. We also demonstrate the effect of varying tracer proportions, and find that larger tracer proportions result in narrower confidence intervals. Finally, the probability of encountering a CFS spore from a specific taxon occurring in moderate concentrations (1,000 spores/unit2) dramatically decreases after a low tracer count (∼50). The uncertainties in concentration estimates caused by calculating tracer proportion are a likely cause of the high observed variance in many CFS time series, especially when CFS or tracer concentrations are low. Thus, we recommend future CFS studies increase counts and report the uncertainty surrounding concentration values. For some records, reporting spore data as presence/absence rather than concentrations or counts is preferable, such as when performing high counts is not feasible.

Pastoralism is considered a crucial factor in the eventual year-round occupation of high-altitude regions (>3000 m asl) of the Qinghai-Tibetan Plateau, and the northeastern Qinghai-Tibetan Plateau (NE-QTP) was an important corridor for early human occupation. We analyzed fossil fungal spore records for the last 8500 years from a high resolution aeolian section at Langgeri (LGR) on the NE-QTP Thirty-two fungal spore types were identified in the LGR section, including seven coprophilous types. We combined analysis of coprophilous fungal spores, Cyperaceae, Artemisia, and Hippophae pollen, and the charcoal >50 µm fraction to explore the timing and controls of pastoralism on the NE-QTP since the middle Holocene. Pastoralism commenced at LGR shortly before ca. 5.5 ka and gradually increased between ca. 5.5–3.5 ka, but markedly intensified after ca. 2.2 ka, with three periods of growth at ca. 2.1–1.9,1.4–1.2, and 0.6–0 ka, and a decline at ca. 1.1–0.6 ka. The timing of changes in pastoral activity on the NE-QTP based on the coprophilous fungal spore record is supported by regional archeology and historical documents. Technological and political developments, rather than climate change, played key roles in the long-term fluctuations of regional pastoralism on the NE-QTP in the late Holocene. [ABSTRACT FROM AUTHOR]

Analyses of high-resolution pollen data, coprophilous fungal spores, microscopic charcoal and sedimentology, combined with radiocarbon dating, allow the assessment of the impact of Sami and Nordic land use in the region surrounding the winter market town of Lycksele in northern Sweden. Such winter markets were established by the Crown during the seventeenth century AD to control the semi-nomadic movements of the Sami who traded here with Finnish settlers and were also taxed and educated. Little is known about Sami and Nordic co-existence beyond these market places, mainly due to a lack of archaeological evidence relating to Sami activity. Vegetation and land-use changes in the region between ~ AD 250 and 1825 reveal no signal for pre-seventeenth century agricultural activity, but the coprophilous fungal spore records suggest the increased regional presence of grazing herbivores (possibly reindeer) between ~ AD 800 and 1100. Sami activity in the parish of Lycksele has been suggested by rich metal finds dated to ~ AD 1000–1350 and they may have been attracted by an abundance of reindeer. [ABSTRACT FROM AUTHOR]

One major challenge in the study of late-Quaternary extinctions (LQEs) is providing better estimates of past megafauna abundance. To show how megaherbivore population size varied before and after the last extinctions in interior Alaska, we use both a database of radiocarbon-dated bone remains (spanning 25–0 ka) and spores of the obligate dung fungus, Sporormiella, recovered from radiocarbon-dated lake-sediment cores (spanning 17–0 ka). Bone fossils show that the last stage of LQEs in the region occurred at about 13 ka ago, but the number of megaherbivore bones remains high into the Holocene. Sporormiella abundance also remains high into the Holocene and does not decrease with major vegetation changes recorded by arboreal pollen percentages. At two sites, the interpretation of Sporormiella was enhanced by additional dung fungal spore types (e.g., Sordaria). In contrast to many sites where the last stage of LQEs is marked by a sharp decline in Sporormiella abundance, in interior Alaska our results indicate the continuance of megaherbivore abundance, albeit with a major taxonomic turnover (including Mammuthus and Equus extinction) from predominantly grazing to browsing dietary guilds. This new and robust evidence implies that regional LQEs were not systematically associated with crashes of overall megaherbivore abundance. [ABSTRACT FROM AUTHOR]

Fossil spores of coprophilous fungi preserved in lake and bog sediments are well-established palaeoecological tracers for the local history of pastoralism and other animal husbandry. However, applications in Africa are limited by the lack of information on the exact relationship between human activity and fungi growing on large-herbivore dung. Here we use 25 small crater-lake basins in western Uganda, which cover the complete regional gradient of natural vegetation and human impact intensity, as a natural laboratory to assess the relationship between coprophilous fungal spores recovered from recently deposited bottom sediments and the presence and abundance of large domestic herbivores in the surrounding crater basins. We find that the summed abundances of Sporormiella, Sordaria, Podospora and Delitschia, or of these four types plus Cercophora, are the most useful and robust fungal-spore indicators for the presence of domestic large herbivores in an East African context. Chaetomium could also be added to this list, particularly as it may help reduce stochasticity in the often low fossil counts of obligate coprophilous types. Our results suggest that domestic large herbivores, at least when associated with some degree of crop agriculture, leave a clearer signature on fungal-spore assemblages in the lakes' sediments than large wildlife. Furthermore, they indicate that the typically small number of livestock kept by mixed-subsistence farmers leave a stronger imprint on fungal-spore assemblages than the much larger herds of specialist pastoralists occasionally visiting the lake to drink or bathe. [ABSTRACT FROM AUTHOR]

The history of permanent human settlement in the high-altitude regions (>3000 m above sea level [masl]) of the Qinghai-Tibetan Plateau (QTP) is important in understanding human adaptation to this cold "Third Pole" region. The Qinghai Lake Basin was an important corridor used by prehistoric humans migrating to the inner QTP. Pastoralism is currently the most important means of sustaining permanent human settlement in the high-altitude regions of the QTP. However, the lack of reliable proxy measures reflecting prehistoric pastoral activities impedes our understanding of how pastoralism developed. The identification of coprophilous fungi in prehistoric cultural deposits may help refine the history of pastoralism. We collected 21 modern domesticated herbivore dung samples and 66 surface soil samples from the Qinghai Lake Basin for fungal spore analyses. We then evaluated how useful such analyses are for identifying grazing activities. Fifty-three samples were also collected from the JXG2 stratigraphic profile (∼10.0–0 ka; 3312 masl) for fungal spore analysis. Results indicate that low and stable values of the total concentration of coprophilous fungi were present from ∼10.0 to ∼5.5 ka. Concentrations gradually increased from ∼5.5 to ∼4.2 ka, significantly increased from ∼4.2 to ∼2.6 ka, and then increased dramatically after ∼2.6 ka. By combining these results with charcoal concentrations (>50 µm), lithic artifacts, bones and potsherds recovered from the JXG2 site, we infer that early pastoralism in Qinghai Lake Basin appeared between ∼6.0 to ∼5.5 ka and gradually intensified throughout the remainder of the Holocene as herding and farming gradually replaced hunting–gathering as the primary subsistence strategies. These results are supported by pollen records, archeological remains and historical records in the northeastern QTP. [ABSTRACT FROM AUTHOR]

The limited availability of historical and archaeological evidence means that much is still unknown about the development of Sami reindeer herding in Fennoscandia in both the recent and more distant past. To address this problem, high-resolution palynological analyses, C and Pb dating were undertaken on two adjacent (<25 m apart) peat profiles collected at a recently abandoned reindeer gathering pen ( renvall) near Jokkmokk (~66.6°N, 19.8°E) in the boreal forest of northern Sweden. The aim was to assess the impact of Sami reindeer herding on the local environment through a study of pollen, coprophilous fungal spores, microscopic charcoal and sedimentology. The samples collected from within an annex to the renvall indicate cycles of use and abandonment of the pen on a multi-decadal timescale between ~ ad 1800-2008, most obviously in the coprophilous fungal spore archive. The pattern and timing of these cycles confirm events previously known only from oral histories. Although the local pollen assemblage zones associated with the phasing of activity were reproducible in a second peat core beyond the boundary of the renvall, the coprophilous fungal spore signal in this paired profile was much less distinctive, possibly due to the typically shorter dispersal distances for these microfossils in comparison to pollen grains. [ABSTRACT FROM AUTHOR]

The type HdV-368 (van Geel et al., 1981) or the Podospora -type (van Geel et al., 2001) is a common representative of non-pollen palynomorphs (NPP) with high palaeoecological potential. Here we present a revision of the type on the basis of a survey of the mycological literature and incubation of related dung fungi. The characteristic truncated base of the type HdV-368 is shared by the ascospores of at least 31 species of the genus Podospora , including one species that probably belongs to the nearly related genus Schizothecium . We arrange these species in morphological spore taxa, which are named after representative species, and provide an identification key to the new types and groups including descriptions of ecology and distribution of each species. The former type of HdV-368 falls into the newly erected Podospora decipiens -type, which covers the spores of the coprophilous species P. decipiens and P. perplexens and partly of P. pleiospora and P. prethopodalis . A short discussion of the spores of coprophilous fungi as palaeoecological indicators for animal diversity, population density, vegetation, climate, local conditions, and transport events is also given. The key's application can provide detailed insights into past environmental and biodiversity changes. [ABSTRACT FROM AUTHOR]

Decadal-scale analyses of fungal spores in a lake-sediment core from Ware Pond, located in the town of Marblehead in northeastern Massachusetts, test the potential of this approach for reconstructing past sheep and cattle grazing in southern New England, USA. The influx of spores of Sordaria and other coprophilous taxa increases at ad 1650, which corresponds with the beginning of European settlement, and subsequent peaks in these taxa at ad 1840 coincide with maximum abundances of weedy and agricultural taxa in the pollen record. Historical data from Marblehead and neighbouring towns indicate that maximum numbers of cattle and sheep occurred at this time. These findings suggest that fungal spores in New England lake sediments can be used to reconstruct changes in grazing pressure over time at the landscape scale. [ABSTRACT FROM AUTHOR]

Megaherbivores are keystone species whose removal from landscapes can cause cascading ecosystem changes, yet the consequences of Late Quaternary megaherbivore extinctions remain uncertain. This paper tests the Megaherbivory Release Hypothesis (MRH), which posits that the decline and extinction of megaherbivores (body size >1000 kg) during the last deglaciation in eastern North America contributed to the expansion of more palatable hardwood tree taxa, the development of vegetation assemblages with no modern analogue, and increased fuel load and fire activity. Coprophilous fungal spores in lake sediment records are used as proxies for megaherbivore abundance and are essential to testing the MRH through analyses of lead/lag relationships among vegetation composition, megaherbivore abundance, fire, and climate. Although some prior analyses of coprophilous fungal spores from individual sites have supported the MRH, these interpretations have been complicated by 1) discrepancies in the timing of coprophilous spore declines versus megaherbivore extinction timing based on dated vertebrate remains, 2) reliance on a single fungal taxon (Sporormiella) rather than a full suite of coprophilous fungi taxa, and 3) uncertainties in the taphonomic processes that influence fungal spore abundances. To examine the spatiotemporal relationships among megaherbivory, vegetation, and fire, we developed five new multi-taxon coprophilous fungal spore records for comparison with existing pollen, spore, and charcoal records from 14 sites across eastern North America. The MRH was well supported in the northeast and central US, with most sites showing a coprophilous spore decline by ∼14.6 ka followed by a rise of hardwood taxa (∼14.4 ka). However, changes in fire regime varied widely among northeast and central US sites and may have preceded spore declines. The MRH was not well supported in the southeastern US, where a smaller rise in hardwood taxa (∼16.1–13.1 ka) generally preceded the decline in coprophilous spores at individual sites (∼15.8–12.7 ka). These site-level and regional differences suggest spatial variations in the strength of couplings among late-Quaternary megaherbivore extinctions, vegetation composition and structure, and fire regime. Possible explanations for the differences between the northern and southeastern US include (1) differences in landscape heterogeneity of canopy openness and palatability, (2) net primary productivity and sensitivity to top-down trophic effects, (3) megaherbivore density, and (4) climate trends and seasonality at orbital to millennial timescales. [ABSTRACT FROM AUTHOR]

To reconstruct the evolution of livestock in SW Greenland over the last two millennia, we measured the concentration of bile acids in a sedimentary sequence retrieved from Lake Igaliku. Deoxycholic acid (DOC) was the sole bile acid and was present throughout the sequence. The DOC flux correlated quantitatively with that of coprophilous fungal spores. Maximum DOC and coprophilous fungal spores flux was recorded during the two periods of human settlement and grazing activity in the region (i.e. the Norse settlement during the Middle Ages and the recent Danish agricultural phase since 1920). These flux values were consistent with the presence of recent livestock around the lake and are attested to by way of archeological data relating to the Norse period. In contrast, the DOC and coprophilous fungal spores background during pre-Norse times and the Little Ice Age (LIA), indicated the presence of wild herbivores. Lower DOC and coprophilous fungal spore flux values after the Norse abandonment, compared with pre-colonization conditions, could indicate that Norse activity in conjunction with climate change, altered the pristine wildlife. Therefore, these quantitative correlations between DOC flux and coprophilous fungal spores flux potentially suggest a quantitative relationship with the livestock grazing in the catchment. The comparison between sedimentary DOC and coprophilous fungal spores provides significant highlights on past pastoral dynamics over the last 2000 yr in SW Greenland. [ABSTRACT FROM AUTHOR]

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 Ø39 at Qinngua in the former Eastern Settlement). Landnám 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. [ABSTRACT FROM AUTHOR]

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 Pinus sylvestris-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. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]