Linking Seasonal Dietary Strategies and Selectivity to Inform Forage Restoration for Przewalski’s Gazelle on the Qinghai–Tibet Plateau
Simple Summary
Abstract
1. Introduction
2. Materials and Methods
2.1. Study Area
2.2. Vegetation Monitoring and Fecal Sample Collection
2.3. Fecal Microhistological Analysis
2.4. Calculation of Vegetation Importance Values (IV)
2.5. Diversity and Niche Breadth Indices
2.6. Dietary Selectivity Index
2.7. Statistical Analysis
3. Results
3.1. Seasonal Variation in Dietary Functional Groups
3.2. Seasonal Variation in Dietary Diversity and Niche Breadth
3.3. Vegetation Composition and Availability
3.4. Dietary Selectivity and Preference Patterns
3.5. Seasonal Priority Forage Taxa Across the Basin
4. Discussion
4.1. Regional-Scale Patterns of Foraging Strategies
4.2. Core Foundation Taxa and Season-Specific Priority Resources
4.3. Implications for Restoration and Management
5. Conclusions
Supplementary Materials
Author Contributions
Funding
Institutional Review Board Statement
Informed Consent Statement
Data Availability Statement
Acknowledgments
Conflicts of Interest
References
- Atwood, T.B.; Valentine, S.A.; Hammill, E.; McCauley, D.J.; Madin, E.M.P.; Beard, K.H.; Pearse, W.D. Herbivores at the Highest Risk of Extinction among Mammals, Birds, and Reptiles. Sci. Adv. 2020, 6, eabb8458. [Google Scholar] [CrossRef]
- Pringle, R.M.; Abraham, J.O.; Anderson, T.M.; Coverdale, T.C.; Davies, A.B.; Dutton, C.L.; Gaylard, A.; Goheen, J.R.; Holdo, R.M.; Hutchinson, M.C.; et al. Impacts of Large Herbivores on Terrestrial Ecosystems. Curr. Biol. 2023, 33, 584–610. [Google Scholar] [CrossRef]
- Villar, N.; Medici, E.P. Large Wild Herbivores Slow down the Rapid Decline of Plant Diversity in a Tropical Forest Biodiversity Hotspot. J. Appl. Ecol. 2021, 58, 2361–2370. [Google Scholar] [CrossRef]
- Ripple, W.J.; Newsome, T.M.; Wolf, C.; Dirzo, R.; Everatt, K.T.; Galetti, M.; Hayward, M.W.; Kerley, G.I.H.; Levi, T.; Lindsey, P.A.; et al. Collapse of the World’s Largest Herbivores. Sci. Adv. 2015, 1, e1400103. [Google Scholar] [CrossRef]
- Yang, G.; Peng, C.; Chen, H.; Dong, F.; Wu, N.; Yang, Y.; Zhang, Y.; Zhu, D.; He, Y.; Shi, S.; et al. Qinghai–Tibetan Plateau Peatland Sustainable Utilization under Anthropogenic Disturbances and Climate Change. Ecosyst. Health Sustain. 2017, 3, e01263. [Google Scholar] [CrossRef]
- Liang, D.; Liu, Y.; Ping, X.; Jiang, Z.; Li, C. Ensuring Recovery for the Przewalski’s Gazelle. Science 2021, 374, 163. [Google Scholar] [CrossRef]
- Liu, T.; Jiang, Z.; Wang, W.; Wang, G.; Song, X.; Xu, A.; Li, C. Changes in Habitat Suitability and Population Size of the Endangered Przewalski’s Gazelle. Glob. Ecol. Conserv. 2023, 43, e02465. [Google Scholar] [CrossRef]
- Liang, D.; Li, C. Habitat Suitability, Distribution Modelling and GAP Analysis of Przewalski’s Gazelle Conservation. Animals 2024, 14, 149. [Google Scholar] [CrossRef] [PubMed]
- Liu, Z.; Wang, X.; Li, Z.; Cui, D.; Li, X. Feeding Habitats of Blue Sheep (Pseudois nayaur) during Winter and Spring in Helan Mountains, China. Front. Biol. China 2007, 2, 100–107. [Google Scholar] [CrossRef]
- Zweifel-Schielly, B.; Leuenberger, Y.; Kreuzer, M.; Suter, W. A Herbivore’s Food Landscape: Seasonal Dynamics and Nutritional Implications of Diet Selection by a Red Deer Population in Contrasting Alpine Habitats. J. Zool. 2012, 286, 68–80. [Google Scholar] [CrossRef]
- Lu, Q.; Hu, X.; Fan, L.; Jia, Y.; Gu, J.; Tang, H.; Lin, Y.; Liu, N. Construction of the Planning System for Qinghai Lake National Park. Nat. Park 2024, 2, 514–523. [Google Scholar] [CrossRef]
- Staver, A.C.; Hempson, G.P. Seasonal Dietary Changes Increase the Abundances of Savanna Herbivore Species. Sci. Adv. 2020, 6, eabd2848. [Google Scholar] [CrossRef]
- Castellaro, G.; Orellana, C.L.; Escanilla, J.P. Summer Diet of Horses (Equus ferus caballus Linn.), Guanacos (Lama guanicoe Müller), and European Brown Hares (Lepus europaeus Pallas) in the High Andean Range of the Coquimbo Region, Chile. Animals 2021, 11, 1313. [Google Scholar] [CrossRef]
- Zhang, N.; Zhang, Z.; Liu, C.; Xiong, Z.; Wei, Y.; Meng, D.; Zhan, M.; Li, Z.; Zhao, Y.; Teng, L.; et al. Seasonal Diet Composition of Goitered Gazelle (Gazella subgutturosa) in an Arid and Semi-Arid Region of Western China. Animals 2024, 14, 663. [Google Scholar] [CrossRef]
- Castellaro, G.G.; Squella, F.N.; Ullrich, T.R.; León, F.C.; Raggi, A.S. Algunas Técnicas Microhistolóigcas Utilizadas En La Determinación de La Composición Botánica de Dietas de Herbívoros. Agric. Téc. 2007, 67, 86–93. [Google Scholar] [CrossRef]
- Li, Z.; Yao, Z.; Zhang, M.; Khattak, R.H.; Han, X.; Sun, J.; Li, Z.; Lang, J.; Chen, C.; Jin, J.; et al. The Dietary Patterns of Water Deer Recently Rediscovered in Northeast China Exhibit Remarkable Similarities to Those Observed in Other Regions. Sci. Rep. 2025, 15, 9351. [Google Scholar] [CrossRef] [PubMed]
- Jacobs, J. Quantitative Measurement of Food Selection: A Modification of the Forage Ratio and Ivlev’s Electivity Index. Oecologia 1974, 14, 413–417. [Google Scholar] [CrossRef]
- Nyamukanza, C.C.; Sebata, A. Effect of Leaf Type on Browse Selection by Free-Ranging Goats in a Southern African Savanna. PLoS ONE 2020, 15, e0242231. [Google Scholar] [CrossRef]
- Pápay, G.; Fehér, Á.; Kiss, O.; Szabó, G.; Zimmermann, Z.; Hufnagel, L.; Járdi, I.; Szemethy, L.; Penksza, K.; Katona, K. Impact of Shrub Cover and Wild Ungulate Browsing on the Vegetation of Restored Mountain Hay Meadows. Tuexenia 2020, 40, 445. [Google Scholar]
- Li, Z.; Jiang, Z.; Li, C. Dietary Overlap of Przewalski’s Gazelle, Tibetan Gazelle, and Tibetan Sheep on the Qinghai-Tibet Plateau. J. Wildl. Manag. 2008, 72, 944–948. [Google Scholar] [CrossRef]
- Liu, B.; Jiang, Z. Dietary Overlap Between Przewalski’s Gazelle and Domestic Sheep in the Qinghai Lake Region and Implications for Rangeland Management. J. Wildl. Manag. 2004, 68, 241–246. [Google Scholar] [CrossRef]
- Gong, J.; Li, J.; Yang, J.; Li, S.; Tang, W. Land Use and Land Cover Change in the Qinghai Lake Region of the Tibetan Plateau and Its Impact on Ecosystem Services. Int. J. Environ. Res. Public Health 2017, 14, 818. [Google Scholar] [CrossRef]
- Qian, D.; Du, Y.; Li, Q.; Guo, X.; Cao, G. Alpine Grassland Management Based on Ecosystem Service Relationships on the Southern Slopes of the Qilian Mountains, China. J. Ethnobiol. Ethnomed. 2021, 288, 112447. [Google Scholar] [CrossRef] [PubMed]
- Li, C.; Li, Y.; Li, X.; Ma, L.; Xiao, Y.; Zhang, C. Differential Responses of Plant Primary Productivity to Nutrient Addition in Natural and Restored Alpine Grasslands in the Qinghai Lake Basin. Front. Plant Sci. 2021, 12, 792123. [Google Scholar] [CrossRef]
- Yang, Y.; Dou, Y.; An, S.; Zhu, Z. Abiotic and Biotic Factors Modulate Plant Biomass and Root/Shoot (R/S) Ratios in Grassland on the Loess Plateau, China. Sci. Total Environ. 2018, 636, 621–631. [Google Scholar] [CrossRef]
- Shen, M.; Wang, S.; Jiang, N.; Sun, J.; Cao, R.; Ling, X.; Fang, B.; Zhang, L.; Zhang, L.; Xu, X.; et al. Plant Phenology Changes and Drivers on the Qinghai–Tibetan Plateau. Nat. Rev. Earth Environ. 2022, 3, 633–651. [Google Scholar] [CrossRef]
- Owen-Smith, N. Functional Heterogeneity in Resources within Landscapes and Herbivore Population Dynamics. Landscape Ecol. 2004, 19, 761–771. [Google Scholar] [CrossRef]
- Williams, O.B. An Improved Technique for Identification of Plant Fragments in Herbivore Feces. J. Range Manag. 1969, 22, 51–52. [Google Scholar] [CrossRef][Green Version]
- Holechek, J.; Vavra, M. The Effect of Slide and Frequency Observation Numbers on the Precision of Microhistological Analysis. J. Range Manag. 2006, 34. [Google Scholar] [CrossRef]
- Aryal, A.; Brunton, D.; Ji, W.; Raubenheimer, D. Blue Sheep in the Annapurna Conservation Area, Nepal: Habitat Use, Population Biomass and Their Contribution to the Carrying Capacity of Snow Leopards. Integr. Zool. 2014, 9, 34–45. [Google Scholar] [CrossRef] [PubMed]
- Aebischer, N.J.; Robertson, P.A.; Kenward, R.E. Compositional Analysis of Habitat Use from Animal Radio-Tracking Data. Ecology 1993, 74, 1313–1325. [Google Scholar] [CrossRef]
- Lechowicz, M.J. The Sampling Characteristics of Electivity Indices. Oecologia 1982, 52, 22–30. [Google Scholar] [CrossRef] [PubMed]
- Hofmann, R.R. Evolutionary Steps of Ecophysiological Adaptation and Diversification of Ruminants: A Comparative View of Their Digestive System. Oecologia 1989, 78, 443–457. [Google Scholar] [CrossRef]
- Parker, K.L.; Barboza, P.S.; Gillingham, M.P. Nutrition Integrates Environmental Responses of Ungulates. Funct. Ecol. 2009, 23, 57–69. [Google Scholar] [CrossRef]
- Aikens, E.O.; Mysterud, A.; Merkle, J.A.; Cagnacci, F.; Rivrud, I.M.; Hebblewhite, M.; Hurley, M.A.; Peters, W.; Bergen, S.; De Groeve, J.; et al. Wave-like Patterns of Plant Phenology Determine Ungulate Movement Tactics. Curr. Biol. 2020, 30, 3444–3449.e4. [Google Scholar] [CrossRef]
- Merkle, J.A.; Monteith, K.L.; Aikens, E.O.; Hayes, M.M.; Hersey, K.R.; Middleton, A.D.; Oates, B.A.; Sawyer, H.; Scurlock, B.M.; Kauffman, M.J. Large Herbivores Surf Waves of Green-up during Spring. Proc. R. Soc. B Biol. Sci. 2016, 283, 20160456. [Google Scholar] [CrossRef]
- Long, R.J.; Apori, S.O.; Castro, F.B.; Ørskov, E.R. Feed Value of Native Forages of the Tibetan Plateau of China. Anim. Feed Sci. Technol. 1999, 80, 101–113. [Google Scholar] [CrossRef]
- Van Beest, F.M.; Mysterud, A.; Loe, L.E.; Milner, J.M. Forage Quantity, Quality and Depletion as Scale-Dependent Mechanisms Driving Habitat Selection of a Large Browsing Herbivore. J. Anim. Ecol. 2010, 79, 910–922. [Google Scholar] [CrossRef]
- Villalba, J.J.; Ramsey, R.D.; Athanasiadou, S. Review: Herbivory and the Power of Phytochemical Diversity on Animal Health. Animal 2025, 19, 101287. [Google Scholar] [CrossRef] [PubMed]
- Felton, A.M.; Felton, A.; Raubenheimer, D.; Simpson, S.J.; Krizsan, S.J.; Hedwall, P.-O.; Stolter, C. The Nutritional Balancing Act of a Large Herbivore: An Experiment with Captive Moose (Alces alces L). PLoS One 2016, 11, e0150870. [Google Scholar] [CrossRef] [PubMed]
- French, K.E. Species Composition Determines Forage Quality and Medicinal Value of High Diversity Grasslands in Lowland England. Agric. Ecosyst. Environ. 2017, 241, 193–204. [Google Scholar] [CrossRef]
- Foley, W.J.; Moore, B.D. Plant Secondary Metabolites and Vertebrate Herbivores—From Physiological Regulation to Ecosystem Function. Curr. Opin. Plant Biol. 2005, 8, 430–435. [Google Scholar] [CrossRef]
- Guo, L.; Zhao, H.; Zhai, X.; Wang, K.; Liu, L.; Wang, K.; Huang, D. Study on Life Histroy Traits of Stellera Chamaejasme Provide Insights into Its Control on Degraded Typical Steppe. J. Environ. Manag. 2021, 291, 112716. [Google Scholar] [CrossRef] [PubMed]
- Hector, A.; Bagchi, R. Biodiversity and Ecosystem Multifunctionality. Nature 2007, 448, 188–190. [Google Scholar] [CrossRef]
- Wagg, C.; Roscher, C.; Weigelt, A.; Vogel, A.; Ebeling, A.; De Luca, E.; Roeder, A.; Kleinspehn, C.; Temperton, V.M.; Meyer, S.T.; et al. Biodiversity–Stability Relationships Strengthen over Time in a Long-Term Grassland Experiment. Nat. Commun. 2022, 13, 7752. [Google Scholar] [CrossRef]
- Guo, C.; Ding, X.; Addi, Y.; Zhang, Y.; Zhang, X.; Zhuang, H.; Wang, Y. An Ethnobotany Survey of Wild Plants Used by the Tibetan People of the Yadong River Valley, Tibet, China. J. Ethnobiol. Ethnomed. 2022, 18, 28. [Google Scholar] [CrossRef]
- Pandey, S.; Chataut, G.; Maharjan, S.; Maidali, D.R.; Bhattarai, K. Unveiling the Potential of Jimbu (Allium przewalskianum): Bioactive Compounds, Antioxidant, and Antimicrobial Properties of a Native Himalayan Spice Herb. Int. J. Appl. Sci. Biotechnol. 2023, 11, 171–180. [Google Scholar] [CrossRef]
- Tang, S.; Ren, J.; Kong, L.; Yan, G.; Liu, C.; Han, Y.; Sun, H.; Wang, X.-J. Ephedrae Herba: A Review of Its Phytochemistry, Pharmacology, Clinical Application, and Alkaloid Toxicity. Molecules 2023, 28, 663. [Google Scholar] [CrossRef]
- Wei, J.; Li, S.; Su, T.; Zhao, J.; Jiang, Y.; Zubarev, Y.A.; Bi, Y. Phenolic Compositions and Antioxidant Activities of Hippophae tibetana and H. rhamnoides ssp. sinensis Berries Produced in Qinghai-Tibet Plateau. Food Chem. X 2022, 15, 100397. [Google Scholar] [CrossRef] [PubMed]
- Shang, Z.; Long, R. Formation Causes and Recovery of the “Black Soil Type” Degraded Alpine Grassland in Qinghai-Tibetan Plateau. Front. Agric. China 2007, 1, 197–202. [Google Scholar] [CrossRef]
- Li, W.; Zhao, R. Mechanisms Driving the Impact of Wolf Poison (Stellera Chamaejasme) in Grasslands of China. Weed Sci. 2025, 73, e45. [Google Scholar] [CrossRef]




| Family | Species | HN | HS | GN | GS | QF | TL | SI | HD | WY |
|---|---|---|---|---|---|---|---|---|---|---|
| Poaceae | Agropyron cristatum | 0.078 | 0.079 | 0.052 | 0.061 | 0.107 | 0.069 | 0.040 | 0.070 | 0.069 |
| Poa pratensis | 0.133 | 0.062 | 0.125 | 0.080 | 0.179 | 0.087 | 0.025 | 0.003 | 0.009 | |
| Agropyron desertorum | 0.001 | 0.044 | 0.029 | 0.083 | 0.010 | 0.032 | 0.101 | 0.005 | 0.035 | |
| Neotrinia splendens | 0.017 | 0.041 | 0.077 | 0.014 | 0.013 | 0.046 | 0.007 | 0.015 | 0.016 | |
| Stipa purpurea | 0.002 | 0.023 | 0.004 | 0.013 | 0.021 | 0.016 | 0.032 | 0.012 | 0.086 | |
| Elymus nutans | 0.040 | 0.074 | - | 0.003 | 0.032 | - | - | - | - | |
| Orinus kokonorica | - | - | - | 0.006 | - | - | 0.250 | 0.311 | 0.110 | |
| Cyperaceae | Carex arcatica | 0.033 | 0.048 | 0.122 | 0.088 | 0.051 | 0.153 | 0.015 | 0.029 | 0.212 |
| Kobresia humilis | 0.056 | 0.010 | 0.060 | 0.040 | 0.009 | 0.043 | 0.012 | 0.039 | 0.097 | |
| Asteraceae | Aster altaicus | 0.068 | 0.026 | 0.013 | 0.049 | 0.107 | 0.003 | 0.034 | 0.051 | 0.029 |
| Artemisia frigida | 0.038 | 0.034 | 0.069 | 0.095 | 0.037 | 0.090 | 0.057 | 0.076 | 0.032 | |
| Artemisia waltonii | 0.016 | 0.048 | 0.014 | 0.002 | 0.031 | 0.043 | 0.014 | 0.026 | 0.005 | |
| Artemisia salsoloides | - | - | - | - | - | - | 0.026 | 0.104 | - | |
| Fabaceae | Astragalus polycladus | 0.041 | 0.022 | 0.069 | 0.030 | 0.047 | 0.051 | 0.012 | 0.021 | 0.005 |
| Rosaceae | Argentina anserina | 0.033 | 0.060 | - | 0.003 | 0.013 | - | 0.002 | - | - |
| Sibbaldianthe adpressa | 0.013 | 0.008 | 0.040 | 0.075 | 0.009 | 0.015 | 0.002 | 0.012 | 0.093 | |
| Sibbaldianthe bifurca | 0.012 | 0.042 | 0.050 | 0.057 | 0.016 | 0.023 | 0.003 | 0.012 | 0.033 | |
| Potentilla multifida | 0.041 | 0.022 | 0.042 | 0.015 | 0.029 | 0.017 | - | - | 0.022 | |
| Allium przewalskianum | 0.010 | 0.025 | 0.021 | 0.046 | 0.020 | 0.003 | 0.066 | - | 0.029 | |
| Others | Lagotis brachystachya | 0.011 | 0.048 | 0.001 | 0.014 | 0.009 | 0.008 | - | 0.002 | - |
| Androsace mariae | 0.023 | 0.046 | 0.033 | - | 0.030 | - | - | - | - | |
| Salsola collina Pall | - | - | - | 0.006 | - | - | 0.102 | 0.013 | - | |
| Stellera chamaejasme | 0.019 | 0.001 | 0.008 | 0.001 | 0.001 | 0.030 | - | 0.033 | 0.007 | |
| Ephedra monosperma | - | - | - | - | - | - | - | 0.018 | 0.003 | |
| Hippophae tibetana | 0.014 | 0.007 | - | - | 0.011 | - | - | - | - |
| Plant | April | July | Categories | ||||
|---|---|---|---|---|---|---|---|
| n | Median D | Rank | n | Median D | Rank | ||
| Agropyron cristatum | 9 | 0.4892 | 1 | 9 | 0.262965 | 4 | Core |
| Aster altaicus | 9 | 0.4646 | 2 | 9 | −0.04698 | 11 | Core |
| Artemisia frigida | 9 | 0.3898 | 3 | 9 | 0.418654 | 1 | Core |
| Kobresia humilis | 9 | 0.291 | 4 | 9 | −0.15035 | 15 | Spring-specific |
| Leymus secalinus | 4 | 0.5905 | 5 | 4 | 0.657147 | 8 | Core |
| Thermopsis lanceolata | 6 | 0.25965 | 6 | 7 | 0.334988 | 2 | Summer-specific |
| Allium przewalskianum | 7 | 0.1141 | 7 | 7 | 0.270824 | 5 | Summer-specific |
| Stellera chamaejasme | 4 | 0.48395 | 8 | 1 | −0.75267 | 26 | Indicator species |
| Poa pratensis | 7 | 0.1619 | 9 | 6 | 0.217588 | 10 | Core |
| Astragalus polycladus | 9 | −0.1173 | 10 | 9 | 0.313621 | 3 | Summer-specific |
| Neotrinia splendens | 5 | 0.3868 | 11 | 4 | −0.13899 | 21 | Spring-specific |
| Agropyron desertorum | 4 | 0.4433 | 12 | 6 | 0.086579 | 12 | Spring-specific |
| Sibbaldianthe bifurca | 6 | 0.21325 | 13 | 8 | 0.124433 | 9 | Summer-specific |
| Taraxacum mongolicum | 4 | 0.163 | 14 | 5 | 0.704576 | 6 | Summer-specific |
| Hippophae tibetana | 2 | 0.7281 | 15 | 3 | 0.72438 | 14 | Local |
| Oxytropis stracheyana | 5 | 0.0592 | 16 | 7 | 0.064692 | 13 | Summer-specific |
| Potentilla multifida | 7 | −0.1521 | 18 | 7 | 0.353171 | 7 | Summer-specific |
Disclaimer/Publisher’s Note: The statements, opinions and data contained in all publications are solely those of the individual author(s) and contributor(s) and not of MDPI and/or the editor(s). MDPI and/or the editor(s) disclaim responsibility for any injury to people or property resulting from any ideas, methods, instructions or products referred to in the content. |
© 2026 by the authors. Licensee MDPI, Basel, Switzerland. This article is an open access article distributed under the terms and conditions of the Creative Commons Attribution (CC BY) license.
Share and Cite
Hou, L.; Xu, M. Linking Seasonal Dietary Strategies and Selectivity to Inform Forage Restoration for Przewalski’s Gazelle on the Qinghai–Tibet Plateau. Animals 2026, 16, 794. https://doi.org/10.3390/ani16050794
Hou L, Xu M. Linking Seasonal Dietary Strategies and Selectivity to Inform Forage Restoration for Przewalski’s Gazelle on the Qinghai–Tibet Plateau. Animals. 2026; 16(5):794. https://doi.org/10.3390/ani16050794
Chicago/Turabian StyleHou, Lili, and Ming Xu. 2026. "Linking Seasonal Dietary Strategies and Selectivity to Inform Forage Restoration for Przewalski’s Gazelle on the Qinghai–Tibet Plateau" Animals 16, no. 5: 794. https://doi.org/10.3390/ani16050794
APA StyleHou, L., & Xu, M. (2026). Linking Seasonal Dietary Strategies and Selectivity to Inform Forage Restoration for Przewalski’s Gazelle on the Qinghai–Tibet Plateau. Animals, 16(5), 794. https://doi.org/10.3390/ani16050794

