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29 October 2026

19 Pages

Towards a Comprehensive Inventory of Freshwater Macroinvertebrates in Northeastern Morocco: Diversity Patterns, Endemism, and Potential Implications for Aquatic Ecosystem Sustainability

,
and
1
Synergy Lab, Higher School of Education and Training, Mohammed I University, BP-410, Oujda 60000, Morocco
2
Molecular Ecology Group (MEG), Water Research Institute (IRSA), National Research Council of Italy (CNR), 28922 Verbania, Italy
3
Laboratory of Biotechnology, Conservation and Valorization of Bioresources, Faculty of Sciences Dhar El Mehraz, Sidi Mohamed Ben Abdellah University, Fes 30000, Morocco
*
Author to whom correspondence should be addressed.
Sustainability2026, 18(21), 11003;https://doi.org/10.3390/su182111003
This article belongs to the Section Sustainability, Biodiversity and Conservation
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Abstract

Freshwater macroinvertebrate diversity remains unevenly documented across North Africa, limiting the use of biodiversity data in conservation planning. Here, we provide a regional synthesis of freshwater macroinvertebrates from the Moulouya River basin and the Eastern Region of Morocco, integrating records from historical literature, expert knowledge, and recent field observations. The dataset documents 525 species, 269 genera, 114 families, and 4924 georeferenced occurrence records compiled between 1914 and 2026. Taxonomic representation was highly uneven, with aquatic insects dominating the dataset and several small-bodied, cryptic, subterranean, and taxonomically difficult groups remaining poorly documented. Chorological analyses revealed a predominance of Palearctic taxa with Mediterranean affinities, together with a substantial proportion of Moroccan, Maghrebian, and Ibero-Maghrebian endemics. Strict Moroccan endemics were especially concentrated in gastropods and were mainly associated with lotic ecosystems and springs. Spatial and temporal patterns revealed apparent richness concentrations in northern and western sectors of the study area, but also strong biases linked to uneven sampling effort. This synthesis provides a baseline for future monitoring, highlights priority habitats and taxonomic gaps, and offers actionable insights to support sustainable water resource management and ecosystem resilience under accelerating climate change and anthropogenic pressures in North African arid and semi-arid regions.

1. Introduction

Morocco occupies an important biogeographical position between Europe and Africa and forms part of the Mediterranean Basin global biodiversity hotspot [1,2]. Influenced by Mediterranean, Atlantic, and Afrotropical biogeographical elements, the country represents a region of faunal exchange and diversification [3,4]. Its complex topography, climatic gradients, and geographic barriers have contributed to high levels of endemism across multiple taxonomic groups, including freshwater fauna [5,6]. Morocco also harbors some of the most extensive and environmentally heterogeneous freshwater systems in North Africa, including river basins, wetlands, springs, temporary streams, groundwater systems, and subterranean aquatic ecosystems [6].
Morocco’s freshwater ecosystems are particularly vulnerable to environmental change, as water availability is constrained by climatic variability and increasing human pressures, including water abstraction and land-use change [7]. Reliable biodiversity data are therefore essential for identifying threatened habitats, assessing ecological risks, and informing conservation strategies [8]. Accurate species identification, georeferenced distribution records, and regularly updated inventories provide the baseline information needed to assess biodiversity patterns and knowledge gaps at regional and national scales [9,10]. Combined with ecological and environmental data, these inventories can inform habitat protection, biodiversity monitoring, invasive species management, and the identification of poorly studied areas [11,12,13].
Freshwater macroinvertebrates represent key components of surface and subterranean aquatic ecosystems, contributing to organic matter processing, nutrient cycling, trophic interactions, and sediment dynamics [14,15]. In groundwater-dependent ecosystems, specialized subterranean fauna may also contribute to detritus processing and microbial regulation under nutrient-limited conditions [16,17]. Surface-dwelling taxa often display habitat specificity and turnover along environmental gradients, including temperature, flow regime, and substrate composition [18,19]. In contrast, groundwater and subterranean fauna frequently exhibit morphological and ecological specialization, limited dispersal, and high endemism associated with long-term isolation in aquifers and karst systems [16,20,21]. These eco-evolutionary characteristics make freshwater macroinvertebrates valuable indicators of ecosystem integrity, hydrological connectivity, habitat degradation, and long-term environmental change.
Updated regional faunal inventories provide essential baseline information for freshwater conservation, documenting species richness, endemism, biogeographical affinities, habitat associations, and gaps in biodiversity knowledge. They can also support monitoring of alien species and help identify habitats requiring further monitoring or conservation attention [6,22,23].
In northeastern Morocco, the Moulouya River basin and the Eastern Region encompass diverse aquatic environments, including permanent and temporary rivers, springs, wetlands, reservoirs, saline and brackish systems, and groundwater-dependent habitats [6]. Here, we compile an updated inventory of freshwater macroinvertebrates from this region, integrating historical literature, previous taxonomic research, and recent field observations. We examine taxonomic richness, chorological composition, endemism, habitat associations, and the spatial and temporal distribution of documented records. Specifically, we aim to (i) characterize the taxonomic and biogeographical composition of the known fauna; (ii) identify habitats and areas with high document richness and endemic; and (iii) highlight taxonomic, geographical and ecological gaps relevant to future conservation planning. This synthesis provides a baseline for improving freshwater biodiversity knowledge and informing the conservation of surface-water and groundwater ecosystems in northeastern Morocco.

2. Materials and Methods

2.1. Study Area

The Moulouya River is one of Morocco’s longest rivers, extending over nearly 600 km across northeastern Morocco before reaching the Mediterranean Sea (Figure 1). Its principal permanent tributaries include the Ansegmir, Melloulou, Za, and Msoun rivers, as well as several intermittent streams characterized by seasonal and episodic flow regimes [24]. The Moulouya River basin covers approximately 56,000 km2, extending mainly across Morocco, with a small portion (3.1%) located in Algeria. On the Moroccan side, this basin heavily intersects the Eastern Region, occupying 34,990 km2 (or 62.48% of the basin). At the national level, this administrative region alone covers 90,130 km2, accounting for 12.7% of the Moroccan territory. The study area, which encompasses the entire Eastern Region plus the portions of the watershed extending beyond its administrative boundaries, thus covers 109,390 km2, representing 15.41% of the country’s total surface area. This vast territory is characterized by remarkable ecological diversity. For the purposes of this study, this geographical and administrative entity is referred to as “northeast Morocco”.
Figure 1. Location of the Eastern Region of Morocco and the Moulouya River basin.
The Moulouya basin has a predominantly Mediterranean hydrological and climatic regime, with peak discharge in late winter and early spring, and low-flow conditions during summer. The regional climate ranges from semi-arid to sub-humid. The study area encompasses a high diversity of aquatic ecosystems, including springs, streams, rivers, ponds, wetlands, reservoirs, and coastal saline and brackish environments. Notable wetlands include Ramsar sites such as the Nador lagoon, the Moulouya estuary, and the Mohammed V dam.

2.2. Data Collection

We compiled freshwater macroinvertebrate occurrence records from the Moulouya River basin and the Eastern Region of Morocco using two principal sources: (i) a targeted review of national and international literature [24,25,26,27,28,29,30,31,32,33,34,35,36,37,38,39,40,41,42,43,44,45,46,47,48,49,50,51,52,53,54,55,56,57,58,59,60,61,62,63,64,65,66,67,68,69,70,71,72,73,74,75,76,77,78,79,80,81,82,83,84,85,86,87,88,89,90,91,92,93,94,95,96,97,98,99,100,101,102,103,104,105,106,107,108,109]; (ii) previous field surveys and additional observations by the authors conducted between 2014 and 2026 [6,110,111,112,113,114,115,116,117,118,119,120,121,122,123,124,125,126,127,128,129,130,131,132,133,134,135,136,137,138,139,140,141,142,143,144,145,146,147,148,149,150,151,152,153,154,155,156,157,158].
For this study, freshwater macroinvertebrates were defined as aquatic and semi-aquatic invertebrates associated with inland aquatic habitats, such as rivers, streams, springs, wetlands, reservoirs, ponds, groundwater-dependent systems, hyporheic habitats, caves, wells, and slightly brackish inland or coastal-associated systems when taxa were considered part of the non-marine aquatic fauna. Strictly marine species were excluded, except for alien species documented in freshwater or brackish inland habitats. Aquatic insects were included when at least one life stage was aquatic or closely associated with aquatic habitats. Semi-aquatic taxa were retained when included in regional aquatic macroinvertebrate inventories or biomonitoring studies. Fossil records, taxa considered as nomina dubia, and records with uncertain taxonomic status were excluded.
Published sources were identified through targeted Google Scholar searches combining geographical terms (“Moulouya”, “Oriental Morocco”, “eastern Morocco”) with habitat-related terms (“freshwater”, “macroinvertebrates”, “aquatic invertebrates”, “groundwater”, “springs”, “caves”), and taxonomic terms (“Coleoptera”, “Odonata”, “Mollusca”, “Crustacea”, “Diptera”, “Ephemeroptera”, “Trichoptera”, “Plecoptera”, “Heteroptera”, “Hydrachnidia”). The search was conducted between 2014 and 2026, with publications retained when they reported identifiable freshwater macroinvertebrate taxa from localities within the study area.

2.3. Data Validation and Standardization

For each record, we extracted the accepted taxon name, original taxon name when available, locality, date or period of collection, habitat information, source reference, and geographic coordinates when provided. Duplicate records referring to the same taxon, locality, date, and source were removed. Records from repeated surveys at the same locality were retained when they represented distinct sampling events.
Geographic coordinates were standardized in ArcGIS. Historical records lacking coordinates were georeferenced when locality descriptions were sufficiently precise. These estimated coordinates are flagged with an asterisk (*) in Supplementary Table S2. Records with ambiguous or insufficiently precise locality information, such as references to entire provinces or river basins, were excluded from spatial analyses. For temporal analyses, records were grouped into 20-year intervals based on the publication year of their source, rather than their collection date.
To ensure data interoperability and long-term accessibility, the final validated dataset was formatted in accordance with the Darwin Core standard. The standardized georeferenced occurrence dataset was subsequently published in the Global Biodiversity Information Facility (GBIF) network and is permanently accessible [159].
Taxonomic names were standardized according to recent taxonomic revisions and verified against authoritative databases (WoRMS, MolluscaBase, and FADA) accessed between June and August 2026. Taxonomic assignments were additionally checked against published revisions and, where applicable, validated in consultation with taxonomic specialists. Literature records were cross-referenced with the authors’ research databases to identify and exclude synonymies, resolve taxonomic inconsistencies, and detect potential duplicates.
For the temporal analysis, occurrences were aggregated into 20-year intervals based strictly on the publication year of the source studies. Field collection dates were not processed in this analysis, ensuring that every record was assigned a definitive temporal baseline corresponding to its publication date.
Chorological categories followed classifications of La Greca [160] and Vigna Taglianti et al. [161], complemented by taxon-specific revisions. Habitat classified in four categories: (A) groundwater, including caves, aquifers, phreatic and interstitial habitats; (B) springs, including spring heads, pools and outflows; (C) lotic waters, including rivers, creeks, rivulets, and streams; and (D) lentic waters, including lakes, ponds, and wetlands. Presence of taxa in each category is indicated with value 1 in Supplementary Table S1. We use the term stygobiont to refer to taxa exclusively found in groundwater habitats; and crenobiont for those exclusively found in springs. For analyses of habitat associations, each species–habitat combination was counted separately; consequently, species recorded in multiple habitats contributed to more than one category. Habitat percentages were calculated relative to the total number of species–habitat associations rather than the number of species.

2.4. Data Visualization

A map illustrating the spatial distribution of species richness was constructed by plotting the coordinates and total species counts per locality in ArcGIS software (version 10.2; Esri, Redlands, CA, USA).

3. Results

3.1. Taxonomic Composition

Our dataset included 525 freshwater invertebrate species belonging to 269 genera and 114 families, represented by 4924 georeferenced occurrence records compiled from historical literature and authors’ field observations. Supplementary Material Table S1 provides the species checklist, including geographical distributions and habitat associations, whereas Supplementary Material Table S2 contains occurrences along with GPS coordinates, collection dates and source references.
A few families accounted for a substantial proportion of records, whereas most families had relatively few recorded species and occurrences (Figure 2). The beetle family Dytiscidae was the most represented family in both species richness and number of records, accounting for 13.55% of all occurrences. Other well-represented beetle families included Hydrophilidae (7.03%) and Hydraenidae (5.28%). Among other insect families, Libellulidae accounted for 8.49% of records (16 species), followed by Coenagrionidae (5.36%; 13 species) and Baetidae (4.43%; 10 species). Elmidae (3.53%; 10 species) and Corixidae (3.45%; 15 species) were moderately represented. In contrast, Hydrobiidae contributed 19 species but only 0.85% of occurrence records. Overall, these results indicate considerable variation in the taxonomic representation of the compiled records, potentially reflecting differences in species richness, detectability, and historical sampling effort among groups.
Figure 2. Species richness and number of occurrences across freshwater invertebrate families in the Moulouya River basin and Eastern Region of Morocco. Each point represents a family, with species richness on the x-axis and the number of occurrence records on the y-axis. Circle size is proportional to the number of records. And percentage families indicate each family’s contribution to the total number of records (N = 4924).

3.2. Chorological Composition

Of the 458 species with chorological assignments, 395 (86.2%) belonged to Palearctic or Mediterranean chorotypes, whereas 63 (13.8%) were widely distributed (Figure 3A). The former comprised 156 strictly Palearctic species (34.1% of the classified fauna) and 239 species with Mediterranean affinities (52.2%).
Figure 3. Chorological composition of freshwater macroinvertebrates in northeastern Morocco. (A) Distribution of the 458 species with chorological assignments among the main chorotypes and their subcategories. (B) Proportion of alien and/or invasive species among widely distributed taxa (n = 63). Percentages in each subcategory are calculated relative to its corresponding parent category. Absolute counts are reported in Section 3.2, and species-level classifications are provided in Supplementary Table S1.
Within this Mediterranean component (n = 239), endemic taxa in the broad sense accounted for 52.7% (n = 126), followed by West Mediterranean (23.8%, n = 57), Holo-Mediterranean (13.8%, n = 33), and Atlanto-Mediterranean taxa (9.6%, n = 23). Among endemic taxa (n = 126), 48 (38.1%) were strictly Moroccan endemics, whereas 36 (23.6%) had Ibero-Maghrebian distribution, including five restricted to the Ibero-Moroccan region.
Among the widely distributed taxa (n = 63), nine (14%) were classified as alien and/or invasive (Figure 3B), including Helobdella europaea Kutschera, 1987; Branchiura sowerbyi Beddard, 1892; Physella acuta (Draparnaud, 1805), Potamopyrgus antipodarum (J.E. Gray, 1843); and Callinectes sapidus (Rathbun, 1896). These nine species represented 2.0% of the fauna with chorological assessments. Species-level classifications are provided in Supplementary Table S1.

3.3. Strictly Endemic Species Composition

A total of 49 species were identified as strict Moroccan endemics, highlighting the importance of the Moulouya River basin and the Eastern Region as a regional center of endemism. The faunistic composition of these endemic species is dominated by Gastropoda (n = 19; 38.8%), followed by Amphipoda (n = 6; 12.2%), Trichoptera (n = 5; 10.2%), and Diptera (n = 4, 8.2%). Ephemeroptera and Coleoptera accounted for three taxa (n = 3; 7%), whereas Hemiptera (n = 2; 4%), Isopoda (n = 2; 4%), Plecoptera (n = 2; 4%), and Bivalvia (n = 2; 4%) each contributed two taxa (4.1%). Hydrachnidia was represented by a single taxon (2%) (Figure 4A). The 49 endemic taxa accounted for 65 species–habitat associations, as some taxa occurred in multiple habitats. Most associations involved lotic ecosystems (n = 26; 40.0%) and springs (n = 23; 35.4%), whereas lentic and groundwater ecosystems each accounted for eight associations (12.3%) (Figure 4B).
Figure 4. Faunistic composition of strictly endemic Moroccan species (N = 49) in the Moulouya River basin and the Eastern Region of Morocco (A), and percentages of their habitat categories (B).

3.4. Temporal Trends in Taxonomic Group Composition

Temporal distribution of published occurrence records was highly uneven (Figure 5). Few records were associated with publications before 1950, followed by a gradual increase between the 1950s and 1980s. Record numbers increased substantially from the 1990s onwards, particularly after 2000, reaching their highest levels during 2010–2020. This increase was accompanied by broader taxonomic representation and is reflected in the steep rise in the cumulative record curve. These patterns reflect the historical accumulation of published biodiversity information rather than temporal changes in species richness or abundance.
Figure 5. Temporal distribution and cumulative accumulation of scientific publications and records over time in the study area database. Stacked bars represent the number of records assigned to each taxonomic group within each period, while the red line indicates the cumulative number of records over time. The figure illustrates both temporal variation in sampling intensity and the progressive growth of the database.

3.5. Spatial Patterns of Documented Species Richness

The highest documented species richness was recorded in the northern and northeastern sectors of the study area, particularly in the lower Moulouya wetlands and the watercourses draining the eastern slopes of the Middle Atlas (Figure 6). Other concentrations of documented richness occurred in the Oued Za basin, the Nador Lagoon area, and the Oued Isly basin. In contrast, relatively few species were recorded in the southern and southeastern sectors, including the Figuig Oasis. These spatial patterns should be interpreted cautiously, as they reflect documented occurrences and may be influenced by uneven historical sampling effort.
Figure 6. The spatial distribution of species within the study area. The red circles indicate observation points, and their size is proportional to the number of species recorded (ranging from 1 to 97).

4. Discussion

4.1. A Regional Baseline Shaped by Uneven Knowledge

This synthesis provides an updated regional inventory of freshwater macroinvertebrates from the Moulouya Basin and the Eastern Region, documenting 525 species, 269 genera, 114 families, and 4924 georeferenced occurrence records. These data support the region’s biodiversity importance but also highlight substantial taxonomic, spatial, and temporal gaps in current knowledge.
Aquatic insects dominate the compiled records, particularly Dytiscidae, Hydrophilidae, Hydraenidae, Libellulidae, Coenagrionidae, and Baetidae. Although this pattern reflects their contribution to the known regional fauna, it may also be influenced by historical research efforts and taxonomic expertise [24,41,44,50]. As reported elsewhere [162], conspicuous and well-studied groups may be more frequently documented than small-bodied, cryptic, subterranean, or taxonomically challenging taxa. The relatively recent documentation of groups such as water mites (Hydracarina), aquatic Curculionidae, Hymenoptera, and Lepidoptera illustrates gaps in historical taxonomic coverage [113,130,131,134,136]. Their late appearance in the inventory rather reflects increasing research attention rather than recent colonization.
Temporal and spatial patterns also reflect uneven documentation. The number of published occurrence records increased substantially after the 1990s, particularly after 2000, coinciding with the expansion of regional hydrobiological research followed by improvements in taxonomic knowledge and sampling coverage. Higher documented richness in the northern, northeastern, and western sectors may partly reflect greater habitat availability, and more permanent aquatic environments, but also differences in sampling effort, accessibility, and research activity. Conversely, the relatively few records from southern and southeastern sectors cannot be interpreted as evidence of intrinsically lower biodiversity, despite the aridity, intermittent flow, and higher environmental stress in these areas. These patterns emphasize the need for more geographically balanced surveys before regional differences in species richness can be assessed reliably [163].

4.2. Endemism and Biogeographical Affinities

The predominance of Palearctic and Mediterranean taxa, together with the substantial proportion of Moroccan, Maghrebian, and Ibero-Maghrebian endemics, highlights the biogeographical affinities of northeastern Morocco’s freshwater macroinvertebrate fauna. These patterns are consistent with other regional faunal groups [49,164,165,166] and with Morocco’s geographical position between Europe and Africa. Accordingly, the known fauna shows stronger affinities with western Mediterranean and European assemblages, whereas Afrotropical elements remain poorly represented [6,167,168].
Historical connections between Europe and North Africa may have contributed to these patterns, facilitating faunal exchanges through the western Mediterranean and potentially other Mediterranean dispersal routes, via the Iberian Peninsula and western Maghreb, and the Italian and Tunisian regions [119]. Changes in geographical connectivity during the Messinian Salinity Crisis may have influenced the dispersal and diversification of freshwater organisms, when the desiccation of the Mediterranean Sea created extensive land bridges and interconnected fluvial systems [169,170]. Subsequent geographical isolation, climatic fluctuations, and topographical barriers may have further contributed to regional differentiation and endemism.
Strict Moroccan endemics were particularly well-represented among gastropods, followed by amphipods and trichopterans. This pattern is consistent with the high endemism reported for Moroccan freshwater molluscs [6] and across the Mediterranean Basin [171]. Limited dispersal, habitat specialization and geographical isolation may contribute to restricted distributions and diversification in freshwater molluscs [6,172,173]. The high number of endemic species documented in rivers and springs suggests that these environments may warrant conservation attention. However, further systematic surveys, particularly in isolated springs, headwaters, hyporheic habitats, and groundwater systems, are needed to address existing knowledge gaps and assess whether additional endemic diversity remains undocumented [174].

4.3. Threats to Endemic and Groundwater-Dependent Biodiversity

Endemic and habitat-specialized taxa may be particularly vulnerable to environmental change because of their restricted distributions and limited dispersal capacity. Major threats include habitat degradation, hydrological alteration, pollution, salinization, and biological invasions [6,127,145]. These pressures are especially relevant in semi-arid regions, where water availability is constrained by climatic variability and increasing human water demand [175].
Freshwater ecosystems in the Moulouya Basin and the Eastern Region are exposed to multiple anthropogenic stressors, including riparian alteration, dam construction, overgrazing, water abstraction, and domestic, industrial, and agricultural pollution [118,176,177]. Groundwater resources are also affected by mineralization and contamination linked to agricultural and industrial activities [178,179,180,181]. Such pressures are particularly concerning for springs, and hyporheic habitats, which may harbor poorly documented endemic and highly specialized taxa.
Salinization is another important pressure, potentially exacerbated by declining precipitation, evaporation, seawater intrusion, and hydrological alteration, particularly in downstream areas [127]. Increasing salinity can alter community composition, reduce suitable habitat for freshwater specialists, and favor salt-tolerant taxa. The vulnerability of habitat-specialized species is illustrated by Aphelocheirus pemae Millán, L’Mohdi & Carbonell, 2016, a geographically restricted aquatic insect reported only from North Africa’s Moroccan fauna [129,140,145]. Its limited distribution and association with running waters make its populations potentially vulnerable to habitat degradation, pollution, and hydrological changes. These pressures highlight the importance of integrating freshwater biodiversity conservation into sustainable water-resource management, particularly in arid and semi-arid regions [118,142,145].
Biological invasions interact with these pressures. Alien or invasive macroinvertebrates reported for the region include Barbronia weberi (Blanchard 1897), Planorbella duryi (Wetherby, 1879), Corbicula fluminalis (O.F. Müller, 1774) and Trichocorixa verticalis verticalis (Fieber, 1851) [182]. Additionally, invasive alien fishes, including Gambusia holbrooki Girard, 1859, Micropterus salmoides Lacépède, 1802, and Cyprinus carpio Linnaeus, 1758 [182], represent additional potential pressures on native aquatic communities. Alien species may affect native fauna through competition, predation or changes in trophic interactions [183,184]. The reported upstream expansion of Callinectes sapidus, facilitated by its broad salinity tolerance, illustrates the potential interaction between biological invasions and salinity gradients [127]. Local disappearance of Potamon algeriense (Herbst, 1785) and Unio foucauldianus Pallary, 1936 have also been reported from parts of the Moulouya River, although their causes remain uncertain [6,127].

4.4. Research Priorities and Potential Frameworks for Long-Term Ecosystem Sustainability

This checklist provides a baseline for future research and conservation planning rather than a complete estimate of freshwater invertebrate diversity in northeastern Morocco. Four main priorities emerge from the gaps identified here.
First, future surveys should broaden their taxonomic coverage to include small-bodied and cryptic invertebrates, particularly meiofaunal groups such as nematodes, small annelids, microcrustaceans, rotifers, and gastrotrichs. These organisms contribute to sediment processes, microbial grazing, and organic matter decomposition [174], but remain poorly documented in regional freshwater inventories [185].
Second, groundwater-dependent ecosystems warrant dedicated attention. Springs, aquifers, wells, caves, hyporheic zones, and interstitial habitats may support specialized and endemic assemblages, potentially serving as refugia during periods of surface-water scarcity [186,187]. However, these habitats are vulnerable to water abstraction, pollution, salinization, and hydrological alteration [175,188]. Standardized surveys would help clarify their biodiversity and conservation relevance [189].
Third, taxonomic and molecular resources should be improved. Identification remains challenging for several freshwater invertebrate groups because of unresolved taxonomy, cryptic diversity, and limited regional reference data [190]. Integrative approaches combining morphology and molecular evidence could improve species delimitation and the reliability of biodiversity inventories [191]. Developing a regional DNA reference library would facilitate future taxonomic research and biomonitoring.
Finally, biodiversity inventories should be integrated with standardized, long-term ecological monitoring. Although occurrence records provide valuable baseline information, they cannot independently establish population trends, local extinctions, or responses to environmental change [192,193]. Repeated biological surveys combined with environmental measurements, including water chemistry, temperature, salinity, flow permanence, land use, and groundwater conditions, would improve assessments of ecological change [19,194]. Publicly accessible, standardized biodiversity data would also enhance reproducibility and facilitate their integration into regional monitoring and conservation initiatives [195]. Such coordinated efforts could strengthen the contribution of freshwater biodiversity information to sustainable water-resource management in northeastern Morocco.

5. Conclusions

This study provides an updated regional synthesis of freshwater macroinvertebrate diversity in the Moulouya River basin and the Eastern Region of Morocco, documenting substantial species richness, predominantly Palearctic–Mediterranean affinities, and a notable proportion of endemic species. Rivers and springs accounted for the highest numbers of documented endemic habitat associations, highlighting their potential conservation importance.
However, the dataset reveals gaps in taxonomic, spatial, and temporal knowledge, particularly small-bodied taxa and subterranean fauna, groundwater-dependent ecosystems and the southern sectors of the study area. Uneven historical sampling also limits the interpretation of spatial richness patterns. These limitations emphasize that the inventory represents a baseline of documented biodiversity rather than a complete assessment of regional species richness or conservation priorities.
Future work should focus on standardized surveys, integrative taxonomy, public deposition of occurrence data, and long-term monitoring. Expanding coverage of poorly studied taxa and habitats, particularly groundwater and hyporheic systems, would improve biodiversity assessments and help integrate freshwater conservation into sustainable water-resource management in northeastern Morocco.

Supplementary Materials

The following supporting information can be downloaded at https://www.mdpi.com/article/10.3390/su182111003/s1, Table S1, taxonomic list; Table S2, records.

Author Contributions

Conceptualization, Y.M., A.M. and F.A.T.; methodology, Y.M. and F.A.T.; software, A.M. and F.A.T.; validation, formal analysis, investigation, Y.M. and F.A.T.; resources, data curation, Y.M. and F.A.T.; writing—original draft preparation, Y.M. and F.A.T.; writing—review and editing, visualization, supervision, Y.M. and A.M. All authors have read and agreed to the published version of the manuscript.

Funding

This research received no external funding.

Institutional Review Board Statement

Not applicable.

Data Availability Statement

The occurrence dataset supporting the findings of this study has been deposited in the GBIF network and is openly accessible through the following https://www.gbif.org/dataset/6b3f90ff-fc9a-4962-9605-8a79b81193e7 (accessed on 25 October 2026). Supplementary Table S1 contains the taxonomic checklist, and Supplementary Table S2 contains occurrence records, coordinates, dates, habitats, and sources.

Acknowledgments

We would like to express our sincere gratitude to the Editor and the Reviewers for their helpful and constructive comments, which have greatly improved the manuscript, as well as to Maria Bogomolova for her careful English revision.

Conflicts of Interest

The authors declare no conflicts of interest.

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