Spatiotemporal dynamics of endemic macroinvertebrates in the Martil Basin (Morocco): Responses to hydroclimatic variability and anthropogenic disturbances
Abstract
This study investigates the responses of endemic macroinvertebrate species in the Martil River Basin (Morocco) to physic o chemical (T, pH, Cond and DO), land use, biotic indices (IBMWP, IHF and QBR), and hydro-morphological factors. Sampling was conducted over four seasons (2017) at 19 stations, revealing 1,768 individuals belonging to 32 endemic species. Hydro p syche iberomaroccana was the most abundant species, while Choroterpes volubilis was the most widespread. Biogeographical analysis highlighted a high proportion of Ibero-Maghrebian endemics. Multivariate analyses revealed significant biotypological differences among stream types, with sensitive endemic species predominantly found in highland / midland permanent stations, positively correlated with dissolved oxygen, altitude, and biotic indices. In contrast, lentic, thermophilic, and eurytopic species were more common in downstream areas, exposed to agricultural, industrial, and urban activities, and associated with higher temperatures and pollution levels. Seasonal and spatial variability in taxonomic composition was strongly influenced by hydr o logical intermittency and human pressures, with endemic species diversity and abundance peaking in spring. Despite their ec o logical importance, with the exception of Odonata, none of the species from other orders are currently listed on the IUCN Red List or in Annex IV of Moroccan Law 29-05 concerning the protection of wild flora and fauna species and the control of their trade. The findings underscore the urgent need for targeted conservation strategies to address threats from climate change, habitat degradation, and anthropogenic activities.References
Arribas, P., Andujar, C., Sánchez-Fernández, D., Abellán, P., & Millán, A. (2013). Integrative taxonomy and conservation of cryptic beetles in the Mediterranean region (Hydrophilidae). Zoologica Scripta, 42(2), 182–200.
Balzan, M. V., Hassoun, A. E. R., Aroua, N., Baldy, V., Dagher, M. B., Branquinho, C., Dutay, J.-C., Bour, M. E., Médail, F., Mojtahid, M., et al. (2020). First Mediterranean assessment report – Chapter 4: Ecosystems. In: Climate and environmental change in the Mediterranean basin – current situation and risks for the future. Union for the Mediterranean, Plan Bleu. Pp. 323–468.
Banegas-Medina, A., Montes, I.-Y., Tzoraki, O., Brendonck, L., Pinceel, T., Diaz, G., Arriagada, P., Arumi, J.-L., Pedreros, P., & Figueroa, R. (2021). Hydrological, environmental and taxonomical heterogeneity during the transition from drying to flowing conditions in a Mediterranean intermittent river. Biology, 10(4), 316.
Belhaj, A., Mingarro, M., Sánchez Fernández, D., Bennas, N., Chergui, B., & Pallarés, S. (2024). Conservation of freshwater biodiversity in North Africa under future climate and land-cover changes. Biodiversity and Conservation, 33, 1145–1163.
Belhaj, A., Pallarés, S., Bennas, N., Chergui, B., & Sánchez-Fernández, D. (2023b). Towards the identification of hotspots of freshwater biodiversity in North-Western Africa: A case study using species distribution models for water beetles in Morocco. Global Ecology and Conservation, 43, e02441.
Belhaj, A., Sánchez-Fernández, D., Chergui, B., & Bennas, N. (2023a). A general lack of complete inventories for aquatic beetles in Morocco. Journal of Insect Conservation, 27, 75–85.
Bemmoussat-Dekkak, S., Abdellaoui-Hassaine, K., Sartori, M., Morse, J. C., & Zamora-Muñoz, C. (2021). Larval taxonomy and distribution of genus Hydropsyche (Trichoptera: Hydropsychidae) in Northwestern Algeria. Zootaxa, 4915(4), 481–505.
Benamar, L., Bennas, N., & Millàn, A. (2011). Les coléoptères aquatiques du Parc National de Talassemtane (Nord-Ouest du Maroc). Biodiversité, degré de vulnérabilité et état de conservation [Aquatic beetles of Talassemtane National Park (Northwestern Morocco), biodiversity, vulnerability level, and conservation status]. Boletín Sociedad Entomológica Aragonesa, 49, 231–242 (in French).
Benamar, L., Bennas, N., Belhaj, A., Boulahfa, N., & Millán, A. (2021b). An updated checklist of Haliplidae, Gyrinidae, Noteridae, Hygrobiidae and Dytiscidae (Coleoptera: Adephaga) of Morocco, with notes on chorology. Aquatic Insects, 42(3–4), 247–371.
Benamar, L., Bennas, N., Hassoun, M., & Millán, A. (2021c). Threatened endemic water beetles from Morocco. Journal of Insect Conservation, 25, 465–477.
Benamar, L., Bennas, N., Hassoun, M., & Millán, A. (2022a). Checklist of Moroccan aquatic beetles (Coleoptera: Hydraenidae Mulsant, 1844). New records and updates. Zootaxa, 5129(4), 451–504.
Benamar, L., Bennas, N., Hassoun, M., & Millán, A. (2022b). Updating the presence, distribution and chorology of Moroccan Dryopoidea (Coleoptera: Elmidae and Dryopidae). Aquatic Insects, 43(4), 335–389.
Benamar, L., Bennas, N., Henegouwen, A. V. B., García-Meseguer, A. J., Hassoun, M., Benali, N., & Millán, A. (2024). The Hydrochidae (Coleoptera) of Morocco with a redescription of Hydrochus cf. obtusicollis Fairmaire, 1877. Zootaxa, 5458(2), 197–228.
Benamar, L., Millán, A., Sáinz-Cantero, C. E., Belhaj, A., & Bennas, N. (2021a). Annotated checklist of water scavenger beetles (Coleoptera: Polyphaga: Hydrophilidae) of Morocco. Aquatic Insects, 42(1), 87–108.
Bennas, N., Sáinz-Cantero, C. E. & Alba-Tercedor, J. (1992). Datos preliminares para un estudio biogeográfico del Macizo Bético-Rifeño basado en Coleópteros acuáticos [Preliminary data for a biogeographical study of the Betic-Rifian Massif based on aquatic beetles]. Zoologica Baetica, 3, 167–180 (in Spanish).
Bennas, N., Sanchez, D., Abellán, P., & Millán, A. (2009). Analyse de la vulnérabilité des coléoptères aquatiques dans la rive sud méditerranéenne: Cas du Rif Marocain [Vulnerability analysis of aquatic beetles in the southern Mediterranean shore: The case of the Moroccan Rif]. Annales de la Société Entomologique de France, 45(3), 309–320 (in French).
Bogan, M. T., Boersma, K. S., & Lytle, D. A. (2013). Flow intermittency alters longitudinal patterns of invertebrate diversity and assemblage composition in an arid-land stream network. Freshwater Biology, 58, 1016–1028.
Bonada, N., Cañedo-Argüelles, M., Gallart, F., von Schiller, D., Fortuño, P., Latron, J., Llorens, P., Múrria, C., Soria, M., Vinyoles, D., & Cid, N. (2020). Conservation and management of isolated pools in temporary rivers. Water, 12(10), 2870.
Bonada, N., Rieradevall, M., & Prat, N. (2007). Macroinvertebrate community structure and biological traits related to flow permanence in a Mediterranean river network. Hydrobiologia, 589(1), 91–106.
Bonada, N., Zamora-Muñoz, C., El Alami, M., Múrria, C., & Prat, N. (2008). New records of Trichoptera in reference Mediterranean-climate rivers of the Iberian Peninsula and North of Africa. Graellsia, 64(2), 189–208.
Bonada, N., Zamora-Muñoz, C., Rieradevall, M., & Prat, N. (2004). Ecological profiles of caddisfly larvae in Mediterranean streams: Implications for bioassessment methods. Environmental Pollution, 132, 509–521.
Boys, C. A., Rayner, T. S., Baumgartner, L. E., & Doyle, K. E. (2021). Native fish losses due to water extraction in Australian rivers. Ecological Management and Restoration, 22(2), 134–144.
Brown, L. E., Céréghino, R., & Compin, A. (2009). Endemic freshwater invertebrates from Southern France: Diversity, distribution and conservation implications. Biological Conservation, 142(12), 2613–2619.
Carey, N., Chester, E. T., & Robson, B. J. (2021). Flow regime change alters shredder identity but not leaf litter decomposition in headwater streams affected by severe, permanent drying. Freshwater Biology, 66(9), 1813–1830.
Carey, N., Chester, E. T., & Robson, B. J. (2023). Loss of functionally important and regionally endemic species from streams forced into intermittency by global warming. Global Change Biology, 29, 3019–3038.
Cid, N., Bonada, N., Carlson, S. M., Grantham, T. E., Gasith, A., & Resh, V. H. (2017). High variability is a defining component of Mediterranean-climate rivers and their biota. Water, 9(1), 52.
Clavero, M., & García-Berthou, E. (2006). Homogenization dynamics and introduction routes of invasive freshwater fish in the Iberian Peninsula. Ecological Applications, 16(6), 2313–2324.
Cummins, K. W. (1974). Structure and function of stream ecosystems. BioScience, 24(11), 631–641.
Dakki, M. (1997). Etude nationale sur la biodiversité. Biodiversité de la faune aquatique continentale (invertébrés et poissons) [National study on biodiversity: Biodiversity of continental aquatic fauna (invertebrates and fish)]. State Secretariat for the Environment, Morocco (in French).
Datry, T., Moya, N., Zubieta, J., & Oberdorff, T. (2016). Determinants of local and regional communities in intermittent and perennial headwaters of the Bolivian Amazon. Freshwater Biology, 61, 1335–1349.
Davies, P. M., & Stewart, B. A. (2013). Aquatic biodiversity in the Mediterranean climate rivers of Southwestern Australia. Hydrobiologia, 719, 215–235.
Domisch, S., Araújo, M. B., Bonada, N., Pauls, S. U., Jähnig, S. C., & Haase, P. (2013). Modelling distribution in European stream macroinvertebrates under future climates. Global Change Biology, 19(3), 752–762.
El Alami, M., El Yaagoubi, S., Gattolliat, J.-L., Sartori, M., & Dakki, M. (2022). Diversity and distribution of mayflies from Morocco (Ephemeroptera, Insecta). Diversity, 14(6), 498.
El Alami, M., Vuataz, L., El Yaagoubi, S., & Gattolliat, J.-L. (2023). A new species of the genus Alainites Waltz & McCafferty, 1994 (Ephemeroptera, Baetidae) from the north of Morocco. ZooKeys, 1176, 221–241.
El Bazi, R., El Alami, M., Khadri, O., Errochdi, S., Slimani, M., & Bennas, N. (2017). Contribution à l’étude de la biodiversité aquatique des aires protégées du Maroc [Contribution to the study of aquatic biodiversity in Morocco's protected areas]. Boletín de la Sociedad Entomológica Aragonesa, 61, 55–66 (in French).
El Haissoufi, M., Knijf, G. D., Bosch, J. V., Bennas, N., & Millán, A. (2015). Contribution to the knowledge of the Moroccan Odonata, with first records of Orthetrum sabina. Odonatologica, 44(3), 225–254.
El Haissoufi, M., L’Mohdi, O., Bennas, N., Mellado, A., & Millán, A. (2008). Les odonates du bassin versant Laou (Rif Occidental, Maroc) [The odonates of the Laou Watershed (Western Rif, Morocco)]. Travaux de l’Institut Scientifique Rabat, 5, 47–59 (in French).
El Yaagoubi, S., Edegbene, A. O., El Alami, M., Errochdi, S., & Harrak, R. (2024). Ephemeroptera, Plecoptera and Trichoptera (EPT) trait-based biomonitoring of rivers within the northwestern Rif of Morocco: Implications for determining riverine ecosystems ecological health in Africa. Aquatic Sciences, 86, 54.
El Yaagoubi, S., Errochdi, S., Edegbene, A. O., Kassout, J., Harrak, R., & El Alami, M. (2024a). Distribution patterns of Ephemeroptera, Plecoptera, and Trichoptera (EPT) species in the Northwestern Rif: Environmental and climate change impacts. Hydrobiologia, 852, 1575–1593.
Ennabili, A., Libiad, M., Haissoufi, M. E., Khabbach, A., & Bennas, N. (2025). Biodiversity, change, and use of Moroccan wetlands. In: Finlayson, A., Finlayson, C. M., & Milton, G. R. (Eds.). Wetlands of tropical and subtropical Asia and Africa. Springer, Cham. Pp. 265–296.
Errochdi, S., El Alami, M., Vinçon, G., Abdaoui, A., & Ghamizi, M. (2014). Contribution to the knowledge of Moroccan and Maghrebin stoneflies (Plecoptera). Zootaxa, 3838(1), 46–76.
Espinar-Herranz, S., Velázquez, J., Pérez-Andueza, G., Çiçek, K., & Rincón, V. (2025). Aquatic macroinvertebrates as indicators of climate change in the riparian environments of the Mediterranean region. Land, 14(3), 521.
Faramarzi, M., Abbaspour, K. C., Ashraf Vaghefi, S., Farzaneh, M. R., Zehnder, A. J. B., Srinivasan, R., & Yang, H. (2013). Modeling impacts of climate change on freshwater availability in Africa. Journal of Hydrology, 480, 85–101.
Feminella, J. W. (1996). Flow permanence and macroinvertebrate assemblages in small streams along a gradient of flow permanence. Journal of the North American Benthological Society, 15(4), 651–669.
Fernández-Calero, J. M., Cunillera-Montcusí, D., Hermoso, V., Quevedo-Ortiz, G., Fortuño, P., Acosta, R., Gomà, J., Cid, N., Vinyoles, D., Ruhí, A., Cañedo-Argüelles, M., & Bonada, N. (2024). Integrating spatiotemporal hydrological connectivity into conservation planning to protect temporary rivers. Freshwater Ecosystems, 34(3), e4139.
Figueroa, R., Bonada, N., Guevara, M., Pedreros, P., Correa-Araneda, F., Díaz, M. E., & Ruiz, V. H. (2013). Freshwater biodiversity and conservation in Mediterranean climate streams of Chile. Hydrobiologia, 719, 269–289.
Filipe, A. F., Lawrence, J. E., & Bonada, N. (2013). Vulnerability of stream biota to climate change in Mediterranean climate regions: A synthesis of ecological responses and conservation challenges. Hydrobiologia, 719, 331–351.
Gasith, A., & Resh, V. H. (1999). Streams in Mediterranean climate regions: Abiotic influences and biotic responses to predictable seasonal events. Annual Review of Ecology and Systematics, 30, 51–80.
Giam, X., Chen, W., Schriever, T. A., Van Driesche, R., Muneepeerakul, R., Lytle, D. A., & Olden, J. D. (2017). Hydrology drives seasonal variation in dryland stream macroinvertebrate communities. Aquatic Sciences, 79(3), 705–717.
Guellaf, A., & Kettani, K. (2021a). Assessing the ecological status using physic-chemical, bacteriological parameters and biotic indices of the Oued Martil River basin in Northwestern Morocco. Biologia, 76, 585–598.
Guellaf, A., & Kettani, K. (2021b). Performance of macroinvertebrates-based biotic indices for assessing water quality in Martil River Basin, Northern Morocco. Environmental Engineering and Management Journal, 20(1), 141–150.
Guellaf, A., El Alami, M., Kassout, J., Errochdi, S., Khadri, O., & Kettani, K. (2021). Diversity and ecology of aquatic insects (Ephemeroptera, Plecoptera, and Trichoptera) in the Martil Basin (Northwestern Morocco). Community Ecology, 22, 331–350.
Hajji, K. (2017). Les Trichoptères du Maroc: Atlas, biogéographie et degré de vulnérabilité [The caddisflies (Trichoptera) of Morocco: Atlas, biogeography, and vulnerability assessment]. Abdelmalek Essaâdi University, Tétouan (in French).
Hajji, K., Zamora-Muñoz, C., Bonada, N., & El Alami, M. (2012). Quelques notes sur l’écologie et distribution des Rhyacophilidae du Rif (Nord du Maroc) [Some notes on the ecology and distribution of Rhyacophilidae in the Rif (Northern Morocco)]. Boletín de la Sociedad Entomológica Aragonesa, 50, 559–562 (in French).
Jáimez-Cuéllar, P., Vivas, S., Bonada, N., Robles, S., Mellado, A., Álvarez, M., Avilés, J., Casas, J., Ortega, M., Pardo, I., Prat, N., Rieradevall, M., Sáinz-Cantero, C. E., Sánchez-Ortega, A., Suárez, L., Toro, M., Vidal-Abarca, M. R., Zamora-Muñoz, C., & Alba-Tercedor, J. (2002). Protocolo GUADALMED (PRECE). Limnetica, 21(3–4), 187–204.
Karrouchi, M., Touhami, M. O., Oujidi, M., & Chourak, M. (2016). Cartographie des zones à risque d’inondation dans la région Tanger-Tétouan: Cas du bassin versant de Martil, Nord du Maroc [Mapping of flood risk areas in the Tangier-Tetouan region: Case of the Martil watershed, northern Morocco.]. International Journal of Innovation and Applied Studies, 14(4), 1019–1035 (in French).
Khelifa, R., Deacon, C., Mahdjoub, H., Suhling, F., Simaika, J. P., & Samways, M. J. (2021). Dragonfly conservation in the increasingly stressed African Mediterranean-type ecosystems. Frontiers in Environmental Science, 9, 660163.
Kroll, S. A., Ringler, N. H., Costa, M. D. L. C. C., & Ibañez, J. D. L. H. (2017). Macroinvertebrates on the front lines: Projected community response to temperature and precipitation changes in Mediterranean streams. Journal of Freshwater Ecology, 32(1), 513–528.
L’Mohdi, O., El Haissoufi, M., Bennas, N., & Millán, A. (2008). Contribution à la connaissance des Hétéroptères aquatiques du bassin versant Laou [Contribution to the knowledge of aquatic Heteroptera in the Laou Watershed]. Travaux de l’Institut Scientifique, Rabat, Série Générale, 5, 67–74 (in French).
Laouina, A. (2006). Gestion durable des ressources naturelles de la biodiversité au Maroc – prospective Maroc 2030 [Sustainable management of natural resources and biodiversity in Morocco – prospective Morocco 2030]. Report of the High Commission for Planning, Morocco (in French).
Lavergne, S., Hampe, A., & Arroyo, J. (2012). In and out of Africa: How did the Strait of Gibraltar affect plant species migration and local diversification? Journal of Biogeography, 40(1), 24–36.
Leland, H. V., & Porter, S. D. (2000). Distribution of benthic algae in the upper Illinois River basin in relation to geology and land use. Freshwater Biology, 44(2), 279–301.
Magoulick, D. D., & Kobza, R. M. (2003). The role of refugia for fishes during drought: A review and synthesis. Freshwater Biology, 48, 1186–1198.
Médail, F., & Quézel, P. (1997). Hot-spots analysis for conservation of plant biodiversity in the Mediterranean Basin. Annals of the Missouri Botanical Garden, 84, 112–127.
Merritt, R. W., & Cummins, K. W. (1996). An introduction to the aquatic insects of North America. 3rd ed. Kendall, Hunt.
Munné, A., & Prat, N. (2011). Effects of Mediterranean climate annual variability on stream biological quality assessment using macroinvertebrate communities. Ecological Indicators, 11(3), 651–662.
Munné, A., Prat, N., Solà, C., Bonada, N., & Rieradevall, M. (2003). A simple field method for assessing the ecological quality of riparian habitat in rivers and streams: QBR index. Aquatic Conservation, 13(2), 147–163.
Munné, A., Sola, C., & Prat, N. (1998). QBR: Un índice rápido para la evaluación de la calidad de los ecosistemas de ribera [QBR: A rapid index for assessing riparian ecosystem quality]. Tecnología del Agua, 175, 20–37 (in Spanish).
Myers, N., Mittermeier, R. A., Mittermeier, C. G., da Fonseca, G. A. B., & Kent, J. (2000). Biodiversity hotspots for conservation priorities. Nature, 403, 853–858.
Nogueira, A. R., Gama, J., & Ferreira, C. A. (2021). Causal discovery in machine learning: Theories and applications. Journal of Dynamics and Games, 8, 203–231.
Pardo, I., Álvarez, M., Casas, J., Moreno, J. L., Vivas, S., Bonada, N., Alba-Tercedor, J., Jáimez-Cuéllar, P., Moyà, G., Prat, N., Robles, S., Suárez, M. L., Toro, M., & Vidal-Abarca, M. (2002). El hábitat de los ríos mediterráneos: Diseño de un índice de diversidad de hábitat [The habitat of Mediterranean rivers: Design of a habitat diversity index]. Limnetica, 21(3–4), 115–133 (in Spanish).
Petchey, O. L., & Gaston, K. J. (2002). Extinction and the loss of functional diversity. Proceedings of the Royal Society B: Biological Sciences, 269(1501), 1721–1727.
Ribera, I., Castro, A., Dı́az, J. A., Garrido, J., Izquierdo, A., Jäch, M. A., & Valladares, L. F. (2011). The geography of speciation in narrow endemics of the ‘‘Haenydra’’ lineage (Coleoptera, Hydraenidae, Hydraena). Journal of Biogeography, 38(3), 502–516.
Riservato, E., Boudot, J. P., Ferreira, S., Jovic, M., Kalkman, V. J., Schneider, W., & Samraoui, B. (2009). The status and distribution of dragonflies of the Mediterranean Basin. IUCN, Gland.
Salhi, A., Martin-Vide, J. M., Benhamrouche, A., Benabdelouahab, S., Himi, M., Benabdelouahab, T., & Casas Ponsati, A. (2019). Rainfall distribution and trends of the daily precipitation concentration index in Northern Morocco: A need for an adaptive environmental policy. SN Applied Sciences, 1(3), 277.
Sánchez-Fernández, D., Abellán, P., Velasco, J., & Millán, A. (2004). Selecting areas to protect the biodiversity of aquatic ecosystems in a semiarid Mediterranean region using water beetles. Aquatic Conservation: Marine and Freshwater Ecosystems, 14(5), 465–479.
Sánchez-Fernández, D., Bilton, D. T., Abellán, P., Ribera, I., Velasco, J., & Millán, A. (2008). Are the endemic water beetles of the Iberian Peninsula and the Balearic Islands effectively protected? Biological Conservation, 141(7), 1612–1627.
Sansoni, G. (1992). Atlante per il riconoscimento dei macroinvertebrati dei corsi d'acqua Italiani [Atlas for the identification of macroinvertebrates in Italian watercourses]. Italian Center for Environmental Biology Studies, Trento (in Italian).
Slimani, M., Bennas, N., El Haissoufi, M., L’Mohdi, O., Khadri, O., El Bazi, R., & El Alami, M. (2016). Contribution à l’étude de la biodiversité aquatique des aires protégées du Maroc. [Contribution to the study of aquatic biodiversity in the protected areas of Morocco.] Boletín de la Sociedad Entomológica Aragonesa, 58, 168–180.
Smith, R. J., Bennun, L., Brooks, T. M., Butchart, S. H. M., Cuttelod, A., Di Marco, M., Ferrier, S., Fishpool, L. D. C., Joppa, L., Juffe‐Bignoli, D., Knight, A. T., Lamoreux, J. F., Langhammer, P., Possingham, H. P., Rondinini, C., Visconti, P., Watson, J. E. M., Woodley, S., Boitani, L., … Scaramuzza, C. A. de M. (2018). Synergies between the key biodiversity area and systematic conservation planning approaches. Conservation Letters, 12(1), 12625.
Strachan, S. R., Chester, E. T., & Robson, B. J. (2015). Freshwater invertebrate life history strategies for surviving desiccation. Springer Science Reviews, 3(1), 57–75.
Stubbington, R., & Datry, T. (2013). The macroinvertebrate seedbank promotes community persistence in temporary rivers across climate zones. Freshwater Biology, 58(6), 1202–1220.
Tachet, H., Richoux, P., Bournaud, M., & Usseglio-Polatera, P. (2002). Invertébrés d'eau douce: Systématique, biologie, écologie [Freshwater invertebrates: Systematics, biology, ecology]. CNRS Éditions, Paris (in French).
Taybi, A. F., Mabrouki, Y., Chavanon, G., Berrahou, A., & Millán, A. (2018). New data on the distribution of aquatic bugs (Hemiptera) from Eastern Morocco with notes on chorology. Zootaxa, 4459(1), 139–163.
Tierno de Figueroa, J. M., López-Rodríguez, M. J., Fenoglio, S., Sánchez-Castillo, P., & Fochetti, R. (2013). Freshwater biodiversity in the rivers of the Mediterranean Basin. Hydrobiologia, 719, 137–186.
Touaylia, S. (2017). Thoughts on water beetles in a Mediterranean environment. In: Shields, V. D. C. (Ed.). Insect physiology and ecology. IntechOpen. Pp. 3–23.
Trizzino, M., Bisi, F., Maiorano, L., Martinoli, A., Petitta, M., Preatoni, D. G., & Audisio, P. (2014). Mapping biodiversity hotspots and conservation priorities for the Euro-Mediterranean headwater ecosystems, as inferred from diversity and distribution of a water beetle lineage. Biodiversity and Conservation, 24(1), 149–170.
Zerrouk, M., Dakki, M., El Agbani, M. A., El Alami, M., Bennas, N., Qninba, A., & Himmi, O. (2021). Evolution of the benthic communities in a North-African river, the upper Sebou (Middle Atlas – Morocco) between 1981 and 2017: Effects of global changes. Biologia, 76, 2683–2696.
