Pastoral Intensification and Peatland Drying in the Northern Tianshan Since 1560: Evidence from Fungal Spore Indicators

Abstract

Reconstructing historical grazing intensity is essential for understanding long-term human–environment interactions in arid and semi-arid regions. However, historical documents often lack continuous, site-specific information on land use and grazing pressure. We present a high-resolution reconstruction of pastoral activity and hydrological evolution since 1560 AD using fungal spore assemblages from a 92 cm lacustrine-peat sequence from the Sichanghu (SCH) peatland on the northern slope of the Tianshan Mountains, Central Asia. Quantitative analysis of coprophilous fungal spores and principal component analysis (PCA) of spore influxes identify three distinct phases of pastoral intensity: gradual intensification from 1560 to 1730 AD, a sharp decline from 1730 to 1770 AD, and rapid intensification from 1770 AD to the present. These transitions are consistent with historical records of land use and human migration in Xinjiang. Additionally, fungal assemblages reveal a long-term drying trend at Sichanghu, broadly consistent with regional aridification in northwestern China. However, centennial-scale discrepancies in humidity between local and regional records—particularly during the late Little Ice Age—indicate that local hydrological responses were strongly influenced by anthropogenic disturbances. This study highlights the value of fungal spores, particularly influx-based interpretations, as robust indicators of both human activities and hydroclimatic variability. It also underscores the importance of integrating local and regional signals when reconstructing past environmental changes in sensitive dryland ecosystems.

1. Introduction

Reconstructing the history of human activities is fundamental to understanding long-term human–environment interactions, adaptive strategies, and sustainable resource management. Xinjiang is an ecologically sensitive region and a key historical corridor along the ancient Silk Road, offering a unique setting for investigating past socio-environmental dynamics [1].

Human occupation in Xinjiang dates back to the Neolithic period, with evidence of agricultural and pastoral activities emerging approximately 3000 years ago [2]. Since then, the region has experienced cycles of sociopolitical transformation [3,4], leaving behind important archeological sites such as Gaochang (Han to Ming Dynasty) [5,6], Shirenziggou (Bronze to Iron Age) [7,8,9], Loulan (Han to Jin periods) [10], Niya (Wei and Jin periods) [11], Beiting (Tang Dynasty) [12], and Kalen (Qing Dynasty) [13]. The decline of sites like Loulan has sparked debate, with proposed causes ranging from overgrazing to environmental degradation [14,15].

Recent paleoecological studies have provided critical insights into past land use strategies. At Shirenziggou, pollen and fungal spore evidence indicates a mixed economy combining barley cultivation with pastoralism in the northern Tianshan Mountains [7]. Historical documents also describe major demographic shifts during the 18th century, when climate-induced droughts and floods prompted large-scale migrations from Shaanxi and Gansu into Xinjiang [1].

Despite this progress, our understanding of human activities in Xinjiang remains hindered by fragmented historical records and a scarcity of continuous, well-dated environmental archives. This has led to uncertainty regarding the timing and intensity of pastoral land use. Nevertheless, integrating documentary sources from the Ming and Qing dynasties [3,4,13] with sedimentary records from peatlands and lakes [16,17] offers valuable opportunities to reconstruct long-term human–environment interactions. Since the mid-20th century, Xinjiang has undergone rapid population growth—exceeding the national average—with marked regional disparities. The western and northern oasis regions are more densely populated, while the east and southern areas remain sparsely inhabited [18]. These demographic dynamics underscore the urgency of developing sustainable management strategies in light of ongoing and future climate challenges.

Fungal spores, particularly those from coprophilous taxa, are increasingly recognized as robust indicators of grazing pressure [19,20,21,22,23]. Genera such as Sordaria, Sporormiella, and Podospora are closely linked to herbivore presence, and their abundance has been used to infer livestock activity in paleoecological records. For instance, a marked increase in such spores at Qinghai Lake by around 3.5 ka coincides with archeological evidence of the onset of grazing [24,25]. Moreover, modern ecological studies reveal that fungal spore production is sensitive to environmental conditions. Certain taxa tend to be more abundant in peatlands than in lacustrine settings [26], while others are favored by nutrient-rich environments [27].

In this study, we analyze a lacustrine-peat sediment core from the northern slope of the Tianshan Mountains in Xinjiang. By examining fungal spore assemblages, we aim to reconstruct grazing activity and hydrological changes since the Little Ice Age. Integrating these findings with regional vegetation and climate records will help clarify the complex interplay between human activities and environmental change in this ecologically fragile region, providing scientific support for future land use and conservation planning. This work also contributes methodologically to the broader application of fungal indicators in long-term assessments of grazing intensity and hydrological dynamics.

2. Study Area

The Sichanghu peatland is situated approximately 20 km north of Jimsar County on the northern slope of the Tianshan Mountains, adjacent to the southeastern margin of the Gurbantunggut Desert (Figure 1). The peatland primarily develops from snowmelt runoff originating in the Tianshan Mountains and is underlain by ancient lacustrine sediments. It spans an area of approximately 20 km² [28]. This is a sedge-dominated peatland located within a shallow depression, with slight surface salinization and partial coverage by dunes in its northern section.

The regional climate is influenced by both the Westerlies and the East Asian Winter Monsoon [29], resulting in a cold, arid environment with strongly seasonal precipitation [17]. The mean annual precipitation is 129 mm, with summer and winter totals of approximately 72 mm and 14 mm, respectively. The mean annual temperature is 5.3 °C, with seasonal extremes of 21 °C in summer and −16 °C in winter [17].

Vegetation varies notably with elevation. Spruce (Picea) forests and alpine meadows dominate elevations above 1300 m, while desert steppe vegetation—comprising Artemisia, Chenopodiaceae, Ephedraceae, Asteraceae, and Poaceae—characterizes the lower slopes and surrounding plains. Within the peatland, vegetation is sparse and primarily consists of Cyperaceae and other hydrophytic herbs [30,31], which provide limited but important forage for grazing. The peat matrix is primarily derived from Cyperaceae remains [17] (Table S1). In recent years, intensified grazing has led to reduced herbaceous cover and increased localized desertification.

The southern part of the study area holds considerable historical significance. It lies near the ancient city of Beiting (Figure 1D), a key administrative center during the Tang Dynasty as part of the Anxi Protectorate (circa 700–1400 AD) [12]. Numerous archeological sites from different historical periods have been identified in the surrounding region [2,32], underscoring its long-standing role in human settlement and land use. This makes Sichanghu a valuable site for exploring the interplay between environmental change and human activity over time.

3. Material and Methods

A 92 cm sediment profile (44°06′29.2″ N, 89°12′07.5″ E, 616 ± 3 m a.s.l.) was collected from the center of the Sichanghu peatland. The profile was sampled at 1 cm intervals. The upper 52 cm consists of well-developed peat, underlain by dark gray silty clay extending to the bottom of the profile. Five accelerator mass spectrometry (AMS) radiocarbon dates on herbaceous macrofossils were used to establish the age–depth model [17]. The chronology spans the past ~450 years (from ca. 1560 AD to the present), with an average sediment accumulation rate of ~0.2 mm yr⁻¹, providing a robust temporal framework for paleoenvironmental reconstruction (Table S1, Figure S1). To facilitate understanding of the sedimentary context and chronological framework, a summary table (

Table 1. Summary of stratigraphy, chronology, and pollen zonation of the SCH Profile [17].

Depth (cm)	Stratigraphic Unit	14C Age (cal yr AD)	Pollen Zones
25–0	Peat	1880–2011	SCH-4: high contents of Chenopodiaceae, Cyperaceae, and Artemisia
56–25		1730–1880	SCH-3: more Typha and Poaceae
84–56	Silt clay	1600–1730	SCH-2: abundant Chenopodiaceae, Cyperaceae, and Artemisia
92–84		1561–1600	SCH-1: high percentages of Chenopodiaceae

) is provided, integrating depth intervals, stratigraphic units, calibrated ¹⁴C ages, and corresponding pollen zones.

Fungal spores were extracted using standard palynological techniques [33]. Approximately 1 mL of wet sediment per sample was processed, with Lycopodium spores added to calculate spore influxes. Samples underwent sequential treatment with 10% HCl (to remove carbonates), NaOH (to remove humic substances), and HF (to dissolve silicates). Cellulose was removed via acetolysis (9:1 acetic anhydride and concentrated sulfuric acid), followed by sieving through a 10 μm mesh and centrifugation to concentrate residues.

Spores were identified under light microscopy of ZEISS (Shanghai, China) following established taxonomic references and morphological criteria, including external morphology, size, symmetry, number of septa, pigmentation, wall thickness, and the presence of apical transparency [20,21,22,23,26,27,34,35,36,37,38,39,40,41,42,43,44,45]. Unidentified morphotypes were labeled “HY-number” (HY for Hengyang) and documented in detail in the Appendix. Fungal types designated as Type- or UAB- follow established classifications from published sources and are characterized by clear morphological and ecological attributes (Table S2). For each sample, a minimum of 300 spores were counted, with relative abundances calculated as percentages of the total spore sum. Zonation of fungal spore assemblages was conducted using constrained hierarchical clustering (CONISS) [46]. To identify ecological gradients and explore variation in fungal assemblages, principal component analysis (PCA) was performed using the ade4 package in R software (R-4.3.3) [47]. PCA was applied to both percentage data and influx data of fungal spores.

4. Results

4.1. Fungal Assemblages

A total of 25 fungal spore taxa were identified (Figure 2), with an average of 380 spores counted per sample. Dominant taxa included Type-200 (~27.7%), UAB-35 (~33.4%), Arnium (~6.2%), Sordaria (~7.7%), Coniochaeta ligniaria (Type-172) (~3.9%), Podospora (~2.8%), Type-201 (~4.4%), Glomus (~3.7%), and HY-1 (~2.5%). The mean fungal spore influx was 418 grains cm⁻² yr⁻¹, with pronounced temporal fluctuations. Based on CONISS and visual inspection, the assemblages were divided into four main zones (or subzones) (Figure 3 and Figure 4).

Zone 1 (92–54 cm; 1560–1730 AD): This interval exhibits consistent changes and was subdivided into Zones 1A and 1B based on shifts in the abundance of Type-200, Type-201, Glomus, HY-3, and Podospora.

Zone 1A (92–80 cm; 1560–1600 AD): Dominated by UAB-35 (~60.1%, ~400.8 grains cm⁻² yr⁻¹; all influx values below are expressed in grains cm⁻² yr⁻¹ unless otherwise noted), Sordaria (~14.8%, ~94.3 grains), Arnium (~5.2%, ~25.3 grains), Coniochaeta ligniaria (~2.8%, ~27.1 grains), Gelasinospora (~2.8%, ~21.4 grains), Podospora (~2.4%, ~10.9 grains), and HY-3 (~4.2%, ~13.3 grains). Delitschia was also present. Notably, Type-200, Type-201, Type-324, Alternaria, HY-1, and HY-2 were absent. Coprophilous spores accounted for ~29.7% of the assemblage but exhibited a relatively low influx (~187.1 grains).

Zone 1B (80–54 cm; 1600–1730 AD): Marked increases in Type-201 (~6.3%, ~58 grains), Glomus (~8.1%, ~75.7 grains), and Type-200 (~10.6%, ~10.4 grains), alongside the disappearance of HY-3. UAB-35 (~48.8%, ~460 grains), Arnium (~5.7%, ~54.6 grains), Sordaria (~10.9%, ~109.6 grains), Coniochaeta ligniaria (~2.7%, ~27 grains), and Gelasinospora (~1.9%, ~17.4 grains) remained abundant. Sporormiella, Valsaria, and Gaeumannomyces appeared sporadically. Although the relative proportion of coprophilous spores declined (~23.4%), their influx increased (~229.3 grains).

Zone 2 (54–17 cm; 1730–1920 AD): Characterized by a marked increase in Type-200 (~49.4%, ~1031.8 grains), Type-324 (~3.8%, ~76.1 grains), Podospora (~3.9%, ~103.4 grains), Type-10 (~1.6%, ~33.1 grains), and HY-2 (~0.8%, ~16 grains). Meanwhile, Glomus (~0.7%, ~10.9 grains), Gelasinospora (~0.1%, ~2.1 grains), and Delitschia (~0.1%, ~3 grains) decreased notably. Arnium (~4.7%, ~93.8 grains), Sordaria (~5.2%, ~95.8 grains), Coniochaeta ligniaria (~4.1%, ~78.1 grains), and HY-1 (~3.7%, ~65 grains) persisted. The proportion of coprophilous fungi was at its lowest (~18.1%), though their influx continued to rise (~376.4 grains).

Zone 3 (17–0 cm; 1920 AD–present): Exhibited elevated abundances and influxes of UAB-35 (~32.4%, ~1374.1 grains), Type-200 (~21.9%, ~2519.2 grains), Arnium (~11.5%, ~389.8 grains), Sordaria (~3.1%, ~99.5 grains), Coniochaeta ligniaria (~6.1%, ~373.1 grains), Glomus (~4.8%, ~213.2 grains), Type-201 (~3%, ~110.7 grains), HY-1 (~4.7%, ~156.6 grains), and HY-4 (~8.2%, ~993.2 grains). Meanwhile, Podospora (~3.3%, ~109.2 grains) and Type-324 (~0.1%, ~5.4 grains) declined. HY-2, HY-3, Sporormiella, and Valsaria were nearly absent, while Gaeumannomyces, Alternaria, Thecaphora, and Tripterospora appeared sporadically. Coprophilous fungal spore proportion rose to ~23.4%, with influx peaking at ~229.3 grains cm⁻² yr⁻¹.

4.2. Results of PCA

PCA of both fungal spore percentages and deposition influxes provided key insights into assemblage variability. The first two principal components explained 48.1% of the total variance for the percentage dataset (Axis 1: 33%, Axis 2: 15.1%) and 42.5% for the influx dataset (Axis 1: 27.5%, Axis 2: 15%).

PCA based on percentage data (Figure 5A) revealed that Axis 1 effectively distinguished samples from Zone 1 (lacustrine phase) and Zone 2 (peatland phase), whereas samples from Zone 3 (recent peatland) overlapped with both. Axis 2 showed limited discriminatory power, with substantial overlap among all three zones. Species composition varied among zones: UAB-35, Glomus, Sordaria, and Gelasinospora were dominant in Zone 1; Type-200, HY-1, Type-324, and Type-10 characterized Zone 2; Zone 3 exhibited a mixed assemblage, including high abundances in Arnium, Sordaria, Coniochaeta ligniaria, Podospora, Type-200, Type-201, Glomus, UAB-35, HY-1, and HY-4.

In contrast, PCA based on deposition influxes (Figure 5B) provided clearer zone separation. Axis 1 distinctly separated Zone 1 from Zones 2 and 3, while Axis 2 effectively distinguished between Zones 2 and 3. Species with high influxes in Zone 1 included Gelasinospora, Sordaria, Sporormiella, and HY-1. Zone 2 was characterized by Type-324, Type-10, HY-2, Type-201, and Valsaria. In Zone 3, dominant taxa included HY-4, Arnium, Gaeumannomyces, Delitschia, and Coniochaeta ligniaria. These patterns indicate that influx-based PCA is more effective in resolving ecological gradients and temporal shifts in fungal communities.

5. Discussion

5.1. Ecological and Environmental Significance of Fungal Spores

A comprehensive literature survey was conducted to interpret the ecological significance of the fungal taxa identified in the SCH profile (Table S2). Their morphological characteristics are documented in previous studies and illustrated with representative images in Figure 2.

Fungal spores serve as sensitive environmental proxies, particularly for reconstructing grazing pressure, and hydroclimatic variability. Coprophilous spores—such as Sordaria and Delitschia—were found in abundance in cultural layers at the Shirenzigou archeological site east of Sichanghu, but were scarce in natural deposits [7]. In both lacustrine and peatland sediments, fungal spores primarily originate from local processes, including the decomposition of nearby vegetation and dung deposition by herbivores. Consequently, both total fungal and coprophilous spore assemblages reflect hydrological conditions and grazing intensity at a local scale [24]. This pattern suggests a strong link between the presence of dung-associated fungi and herbivore activity, most likely driven by pastoralism in the region.

In addition to human influences, fungal communities also respond to local moisture conditions. For example, Gaeumannomyces and Trichocladium opacum are often observed in peat deposits following water table declines [21,35,40], while Type-200 spores tend to increase in sediments during drying phases [37]. These responses suggest that certain fungal taxa may serve as indirect indicators of hydrological conditions and moisture availability. Thus, spore assemblages provide valuable insights not only into grazing intensity but also into long-term shifts in local environmental moisture.

5.2. Pastoral Activities at Sichanghu Peatland Since 1560 AD

Multiple coprophilous fungal taxa—including Delitschia, Arnium, Sordaria, Coniochaeta ligniaria, Gelasinospora, Tripterospora, Podospora, and Sporormiella—were identified in the SCH profile, indicating a persistent presence of herbivorous animals in the region [7,24,25,48]. Their abundance and influx varied asynchronously through time (Figure 3 and Figure 4), suggesting the influence of both grazing intensity and environmental conditions. For example, Gelasinospora is associated with dung but sensitive to moisture variability [26]. In addition, variations in sedimentation rates associated with changes in sedimentary facies may have further contributed to these discrepancies.

To disentangle these signals, we performed PCA on both percentage and influx data. In the percentage-based PCA (Figure 5A), vectors for coprophilous fungi pointed in varying directions along both principal axes, mirroring the behavior of xerophilic taxa. This suggests that the percentage data do not capture a clear ecological gradient, likely due to differences in ecological tolerances among fungal types, rendering the pollen-style percentage approach less appropriate for these proxies.

In contrast, PCA based on spore influxes (Figure 5B) yielded more interpretable results. Coprophilous fungi were primarily aligned along the negative direction of Axis 2, while xerophilic fungi aligned along the negative direction of Axis 1. These findings suggest that Axis 2 reflects pastoral activity intensity and Axis 1 captures local aridification. Therefore, spore influx—rather than percentage—is a more robust indicator of herbivore abundance and environmental context [49,50].

The SCH profile reveals three distinct phases of pastoral activity (Figure 6C):

During 1560–1730 AD, PCA Axis 2 scores remained low but gradually decreased, while coprophilous fungal influx increased steadily, indicating slow intensification in grazing pressure. During 1730–1770 AD, a rapid increase in Axis 2 scores coincided with a marked decline in coprophilous influx, suggesting a temporary decline in grazing intensity. During 1770 AD–present, Axis 2 scores dropped sharply while coprophilous influx rose dramatically and fluctuated, reflecting a period of rapid and intensified pastoral expansion.

These phases align with regional historical records. For example, during the early Qianlong period (~1736 AD), migration and land cultivation policies were implemented in frontier regions, and Sichang Lake was designated as pastureland [2]. The subsequent establishment of a state-managed horse pasture in 1807 AD further intensified grazing, likely accounting for the sharp post-1770 AD spike in coprophilous fungal influx and drop in PCA-2 scores (Figure 6B,C). Similar trends have been observed at Lake Qinghai, where coprophilous spores display high-frequency fluctuations over the past five centuries [24,25], corroborating the widespread increase in pastoral activities across the region during and after the Little Ice Age.

Additionally, charcoal records from the SCH profile show high fire activity both prior to 1730 AD and after 1850 AD, suggesting anthropogenic disturbance consistent with periods of intensified grazing (Figure 6C,H). The alignment of fungal data, historical events, and charcoal concentrations reinforces the role of human activity—especially livestock management—as a primary driver of ecosystem changes at Sichanghu over the past four centuries.

5.3. Divergence and Implications of Local and Regional Environmental Evolution at Sichanghu Peatland Since 1560 AD

From 1560 to 1600 AD, the region experienced pronounced aridity, evidenced by coarse grain sizes, intensified aeolian activity (Figure 6J), and elevated charcoal concentrations indicative of frequent wildfires (Figure 6H). Vegetation was dominated by drought-adapted taxa such as Chenopodiaceae, Nitraria, Ephedra, and Asteraceae, with a low Artemisia/Chenopodiaceae (A/C) ratio (Figure 6I) [17]. Lithologically, this interval corresponds to lacustrine clay with minimal organic matter and sparse hygrophilous plant indicators (e.g., Cyperaceae, Typha) (Figure 6E–G). PCA Axis 1 scores were positive (Figure 6D), reflecting a lake-dominated environment with relatively high water levels but dry regional conditions that limited surrounding hydrophytic development.

From 1600 to 1730 AD, arid conditions persisted. Sediments remained coarse, and aeolian input and charcoal influxes continued to be high (Figure 6H,J). While drought-tolerant taxa remained dominant, the A/C ratio increased slightly, suggesting modest vegetation shifts (Figure 6I) [17]. Hygrophilous taxa and organic matter content increased marginally (Figure 6E–G), implying a slight contraction in lake extent and a transitional phase toward peatland formation. PCA-1 scores declined, indicating a modest increase in drought-tolerant fungi (Figure 6D).

From 1730 to 1840 AD, during the late Little Ice Age, regional humidity increased significantly. This is reflected by finer sediment grain sizes, reduced dust influx (Figure 6J), and lower charcoal concentrations (Figure 6H). Vegetation composition shifted, with reductions in drought-tolerant taxa and increases in Artemisia and Poaceae [17]. PCA-1 scores became negative, reflecting increased abundance in drought-tolerant fungi associated with semi-terrestrial conditions (Figure 6D). Lithological changes mark the onset of peat accumulation, with notable rises in organic matter and wetland plant abundance (Figure 6E–G), signaling a transition from a lacustrine to a marsh–meadow landscape.

Since 1840 AD, the region has undergone renewed and intensifying aridity. Grain sizes coarsened again, aeolian indicators and charcoal peaked (Figure 6H,J), and xerophytic taxa such as Chenopodiaceae, Asteraceae, and Ephedra reached high abundances [17]. Despite regional drying, wetland herbs (notably Typha and Cyperaceae) remained persistent or increased, suggesting a complex mosaic of drying peatlands with local water retention (Figure 6F,G). Organic content rose sharply, surpassing 40% in recent layers (Figure 6E). PCA-1 scores remained negative, with a slight increase in xerophilic fungal taxa during the modern era (Figure 6D).

A broader regional drying trend since 1645 AD is corroborated by multiple records [51], including pollen data from Lake Dongdaohaizi near Urumqi [52], rising dust storms around Lake Sugan [53], and fire history from the Altai Mountains [54]. These findings are broadly consistent with the SCH profile, which shows a lithological shift from lacustrine to peat-dominated environments. However, a discrepancy emerges during the late Little Ice Age (ca. 1730–1840 AD): while regional records such as carbonate declines in Lake Bosten [55], chironomid-based reconstructions from Lake Sugan [56], and dendrochronological evidence from the Tarim Basin [57] suggest a temporary moistening phase, the SCH profile exhibits high abundances in drought-tolerant fungal spores during this period (Figure 6D).

This divergence implies that fungal assemblages at Sichanghu respond more sensitively to local microenvironmental conditions than to broad regional climatic signals. For instance, Gaeumannomyces and Trichocladium opacum, which dominate this interval (ca. 1730–1840 AD), are commonly associated with decaying wood and persist in oxygenated but moist peat layers [21,35,40]. Their presence may reflect a localized shift from aquatic to semi-terrestrial conditions despite broader climatic amelioration. Additionally, intensified land use since the late 18th century may have exacerbated hydrological stress at the site. Historical documents report increased population migration from Jimusa’er County toward better-watered lowland areas during this time [2], possibly reducing water inflow into Sichanghu and amplifying peatland desiccation.

Therefore, while drought-tolerant fungi broadly track regional aridification since 1560 AD, their behavior during centennial-scale wet phases reveals the masking effects of anthropogenic landscape modification. These findings underscore the need for caution when applying fungal proxies in isolation for regional climate reconstructions and highlight their strength in capturing local ecological trajectories shaped by both climate and land use [58].

6. Conclusions

This study reconstructs the dynamics of pastoral activity and hydrological evolution at the Sichanghu peatland on the northern slope of the Tianshan Mountains since 1560 AD, using a high-resolution record of fungal spore assemblages. The key findings are as follows:

(1) Coprophilous fungal influxes serve as a reliable proxy for grazing intensity. Compared to percentage data, spore influx better reflects ecological gradients and herbivore abundance, particularly when interpreted through PCA.

(2) Three phases of pastoral activity were identified over the past ~460 years. Grazing pressure gradually intensified from 1560 to 1730 AD, declined between 1730 and 1770 AD, and increased rapidly from 1770 AD to the present. These shifts correspond well with historical records of land cultivation and pasture establishment.

(3) The Sichanghu peatland experienced a long-term drying trend consistent with regional aridification, but local responses diverged. Centennial-scale discrepancies—especially during the late Little Ice Age—suggest that fungal spores responded more sensitively to local hydrological conditions, which were heavily influenced by human settlement and water management. Thus, while fungal indicators align with regional climate trends over multi-centennial scales, they also capture localized ecological responses to anthropogenic pressures.

This research demonstrates the utility of fungal spores as high-resolution paleoenvironmental indicators and emphasizes the need to consider both regional climate signals and local human impacts when interpreting past environmental changes in arid landscapes.