Article

Alami, A. 2022: Biodiversity Loss in the Moroccan central High Atlas, its Impact on Local Ecosystems and National Economy , and Wildlife Conservation Strategy : Findings from 20 years of Research. – J. Anal. Sci. Appl. Biotechnol. 4: 81-96.

Alderotti, F. &Verdiani, E. 2023: God save the queen! How and why the dominant evergreen species of the Mediterranean Basin is declining? – AoB PLANTS 15: plad051.

Allen, C. & Breshears, D. D. 1998: Drought-induced shift of a forest-woodland ecotone: Rapid landscape response to climate variation. – Proc. Nat. Acad. Sci. USA 95: 14839-14842. https://doi.org/10.1073/pnas.95.25.14839

Allen, R. G., Pereira, L. S., Raes, D. & Smith, M. 1998: Crop evapotranspiration-Guidelines for computing crop water requirements-FAO Irrigation and drainage paper – Rome.

Anderegg, W. R., Hicke, J. A., Fisher, R. A., Allen, C. D., Aukema, J. & al. 2015: Tree mortality from drought, insects, and their interactions in a changing climate. – New Phytol. 208: 674-683. https://doi.org/10.1111/nph.13477

––, Kane, J. M. & Anderegg, L. D. L. 2013: Consequences of widespread tree mortality triggered by drought and temperature stress. – Nat. Clim. Change 3: 30-36. https://doi.org/10.1038/nclimate1635

Barbeta, A. & Peñuelas, J.. 2016: Sequence of plant responses to droughts of different timescales: lessons from holm oak (Quercus ilex) forests. – Pl. Ecol. Divers. 9: 321-338. https://doi.org/10.1080/17550874.2016.1212288

Batllori, E., Lloret, F., Aakala, T., Anderegg, W. R., Aynekulu, E. & al. 2020: Forest and woodland replacement patterns following drought-related mortality. – Proc. Natl. Acad. Sci. U.S.A. 117: 29720-29729. https://doi.org/10.1073/pnas.2002314117

Belko, N., Zaman‑Allah, M., Diop, N. N., Cisse, N., Zombre, G. & al. 2013: Restriction of transpiration rate under high vapour pressure deficit and non-limiting water conditions is important for terminal drought tolerance in cowpea. – Pl. Biol. 15: 304-316. https://doi.org/10.1111/j.1438-8677.2012.00642.x

Bell, B. A., Hughes, P. D., Fletcher, W. J., Cornelissen, H. L., Rhoujjati, A. & al. 2022: Climate of tthe Marrakech High Atlas, Morocco: Temperature lapse rates and precipitation gradient from piedmont to summits. – Arct. Antarct. Alp. Res. 54: 78-95. https://doi.org/10.1080/15230430.2022.2046897

Benabid, A. 1982: Bref aperçu sur la zonation altitudinale de la végétation climacique du Maroc. – Ecol. Medit. 8: 301-315.

Benabid, A. 1985: Les écosystèmes forestiers, préforestiers et presteppiques du Maroc : diversité, répartition biogéographique et problèmes posés par leur aménagement. – Forêt Médit. 7: 53-64.

Bennett, A. C., McDowell, N. G., Allen, C. D. & Anderson‑Teixeira, K. J. 2015: Larger trees suffer most during drought in forests worldwide. – Nat. Pl. 1: 1-5. https://doi.org/10.1038/nplants.2015.139

Braun-Blanquet, J. & Maire, R. 1921: Études sur La végétation et La flore marocaines: comptes-rendus des herborisations de la société botanique de France, session du Maroc, 1921. – Bull. Soc. Bot. Fr. 68: 5-7.

Caliskan, S. 2014: Germination and seedling growth of holm oak (Quercus ilex L.): effects of provenance, temperature, and radicle pruning. – iForest 7: 103-109. https://doi.org/10.3832/ifor0967-007

Camarero, J. J., Sánchez‑Salguero, R., Sangüesa‑Barreda, G., Lechuga, V., Viñegla, B. & al. 2021: Drought, axe and goats. More variable and synchronized growth forecasts worsening dieback in Moroccan Atlas cedar forests. – Sci. Total Environ. 765: 142752. https://doi.org/10.1016/j.scitotenv.2020.142752

Choat, B., Brodribb, T. J., Brodersen, C. R., Duursma, R. A., López, R. & Medlyn, B. E. 2018: Triggers of tree mortality under drought. – Nature 558: 531-539. https://doi.org/10.1038/s41586-018-0240-x

Da Costa, A. C. L., Galbraith, D., , Almeida, S., Portela, B. T. T., da Costa, M. & al. 2010: Effect of 7 yr of experimental drought on vegetation dynamics and biomass storage of an eastern Amazonian rainforest. – New Phytol. 187: 579-591. https://doi.org/10.1111/j.1469-8137.2010.03309.x

Daget, P. 1977: Le bioclimat méditerranéen : analyse des formes climatiques par le système d’Emberger. – Vegetatio 34: 87-103.

Dallahi, Y., Boujraf, A., Meliho, M. & Orlando C. A. 2023: Assessment of forest dieback on the Moroccan Central Plateau using spectral vegetation indices. – J. For. Res. 34: 793-808. https://doi.org/10.1007/s11676-022-01525-x

DeHayes, D. H., Jacobson, G. L., Schaberg, P. G., Bongarten, B., Iverson, L. & Dieffenbacher-Krall, A. C. 2000: Forest Responses to Changing Climate: Lessons from the Past and Uncertainty for the Future. – Pp. 495-540. In: Mickler, R.A., Birdsey, R.A., Hom, J. (eds), Responses of Northern U.S. Forests to Environmental Change, 139. – New York.

Dorado, F. J., Corcobado, T., Brandano, A., Abbas, Y., Alcaide, F. & al. 2022: First Report of Dieback of Quercus suber Trees Associated with Phytophthora quercina in Morocco. –Pl. Dis. 107: 1246. https://doi.org/10.1094/PDIS-08-22-1795-PDN

Dyer, J. M. 2009: Assessing topographic patterns in moisture use and stress using a water balance approach. – Landsc. Ecol. 24: 391-403. https://doi.org/10.1007/s10980-008-9316-6

Eliades, F., Sarris, D., Bachofer, F., Michaelides, S. & Hadjimitsis, D. 2024: Understanding Tree Mortality Patterns: A Comprehensive Review of Remote Sensing and Meteorological Ground-Based Studies. – Forests 15: 1357. https://doi.org/10.3390/f15081357

Fernández-de-Uña, L., Martínez-Vilalta, J., Poyatos, R., Mencuccini, M. & McDowell N. G. 2023: The role of height-driven constraints and compensations on tree vulnerability to drought. – New Phytol. 239: 2083-2098. https://doi.org/10.1111/nph.19130

Fischbein, D., & Corley, J. C. 2022: Population ecology and classical biological control of forest insect pests in a changing world. – For. Ecol. Manag. 520: 120400. https://doi.org/10.1016/j.foreco.2022.120400

García-Angulo, D., Hereş, A.-M., Fernández-López, M., Flores O., Sanz, M. J. & al. 2020: Holm oak decline and mortality exacerbates drought effects on soil biogeochemical cycling and soil microbial communities across a climatic gradient. – Soil Biol. Biochem. 149: 107921. https://doi.org/10.1016/j.soilbio.2020.107921

Gea-Izquierdo, G., Natalini, F., & Cardillo, E. 2021. Holm oak death is accelerated but not sudden and expresses drought legacies. – Sci. Total Environ. 754: 141793. https://doi.org/10.1016/j.scitotenv.2020.141793

Gharnit, Y., Moujane, A., Outourakhte, A., Hassan, I., El Amraoui, K. & al. 2025: Plant Richness, Species Assessment, and Ecology in the M’goun Geopark Rangelands, High Atlas Mountains, Morocco. – Rangel. Ecol. Manag. 98: 357-376. https://doi.org/10.1016/j.rama.2024.08.022

––, Outourakht, A., Boulli, A. & Hassib, A. 2023. Biodiversity, Autecology and Status of Aromatic and Medicinal Plants in Geopark M’Goun (Morocco). – Ann. Bot. 13: 39-54. https://doi.org/10.13133/2239-3129/18027

––, Outourakhte, A., Moujane, A., Ikhmerdi, H., Hasib, A., & Boulli, A. 2024: Habitat diversity, ecology, and change assessment in the geoparc M’goun in High Atlas Mountains of Morocco. – Geol. Ecol. Landsc.: 1-22. https://doi.org/10.1080/24749508.2024.2431354

Grossiord, C., Buckley, T. N., Cernusak, L. A., Novick, K. A., Poulter, B. & al. 2020: Plant responses to rising vapor pressure deficit. – New Phytol. 226: 1550-1566. https://doi.org/10.1111/nph.16485

Huang, S., Tang, L., Hupy, J. P., Wang, Y. & Shao, G. 2021: A commentary review on the use of normalized difference vegetation index (NDVI) in the era of popular remote sensing. –J. For. Res. 32: 1-6. https://doi.org/10.1007/s11676-020-01155-1

Jung, T., Blaschke, H., & Oßwald, W. 2000: Involvement of soilborne Phytophthora species in Central European oak decline and the effect of site factors on the disease. – Pl. Pathol. 49: 706-718. https://doi.org/10.1046/j.1365-3059.2000.00521.x

Jurskis, V. 2005: Eucalypt decline in Australia, and a general concept of tree decline and dieback. – For. Ecol. Manag. 215: 1-20. https://doi.org/10.1016/j.foreco.2005.04.026

Kassie, A. W., Moges, A., Alelign, T. & G. Bayeh. 2024. Economic Valuation of Ecosystem Services and Its Association with Socioeconomic Factors in the Wof-Washa Natural State Forest, North Shewa Zone, Ethiopia. – Scientifica 2024: 6607551. https://doi.org/10.1155/2024/6607551

Khatouri, M. 1992: Growth and yield of young Quercus ilex coppice stands in the Tafferte forest (Morocco). – Pp. 77-82 in: Romane, F., Terradas, J. (eds), Quercus ilex L. ecosystems: function, dynamics and management. – Dordrecht.

Khayi, S., Armitage, A. D., Gaboun, F., Meftah-Kadmiri, I. Lahlali, R. & al. 2023: Chromosome-scale assembly uncovers genomic compartmentation of Fusarium oxysporum f. sp. albedinis, the causal agent of Bayoud disease in date palm. – Front. Microbiol. 14: 1268051. https://doi.org/10.3389/fmicb.2023.1268051

Kholová, J., Nepolean, T., Tom Hash, C., Supriya, A., Rajaram, V.,  Senthilvel, S., & Kakkera, A. 2012: Water saving traits co-map with a major terminal drought tolerance quantitative trait locus in pearl millet [Pennisetum glaucum (L.) R. Br.]. – MMol. Breed. 30: 1337-1353. https://doi.org/10.1007/s11032-012-9720-0

Kichas, N. E., Hood, S. M., Pederson, G. T., Everett, R. G. & McWethy D. B. 2020: Whitebark pine (Pinus albicaulis) growth and defense in response to mountain pine beetle outbreaks. – For. Ecol. Manag. 457: 117736. https://doi.org/10.1016/j.foreco.2019.117736

Kodero, J. M., Felzer, B. S. & Shi, Y. 2024: Future transition from forests to shrublands and grasslands in the western United States is expected to reduce carbon storage. – Commun. Earth Environ. 5: 78. https://doi.org/10.1038/s43247-024-01253-6

Landsberg, J. & Wylie, F. R. 1983: Water stress, leaf nutrients and defoliation: a model of dieback of rural eucalypts. – Aust. J. Ecol. 8: 27-41. https://doi.org/10.1111/j.1442-9993.1983.tb01516.x

Lihui, M., Xiaoli, L., Jie, C., Youke, W. & Jingui, Y. 2021: Effects of Slope Aspect and Rainfall on Belowground Deep Fine Root Traits and Aboveground Tree Height. – Front. Pl. Sci. 12: 684468. https://doi.org/10.3389/fpls.2021.684468

Liñán, J., Cantos, M., Troncoso, J., García, J. L., Fernández, A. & Troncoso, A. 2011: Some propagation methods for cloning holm oak (Quercus ilex L.) plants. – Cent. Eur. J. Biol. 6: 359-364. https://doi.org/10.2478/s11535-011-0007-y

Linares, J. C. & Tíscar, P. A. 2010: Climate change impacts and vulnerability of the southern populations of Pinus nigra subsp. salzmannii. – Tree Physiol. 30: 795-806. https://doi.org/10.1093/treephys/tpq052

––, Taïqui L. & Camarero J. J. 2011: Increasing drought sensitivity and decline of Atlas cedar (Cedrus atlantica) in the Moroccan Middle Atlas forests. – Forests 2: 777-796. https://doi.org/10.3390/f2030777

––, ––, Sangüesa-Barreda G., Seco J. I. & Camarero J. J. 2013: Age-related drought sensitivity of Atlas cedar (Cedrus atlantica) in the Moroccan Middle Atlas forests. – Dendrochronologia 31: 88-96. https://doi.org/10.1016/j.dendro.2012.08.003

Lindenmayer, D. B. & Laurance, W. F. 2017. The ecology, distribution, conservation and management of large old trees. – Biol. Rev. 92: 1434-1458. https://doi.org/10.1111/brv.12290

Liyaqat, I., Balzano, A., Niccoli, F., Kabala, J. P., Merela, M. & Battipaglia, G. 2024: Xylogenesis Responses to a Mediterranean Climate in Holm Oak (Quercus ilex L.). –Forests 15: 1386. https://doi.org/10.3390/f15081386 

Magalhães, J. G. de S., Amoroso, M. M. & Larson B. C. 2021: What evidence exists on the effects of competition on trees’ responses to climate change? A systematic map protocol. – Environ. Evid. 10: 34. https://doi.org/10.1186/s13750-021-00249-5

McDowell, N. G., Sapes, G., Pivovaroff, A., Adams, H. D., Allen, C. D., Anderegg, W. R. & Xu, C. 2022: Mechanisms of woody-plant mortality under rising drought, CO2 and vapour pressure deficit. – Nat. Rev. Earth Environ. 3: 294-308. https://doi.org/10.1038/s43017-022-00272-1

Medina, S., Vicente R., Nieto-Taladriz, M. T., Aparicio N., Chairi F. & al. 2019: The Plant-Transpiration Response to Vapor Pressure Deficit (VPD) in Durum Wheat Is Associated With Differential Yield Performance and Specific Expression of Genes Involved in Primary Metabolism and Water Transport. – Front. Pl. Sci. 9: 1994. https://doi.org/10.3389/fpls.2018.01994

Meneses-Tovar, C. L. 2011: NDVI as indicator of degradation. – Unasylva 62: 39-46.

Michaud, H., Toumi, L., Lumaret, R., Li, T. X., Romane, F. & Di Giusto, F. 1995: Effect of geographical discontinuity on genetic variation in Quercus ilex L. (holm oak). Evidence from enzyme polymorphism. – Heredity 74: 590-606. https://doi.org/10.1038/hdy.1995.83

Molina, M. O., Sánchez, E. & Gutiérrez, C. 2020: Future heat waves over the Mediterranean from an Euro-CORDEX regional climate  model ensemble. – Sci. Rep. 10: 8801. https://doi.org/10.1038/s41598-020-65663-0

Montes-Vega, M. J., Guardiola-Albert, C. & Rodríguez-Rodríguez, M. 2023: Calculation of the SPI, SPEI, and GRDI Indices for Historical Climatic Data from Doñana National Park: Forecasting Climatic Series (2030–2059) Using Two Climatic Scenarios RCP 4.5 and RCP 8.5 by IPCC. – Water 15: 2369. https://doi.org/10.3390/w15132369

Moujane, A., Outourakhte, A., Gharnit Y., Boulli, A. & Ouhammou, A. 2023: Geographical distribution of the Thuriferous juniper (Juniperus thurifera L.) and the influencing factors in the Atlas Mountains of Azilal (Central High Atlas, Morocco). – Geology, Ecology, Landscapes 7: 1-12. https://doi.org/10.1080/24749508.2023.2203281

Mueller-Dombois, D. 1988: Forest decline and dieback : A global ecological problem. – Trends Ecol. Evol. 3: 310-312. https://doi.org/10.1016/0169-5347(88)90108-5

Natalini, F., Alejano, R., Vázquez-Piqué, J., Cañellas, I. & Gea-Izquierdo, G. 2016: The role of climate change in the widespread mortality of holm oak in open woodlands of Southwestern Spain. – Dendrochronologia 38: 51-60. https://doi.org/10.1016/j.dendro.2016.03.003

Niu, S., Luo, Y., Li, D., Cao, S., Xia, J., Li, J. & Smith, M. D. 2014: Plant growth and mortality under climatic extremes: An overview. – Environ. Exp. Bot. 98: 13-19. https://doi.org/10.1016/j.envexpbot.2013.10.004

Novák, V. 2011. Evapotranspiration. – Pp. 280-283 in: Gliński, J., Horabik, J., Lipiec, J. (eds), Encyclopedia of Agrophysics – Dordrecht.

Oliva, J., Boberg, J. B., Hopkins, A. J., & Stenlid, J. 2013: Concepts of epidemiology of forest diseases. – Pp. 1-28 in Gonthier, P. & Nicolotti, G. (eds), Infectious forest diseases, 1. – Wallingford.

Ouchbani, S. & Romane, F. 1995: Gradient climatique et répartition de la végétation dans l’Atlas de Béni Mellal (Maroc). – Bull. Inst. Sci. 19: 53-64.

Outourakhte, A., Gharnit, Y., Moujane, A., El Haddany, K., Hasib, A. & Boulli, A. 2025: The Floristic Composition and Phytoecological Characterization of Plant Communities in the M’Goun Geopark, High Atlas, Morocco. – Ecologies 6(2): 29. https://doi.org/10.3390/ecologies6020029

Panaïotis, C., Carcaillet, C. & M’Hamedi, M. 1997: Determination of the natural mortality age of an holm oak (Quercus ilex L.) stand in Corsica (Mediterranean Island). – Acta Oecol. 18: 519-530. https://doi.org/10.1016/S1146-609X(97)80038-0

Paredes, P. & Pereira, L. S. 2019: Computing FAO56 reference grass evapotranspiration PM-ETo from temperature with focus on solar radiation. Agric. –Water Manag. 215: 86-102. https://doi.org/10.1016/j.agwat.2018.12.014

Peguero‑Pina, J.  J., Morales, F., Flexas, J., Gil‑Pelegrín, E. & Moya, I. 2008: Photochemistry, remotely sensed physiological reflectance index and de-epoxidation state of the xanthophyll cycle in Quercus coccifera under intense drought. – Oecologia 156: 1-11. https://doi.org/10.1007/s00442-007-0957-y

Peñuelas, J., Lloret, F. & Montoya, R. 2001: Severe drought effects on Mediterranean woody flora in Spain. – For. Sci. 47: 214-218. https://doi.org/10.1093/forestscience/47.2.214

Pretzsch, H., del Río, M., Arcangeli, C., Bielak, K., Dudzinska & al. 2023: Competition-based mortality and tree losses. An essential component of net primary productivity. – For. Ecol. Manag. 544: 121204. https://doi.org/10.1016/j.foreco.2023.121204

Rafiqi, M., Jelonek, L., Diouf, A.  M., Mbaye, A., Rep, M. & Diarra, A. 2022: Profile of the in silico secretome of the palm dieback pathogen, Fusarium oxysporum f. sp. albedinis, a fungus that puts natural oases at risk. – PLOS ONE 17: e0260830. https://doi.org/10.1371/journal.pone.0260830

Randhir, T. O. & Erol, A. 2013: Emerging threats to forests: resilience and strategies at system scale. – Amer. J. Pl. Sci. 4: 739-748. https://doi.org/10.4236/ajps.2013.43A093

Rita, A., Bonanomi, G., Allevato, E., Borghetti, M., Cesarano & al. 2021: Topography modulates near-ground microclimate in the Mediterranean Fagus sylvatica treeline. – Sci. Rep. 11: 8122. https://doi.org/10.1038/s41598-021-87661-6

Rivas Martínez, S. & Gandullo J. M. 1987: Memoria del mapa de series de vegetación de España: 1: 400.000. – Madrid.

Rodríguez‑Romero, M., Gallardo, A., Pérez, A. & Pulido, F. 2022: Interactive effects of biotic stressors and provenance on chemical defence induction by holm oak (Quercus ilex). –Trees 36: 227-240. https://doi.org/10.1007/s00468-021-02201-z

Roncé Le, I., Dardevet, E., Venner, S., Schönbeck, L., Gessler, A. & al. 2023: Reproduction alternation in trees: testing the resource depletion hypothesis using experimental fruit removal in Quercus ilex. – Tree Physiol. 43: 952-964. https://doi.org/10.1093/treephys/tpad025

––, Gavinet, J., Ourcival, J.-M., Mouillot, F., Chuine, I. & Limousin, J.-M. 2021; Holm oak fecundity does not acclimate to a drier world. – New Phytol. 231: 631-645. https://doi.org/10.1111/nph.17412

Ruiz-Gómez, F. J. & Miguel-Rojas, C. 2021: Antagonistic Potential of Native Trichoderma spp. against Phytophthora cinnamomi in the Control of Holm Oak Decline in Dehesas Ecosystems. – Forests 12: 945. https://doi.org/10.3390/f12070945

Salvador, C., Nieto, R., Kapwata, T., Wright, C. Y., Reason, C. & al. 2024: Analyzing the effects of drought at different time scales on cause-specific mortality in South Africa. – Environ. Res. Lett. 19: 54022. https://doi.org/10.1088/1748-9326/ad3bd2

Sena, K., Crocker, E., Vincelli, P. & Barton, C. 2018: Phytophthora cinnamomi as a driver of forest change: Implications for conservation and management. – For. Ecol. Manag. 409: 799-807. https://doi.org/10.1016/j.foreco.2017.12.022

Terradas, J. 1999: Holm Oak and Holm Oak Forests: An Introduction. – Pp. 3-14 in: Rodà, F., Retana, J., Gracia, C.A., Bellot, J. (eds), Ecology of Mediterranean Evergreen Oak Forests, 137. – Berlin.

Thompson, I., Mackey, B., McNulty, S. & Mosseler, A. 2009: Forest resilience, biodiversity, and climate change. A synthesis of the biodiversity/resilience/stability relationship in forest ecosystems – Montreal.

Tkaczyk, M. & Sikora K. 2024: Interaction between Brenneria goodwinii and Phytophthora species on young Quercus robur seedlings. – Eur. J. For. Res. 143(6): 1893-1901. https://doi.org/10.1007/s10342-024-01735-4

Vieira, J., Nabais, C. & Campelo, F. 2022: Dry and hot years drive growth decline of Pinus halepensis at its southern range limit in the Moroccan High Atlas Mountains. – Trees 36: 1585-1595. https://doi.org/10.1007/s00468-022-02314-z

Wang, Z., Lyu, W., Liu, H., Liang, J., Huang, J. & Zhang, Q.-B. 2021: Topographic patterns of forest decline as detected from tree rings and NDVI. – CATENA 198: 105011. https://doi.org/10.1016/j.catena.2020.105011

Yılmaz, O. Y., Akkemik, Ü., Doğan, H. O., Yılmaz, H., Sevgi, O. & Sevgi, E. 2024: The missing part of the past, current, and future distribution model of Quercus ilex L.: the eastern edge. – iForest 17(2): 90.  https://doi.org/10.3832/ifor4350-016

Youssef, G., Abdelaziz, M., Aboubakre, O., Abdelali, B. & Aziz, H. 2024: Impact of climate and demographic changes on the vegetation of the M’goun Geopark UNESCO of Morocco (1984-2021). – Investig. Geogr. (Alicante) 81: 225-243. https://doi.org/10.14198/INGEO.25433 

Zhang, Y., Su, T., Ma, Y., Wang, W., Zha, N. & Shao, M. 2022: Forest ecosystem service functions and their associations with landscape patterns  in Renqiu City. – PloS one 17: e0265015. https://doi.org/10.1371/journal.pone.0265015

––, Yu, Z. & Niu, H. 2018: Standardized Precipitation Evapotranspiration Index is highly correlated with total water storage over China under future climate scenarios. – Atmos. Environ. 194: 123-133. https://doi.org/10.1016/j.atmosenv.2018.09.028

Zheng, Y., Han, J., Huang, Y., Fassnacht, S.  R., Xie, S. & al. 2018: Vegetation response to climate conditions based on NDVI simulations using stepwise cluster analysis for the Three-River Headwaters region of China. – Ecol. Indic. 92: 18-29. https://doi.org/10.1016/j.ecolind.2017.06.040