Biological invasions and their potential economic costs in Morocco

Biological invasions and their potential economic costs in Morocco | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Article Biological invasions and their potential economic costs in Morocco Jazila El Jamaai, Ahmed Taheri, Liliana Ballesteros-Mejia, Danish Ahmed, and 4 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-4731421/v1 This work is licensed under a CC BY 4.0 License Status: Published Journal Publication published 15 Jan, 2026 Read the published version in Scientific Reports → Version 1 posted 11 You are reading this latest preprint version Abstract Biological invasions pose substantial economic threats globally, yet detailed cost assessments for many Global South nations, especially in Africa, remain scarce. This study presents the first comprehensive breakdown of the potential costs of biological invasions in Morocco. We identified 551 invasive alien species, comprising approximately 1.76% of Morocco's biodiversity. Using the InvaCost database, we gathered cost data for the 12 most frequent invasive species with available data. Calculating the mean annual cost for each species and extrapolating based on their prevalence in Morocco, we estimated the potential annual economic impact to be US $ 1.61 billion. Urban species management and damage, especially pigeons, accounted for a substantial portion of this impact. Invasive plant species also emerged as prevalent and costly. Annual management costs exceeded damage costs (US $ 1.50 billion vs. 856.80 million), driven mainly by pigeon management. Costs by authorities and stakeholders outweighed agricultural costs (US $ 1.49 billion vs. 859.10 million). Despite challenges in extrapolating cost data from other regions, this study underscores the urgent need for targeted management and policy interventions to minimize the spread of invasive species and reduce their economic toll. Morocco can implement proactive management measures and foster international collaborations to tackle this socio-ecological crisis, ensuring long-term sustainability and prosperity. Invasive species Economic impacts Management strategies Morocco Biological invasions InvaCost Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 Introduction Globalization has led to unprecedented growth in the number, reach, and impact of biological invasions, with adverse effects on ecological and socioeconomic systems worldwide [ 1 , 2 ]. The increase in biological invasions is primarily due to the intensification of international trade and transport networks, resulting in the proliferation of pathways for the introduction and establishment of invasive alien species [ 3 , 4 , 5 ]. Biological invasions are considered as a major driver of global change, along with land-use and climate changes, which threaten ecosystems by altering biodiversity and functioning of ecosystems [ 6 , 7 ], resulting in severe negative impacts on ecosystem services, human health, food security, and national economies [ 1 ]. Despite the substantial threat, the ability to prevent and effectively mitigate biological invasions has remained inadequate in numerous countries [ 8 , 9 ]. For most taxonomic groups, the rate of first records has been steadily increasing annually, with no sign of saturation in the long term [ 10 ]. Future projections indicate that this upward trend will persist, and often accelerate, at least for the next three decades [ 11 ]. Countries with a high level of economic activity face increased vulnerability to harm from biological invasions, and more often document the presence of alien species [ 12 , 13 ]. However, they also possess greater potential to mitigate such damage, rendering them at lower risk than nations with more limited resources and lower import levels. Consequently, a nation's economic capability partially determines the efficacy of investing in the detection, control and prevention of invasive species. Developing economies such as those in the Africa continent, with fewer resources directed towards tackling ecological crises, will likely experience more dire impacts of biological invasions [ 14 , 15 ]. Intense research efforts in invasion science over the last few decades have provided a more comprehensive understanding of the environmental impacts of invasive alien species [ 16 ], but only recently have there been major developments in the evaluation of their economic impacts [ 17 , 18 ]. Invasive species are estimated to have cost more than US $ 423 billion annually globally and have been increasing four-fold every decade since the 1970s [ 9 ]. With much progress in cost collection and standardization, economic analyses have provided cost assessments at various geographical scales, for different countries and taxonomic groups, various habitats, and several economic sectors [ 18 , 19 , 20 , 21 , 22 , 23 ]. Moreover, numerous studies have shown that allocating greater investments toward prevention and other management measures during the initial stages of biological invasions increases the chances of attaining overall net benefits [ 24 , 25 , 26 , 27 , 28 ]. Given that prevention outweighs the efficacy of post-invasion mitigation and restoration, management strategies should prioritize early intervention in the invasion process [ 15 , 27 ]. Such an intervention would involve rapid response, risk assessments, managing pathways and vectors, and early detection [ 1 ]. Building global collaborations and evidence-based innovations for effectively mitigating the ecological and economic impacts of invasive species is a necessity. To that end, resources for minimizing the negative effects of invasives should be allocated based on comprehensive assessments of their distribution and economic impact, a practice currently deficient in most countries worldwide [ 29 ]. However, to fully realize the benefits of effective management, more effort must be made to improve data availability, particularly in emerging countries integrated into global trade [ 26 ]. Morocco, situated in the northwest corner of Africa, is just 15 kilometers from Europe across the Strait of Gibraltar. It boasts a rich history as an international commercial hub and is one of Europe’s largest trading partners in Africa. Indeed, it serves as a vital transportation node and commercial intermediary for numerous African nations, particularly those located in the hinterland. The country’s geographical diversity is evident in its extensive coastline, stretching 3500 kilometers along the Atlantic and the Mediterranean oceans. Morocco features three major topographical zones, each with its unique characteristics. The verdant plains in the north thrive as agricultural centers, while the Rif region in the extreme north showcases a blend of plains and mountains. Dominating the landscape, the Atlas Mountains serve as the country’s backbone. The climate across Morocco exhibits significant variation, transitioning from a Mediterranean climate in the northern coastal regions to mountainous areas with winter snow, and extending to extremely arid deserts in the south. This diversity in ecosystems, alongside high arrival rates of incoming alien species via trans-Mediterranean and trans-Atlantic routes of trade, as well as transport and tourism, makes the country increasingly susceptible to the introduction of many alien species, some of which may become invasive. However, there are currently no studies estimating the distribution and economic impacts of these species in Morocco, nor identifying the regions and sectors most adversely affected. From there, more research effort is required, given, for example, the presence of invasive species with considerable negative economic impacts [ 30 ], such as the tomato leafminer [ 31 ] and the silver nightshade, which pose a threat to various crops in Morocco [ 32 ]. Furthermore, the most recent evaluation of the cost of environmental degradation to Moroccan society, estimated at approximately US $ 3.9 billion or 3.5% of the country’s GDP in 2014, did not include an assessment of biological invasions [ 33 ]. Currently, the management of biological invasions in Morocco and the wider Arab Maghreb region primarily revolves around implementing restrictive measures, preventing introductions, and controlling and combating harmful exotic species. However, these efforts are across various legal texts from different institutional structures and enforced by various competent services, often lacking comprehensive knowledge of invasive species. Better coordination among these services is required, leading to less fragmented governance and more effective collaboration. Additionally, the scarcity of data on invasive species, including inventories, prioritization, introduction pathways, propagation dynamics, and their responses to environmental pressures, hampers integrated research efforts. As a result, the effective management of these species remains challenging, making it difficult to accurately estimate the economic and biodiversity costs [ 34 ]. The InvaCost database provides the most current, comprehensive, and standardized compilation of global economic costs linked to biological invasions [ 17 ]. Through various analyses, this database has facilitated descriptive studies on the economic impacts of invasive species across different regions and countries [ 35 , 36 , 37 , 38 , 39 ]. In particular, the costs of invasive species in Africa have been examined [ 14 ], revealing lower costs than in other continents such as North America [ 39 ], Europe [ 36 ], or Asia [ 38 ]. Also, there is a notable geographic imbalance in research efforts and financial resources, with a significant portion of African costs originating from South Africa [ 40 ]. Despite concerted efforts, the availability of cost data specific to Morocco within the InvaCost database remains limited, even following thorough searches in non-English languages [ 19 ]. Such unavailability of data underscores the need for compiling comprehensive lists of invasive species and mapping their distribution across Morocco to understand their potential economic impacts better. Here, we provide a projected cost assessment, drawing upon a global analysis of invasive species currently established within the country. Results Observed invasive alien species in Morocco After reviewing biodiversity databases, published papers, and other materials, we identified a total of 551 invasive species established in Morocco. Of these, 141 species were documented in the InvaCost database, whereas 410 lacked cost entries. Also, out of the total 551 species, only 188 had location data in the GBIF database (Table S1 ). Among the 141 invasive species with recorded costs, 107 had documented occurrences. These occurrences were primarily associated with vertebrates (76.0%, 25 species), plants (23.5%, 68 species), and arthropods (0.4%, 11 species). In contrast, among the 410 invasive species with no recorded costs in InvaCost , 81 had documented occurrences in Morocco. These occurrences were mainly comprised of vertebrates (90.1%, ten species), followed by plants (9.3%, 56 species), red algae (0.3%, two species), and arthropods (0.2%, ten species). The taxonomic distribution of these species, ranked by the number of locations in Morocco across both lists, is illustrated in Fig. 1 a. Considering the invasive alien species established in Morocco with over 50 recorded locations, vertebrates emerged as the predominant taxonomic group (Fig. 2 ). Among these, three avian species, namely the Eurasian collared-dove ( Streptopelia decaocto ), the cattle egret ( Bubulcus ibis ), and the pigeon ( Columba livia ), were observed at over 8,000 locations collectively, making them notably the most extensively distributed invasive alien species in the country. Additionally, two other bird species, the mallard ( Anas platyrhynchos ) and the purple swamphen ( Porphyrio porphyrio ), each exceeded 500 occurrences. Among these species, only the pigeon had documented economic costs (Fig. 2 a,b). Among the invasive alien plants, the oleander ( Nerium oleander ), tree tobacco ( Nicotiana glauca ), castor bean ( Ricinus communis ) and common storksbill ( Erodium cicutarium ) were notably widespread, each ranging between 200 and 350 locations in Morocco, all with reported costs outside Morocco (Fig. 2 c). Also, numerous other plant species had fewer than 200 locations, with most lacking reported costs in InvaCost (Fig. 2 d). Cost breakdowns by taxonomy, typology and impacted sector Taxonomic distribution of the costs Out of the 14 most widespread invasive species in Morocco that incur economic costs outside the country (Fig. 2 a,c), two had reported costs not only for the target species but also for multiple species collectively. This was observed for the false yellowhead ( Dittrichia viscosa ) and the feral cat ( Felis catus ), whose management costs were documented alongside those of similar species (in the case of the former, with only two cost records), or alongside all invasive species in the region (in the case of the latter, with 26 cost records). Despite their inclusion at the bottom of Table 1 , these two species were excluded from the subsequent calculations. Considering annual costs in InvaCost , the remaining 12 species accounted for 480 entries. The cumulative global economic cost for these species amounted to US $ 4.26 billion. The annual costs of these species in Morocco were estimated by multiplying the annual cost per site (or per country in the cases of country-level costs) by the number of locations in Morocco. Thus, we estimated a total economic impact of US $ 1.61 billion (Table 1 ). Table 1 Cost estimates in InvaCost for the 14 invasive alien species in Morocco, using the expanded dataset. Species are ordered by their total economic impact reported in InvaCost (US $ in 2017), except for two species with costs that include other species (the false yellowhead, Dittrichia viscosa, and the feral cat, Felis catus). We also provide the number of cost entries in InvaCost for each species, the type of costs (D: damage, M: management, DM: both), the spatial scale of the costs reported and the countries in which these costs incurred, and the annual costs estimated for Morocco. Annual costs for species within Morocco were estimated by multiplying the annual costs by the number of sites, for all species except those with country as spatial scale. Labels for the spatial scale: C: country level, S: site, within country level, R: river basin, Ha: hectare, G: global. Labels for the countries: Sp: Spain, Po: Portugal, Fr: France, UK: United Kingdom; Mx: Mexico; SA: South Africa; US: United States of America; Au: Australia. Costs estimated in Morocco are annual costs in US $ 2017; b: billion, m: million, t: thousand. Species Annual cost in InvaCost Number of entries Cost type Spatial Scale Country Cost in Morocco Columba livia 1 106 745 250.67 3 DM C US, UK 1.107b Solanum elaeagnifolium 456 122 075.40 2 D C Russia 456.12m Erodium cicutarium 96 165.07 12 D S Au 20.58m Arundo donax 65 126.88 229 M S, R, Ha Fr, Po, Mx, SA, Sp 4.77m Ricinus communis 58 091.49 44 M S SA, Sp 10.90m Arctotheca calendula 42 738.66 115 DM C, S, R Sp, Au 4.41m Oxalis pes-caprae 35 085.28 12 M S, G Sp, Po 5.72m Nicotiana glauca 1 105.68 28 M S Sp 323.96t Gambusia holbrooki 918.90 10 M S Sp 51.46t Nerium oleander 149.28 20 M S SA 50.16t Solanum linnaeanum 128.60 4 M S Sp 14.27t Schinus molle 29.37 1 M S Sp 3.64t Subtotal 4.256b 450 1.610b Felis catus 49 107.52 2 M S Sp 2.60m Dittrichia viscosa 6 258.37 26 M S Sp 657.13t Total 4.257b 495 1.613b Species with only country-level cost estimates emerged as the most financially burdensome when annual costs were calculated across various spatial scales. Notably, the pigeon ( Columba livia ), which experienced a relatively large range expansion in Morocco, incurred the highest economic toll. The reported expenses of pigeon invasions included two instances from the United States (management) and one from the United Kingdom (damage), all documented in the early 2000s. Similarly, the silverleaf nightshade ( Solanum elaeagnifolium ), for which country-level cost estimates were available in Russia, were extrapolated to represent potential damages should the species invade the country, likely contributing to the considerable costs reported. Furthermore, costs associated with capeweed ( Arctotheca calendula ) were reported across various spatial scales, including country (e.g., natural areas in Spain), site, and river levels. While expenses were relatively modest at the country level, ranging around US $ 2,173.11 annually (N = 11), costs were substantially higher at lower spatial scales in Spain and South Africa, reaching US $ 47,439.19 at the site level (N = 103) and US $ 4,805.56 at the river scale (N = 1). As a result, capeweed was classified as having costs predominantly at the site level (all costs were averaged independently of the spatial scale). While the pigeon and the silverleaf nightshade collectively incurred a projected annual cost exceeding US $ 1.5 billion in Morocco, the remaining invasive alien species accounted for an annual cost of US $ 49.91 million in the same region during the same period (Fig. 1 b) The lower expenses associated with the remaining species can be due to costs primarily comprising management expenses rather than direct damages. The taxonomic breakdown of estimated annual costs for Morocco is as follows: US $ 20.58 million from 12 management costs reported for the common storksbill ( Erodium cicutarium ), across three different states in Australia; US $ 10.90 million from 44 management costs reported for the castor bean ( Ricinus communis ) across various villages in Spain and Kruger National Park in South Africa; US $ 5.72 million from 12 management costs reported for the Bermuda buttercup ( Oxalis pes-caprae ) with expenditures on research projects conducted in Spain and Portugal; US $ 4.76 million from 229 management costs reported for the giant reed ( Arundo donax ), covering various regions in Spain, Portugal, and France, as well as specific protected areas in Mexico and South Africa; and US $ 4.41 million from 71 management costs reported for the capeweed ( Arctotheca calendula ), across different sites in Spain and various states in Australia, along with 44 cost entries accounting for damage costs across different states in Australia. The projected annual expenses for the remaining invasive species were less than US $ 1 million. Specifically, this included US $ 323.96 thousand coming from 28 management expenditures associated with the tree tobacco ( Nicotiana glauca ), across various locations in Spain; US $ 51.46 thousand from 10 management costs linked to the mosquitofish ( Gambusia holbrooki ), from different sites in Spain; and US $ 50.16 thousand from 20 management costs for the oleander ( Nerium oleander ), situated in the Kruger National Park in South Africa. The apple of Sodom ( Solanum linnaeanum ) with four management costs, and the false pepper tree ( Schinus molle ) with a single management cost reported, both at sites in Spain, represented the remaining cost entries totaling US $ 20.70 thousand and US $ 15.95 thousand. Management and damage costs In terms of cost type (either damage or management), only four species have reported damage costs: pigeons ( Columba livia , N = 1), capeweed ( Arctotheca calendula , N = 44), the silverleaf nightshade ( Solanum elaeagnifolium , N = 2) and the common storksbill ( Erodium cicutarium , N = 12). By extrapolating, we estimate an annual damage cost of US $ 856.80 million in Morocco, presuming that all occurrences of pigeons and capeweed resulted in direct damages. Based on the annual management costs for pigeons and capeweed, we estimate a potential annual expenditure amounting to US $ 1.5 billion (Fig. 3 ). Notably, projected management expenses surpass damage costs, and this difference is mainly influenced by the reported costs associated with pigeons. If we remove pigeons, damage costs surpass management costs (US $ 483.03m vs US $ 27.09m). Sectoral distribution of cost All damage costs were reported for agriculture, except for pigeons, where agriculture and public/social welfare were jointly reported. Regarding management costs, the majority were borne by the authorities and/or stakeholders, including governmental services and official organizations such as conservation agencies, forest services, or associations involved in the broad management of biological invasions, e.g., control programs, eradication campaigns, research funding (Table S2). Two exceptions were noted, capeweed ( Arctotheca calendula ) and giant reed ( Arundo donax ), which also reported management costs in agriculture (Fig. 3 b). On assessing the annual costs per site for the five invasive species affecting the agricultural sector outside of Morocco, and extrapolating these figures based on occurrences in Morocco, the estimated annual cost impacting agriculture totaled US $ 859.10 million. Similarly, in this context, the economic impact on authorities and stakeholders amounted to US $ 1.49 billion annually. Geographic distribution of cost The geographic distribution of costs is dependent on the locations of the occurrences of the twelve invasive alien species considered (Fig. 4 ). Besides, it is also dependent on the scale of the cost that these species have. Summing the extrapolated annual costs per site for all the species, we observe that the potential economic costs in Morocco are unevenly distributed across the country. Higher costs are concentrated in more densely populated regions, such as Casablanca, Rabat, and Marrakech, as well as along the Atlantic coast, including Casablanca and Rabat. Discussion This study presents the first comprehensive global analysis of invasive species in Morocco, offering insights into the potential economic costs associated with biological invasions in the country and the geographic distribution of the estimated costs. Urgent research was prompted by the necessity to identify invasive alien species and map their distribution across Morocco, an essential step for assessing their economic impacts. A total of 551 invasive alien species were identified, representing 1.76% of the nation's plant and animal biodiversity. The absence of cost data for Morocco in the InvaCost database highlighted the need for promoting cost reporting, and based on extrapolation of costs from other countries, we estimated a potential annual cost amounting to at least US $ 1.61 billion. Compared to the estimated annual mean cost for the entire African continent, estimated between US $ 2.6 and US $ 8.6 billion in 2019 [ 1 , 2 ], the potential economic toll of invasive species in Morocco is, therefore, substantial. However, this estimate is likely conservative, as they were predominantly based on data from a single country, with South Africa accounting for approximately 86% of the total number of cost entries [ 14 ]. Besides, Eschen et al. [ 43 ] projected the annual cost of invasive alien species in the Moroccan agricultural sector alone to be US $ 1.66 billion, drawn from literature data and stakeholder interviews. Another example of the underestimation of the costs from Africa generally, is the descriptions of the costs caused by biological invasions in the Mediterranean basin, which were available for only 15 out of 26 countries, with most reports arising from Western European nations, i.e., France, Spain, and Italy, and only 11 records from African countries such as Libya (US $ 593,04 million, N = 8) and Egypt (US $ 147.16 million, N = 3) [ 35 ]. While both studies relied on the InvaCost database, note that none of them incorporated cost data for Morocco. Thus, using known cost estimates for documented species appears to be the only practical and straightforward approach for estimating missing costs in other countries affected by invasions [ 44 ]. We estimated a potential annual cost of US $ 1.61 billion in Morocco, and as a first assessment for the country, it should help develop effective national policies and global management initiatives to tackle this issue. Our assessment is essential for informed decision-making at both national and international levels. Birds are the most prevalent invasive alien species nationwide; however, only the pigeon ( C. livia ) has recorded economic costs. The lack of cost data for most exotic bird species may stem from their occurrence in regions with limited resources for researching their impacts or in remote areas where tracking their effects proves challenging [ 29 ]. Alternatively, it could be attributed to research prioritizing well-known taxa [ 1 , 2 ] or the most threatening alien species [ 36 , 46 ], given that exotic birds often have minimal environmental and socioeconomic impacts [ 47 , 48 ]. Despite this, pigeons accounted for over half of all exotic bird costs globally [ 49 ], reflecting their significant economic burden - with an estimated cost of US $ 1.11 billion in Morocco. In contrast, invasive alien plants such as the oleander ( N. oleander ), tree tobacco ( N. glauca ), castor bean ( R. communis ), and common storksbill ( E. cicutarium ), were the most widespread, each observed ranging between 200 and 350 locations across Morocco. While some authors consider oleander and the common storksbill to be native to Morocco [ 50 ], projected costs for those species are still very conservative. The tree tobacco is native to northwest Bolivia and Argentina, and is recognized as an invasive plant in Morocco [ 51 ], whereas the castor bean is native to Northeastern Africa; both probably introduced before 1931 [ 52 ]. Notably, the silverleaf nightshade, a global threat and the second most expensive plant species in Morocco, is estimated to cost approximately US $ 456.12 million. This plant, introduced unintentionally around 1949, profoundly impacts agriculture, biodiversity, and the environment. For example, losses of up to 64% in maize production have been recorded in Morocco [ 53 ]. While costs for the silverleaf nightshade were not directly assessed in Morocco, estimates were derived from economic costs reported in Russia, with extrapolations made to predict potential damages if the species spreads there [ 54 ]. Even though it is challenging to standardize costs across different economies, using exchange rates provides a practical method, albeit with limitations due to the lack of data on Purchasing Power Parity (PPP). While the pigeon and silverleaf nightshade incurred a combined annual cost surpassing US $ 1.5 billion, the other invasive alien species accrued an estimated annual cost of US $ 49.91 million. The comparatively lower costs associated with the remaining species can be attributed to the fact that most of these expenses are primarily related to management rather than direct damages. This aligns with earlier findings regarding the costs of invasive alien species, where reported management expenses typically fall well below reported damages [ 28 ]. It must be considered that the costs for Morocco are estimated costs using the costs from other countries. Thus, whether costs are typified as management or damage corresponds to the availability of costs in InvaCost for the species invading Morocco. Thus, as many of the costs come from Spain (many of the invasive species are the same, very likely because of the proximity), where management costs surpass damages [ 19 ], this could be one of the reasons why most invasive species have management costs available for Morocco. Given that at the global scale, management costs are about ten times lower than damage costs [ 28 , 29 ], the actual costs for Morocco might be much higher than estimated here. The uneven geographic distribution of potential economic costs from invasive species in Morocco can be attributed to several factors. Firstly, more densely populated regions like Casablanca, Rabat, and Marrakech have higher economic activity, infrastructure, and assets at risk, leading to greater potential costs when invasive species cause damage. These urban centers serve as major hubs of human and commercial activity, facilitating the introduction and proliferation of invasive species through increased transportation, trade, and tourism. Secondly, the Atlantic coast, where Casablanca and Rabat are located, features ecosystems that are particularly vulnerable to certain invasive species (Fig. 4 ), which can thrive in these environments and cause significant economic harm. For example, Solanum elaeagnifolium was introduced to Morocco in the mid-20th century, first establishing itself in the Tadla region (Beni Mellal-Khenifra region) [ 55 ] and spreading widely by the 1990s. The estimated potential cost incurred thus far stands at US $ 0.45 billion annually (Fig. 4 ), emphasising the significant economic consequences of its spread. Currently, all six geographical zones in Morocco are infested, with the least impact in the Saharan and mountainous areas. Around 70% of infestation sites are in major agricultural districts, and more than 15% along the Rabat-Casablanca axis. Moreover, this species is rapidly adapting and spreading to new regions [ 56 , 57 , 58 ]. Coastal regions often host diverse and economically significant industries such as fisheries, tourism, and agriculture that can be severely impacted by invasive species, thereby driving up the costs, such as the Marrakech-Safi region, with an estimated potential cost of US $ 361.62 million (Fig. 5 ). As the primary tourist destination in the kingdom, Marrakech is home to Morocco’s busiest airport, which served over 6.3 million passengers in 2019, and includes Safi, the country’s oldest port. Additionally, the region accounts for 22% of Morocco’s useful agricultural area. Lastly, urban areas and economically vibrant regions tend to have more comprehensive monitoring and reporting systems in place, which can result in better documentation and more frequent records of invasive species presence, compared to less populated and economically active areas. Given that the failure to act on invasive species management can lead to substantial avoidable costs [ 27 ], Morocco must establish proactive measures aimed at preventing the introduction of new invasive alien species. To achieve this, future studies should focus on identifying and cataloging these potential invasive species. With a better understanding of which species pose a threat and early detection of possible introduction pathways, the country will be better positioned to enable policymakers and environmental managers to develop targeted strategies for interception and control. By taking timely action, Morocco can effectively reduce the economic costs linked to the impacts of invasive species in the future. On synthesis, although the estimated annual economic impact of invasive species in Morocco is US $ 1.61 billion, this figure likely underestimates the actual economic burden for several reasons: it is derived from data on only 2% of the invasive species present in Morocco, there is a lack of Morocco-specific cost data, and many invasive species worldwide have relatively low reported costs, including the 12 invasive alien species for which we found cost data. The disproportionate focus on specific taxa, such as pigeons, highlights the need for a more comprehensive approach to invasive species management that considers the full spectrum of ecological and economic impacts. Improved cost reporting, early detection, and effective management strategies are essential for minimizing the economic and environmental consequences of biological invasions. Enhanced collaboration between researchers, policymakers, and stakeholders is necessary to develop targeted laws and interventions, and allocate resources efficiently. By addressing these challenges, we can better protect ecosystems, safeguard biodiversity, and mitigate the economic risks of invasive species in Morocco and the wider region. Methods Data collection, compilation and filtering To assess the economic costs associated with invasive alien species in Morocco, we first identified existing species within the country using comprehensive databases such as GISD, GRIIS, and the ‘Standardising and Integrating Alien Species workflow’ [ 41 ]. Additionally, we utilized resource lists such as ‘Terrestrial Invasive Alien Species in the Arab Maghreb’ and ‘Elements for Reflection on Primary Invasive Alien Species in Morocco’. Supplementary systematic searches in online repositories (Web of Science, Google Scholar and Google search engine), and gray literature were performed. We further engaged with national experts and stakeholders to gather additional information, e.g., official national managers or researchers that could provide cost data. We removed duplicates and considered only those that were taxonomically ranked at the species level (i.e., discarding cases with only genus information). We then cross-checked the InvaCost v4.1 database [ 17 ] ( https://doi.org/10.6084/m9.figshare.12668570.v5 ) to determine whether any of these species incurred economic costs somewhere in the world. Although InvaCost compiles cost information obtained from non-English sources, including French and Arabic [ 19 ], the database still did not feature any reported costs for Morocco. Comparing the list of invasive alien species detected in Morocco with the invasive species having costs in InvaCost , we separated this information into two categories: invasive alien species detected in Morocco lacking cost information in InvaCost , and species observed in Morocco with documented costs in InvaCost , but arising from cost occurrences outside of Morocco (Table S1 ). Occurrence records detailing their current invasive distribution in the country were retrieved from the Global Biodiversity Information Facility (GBIF; https://www.gbif.org ) for all species listed in both datasets. In order to have a reliable dataset, we removed duplicated geographic coordinates, occurrences without GPS coordinates, and those with erroneous coordinates falling outside terrestrial borders that could not be georeferenced from the information provided. Taxonomic descriptions were provided for species in both lists, and those with more than 50 occurrences (N = 14 species) were selected. Estimating costs of invasive alien species in Morocco By filtering the ‘Species’ column in the InvaCost database, we obtained the reported costs for the 14 invasive alien species. We considered only high-reliability cost entries from the ‘Reliability’ column, i.e., cost estimates sourced from official peer-reviewed material and/or material with reproducible methods [ 17 ]. Costs were extracted from the ‘Cost_estimate_per_year_2017_USD_exchange_rate’ column, such that all cost entries were adjusted to a common currency and year for comparability, i.e., in 2017 US dollars. This adjustment also factored in inflation, referencing the Consumer Price Index of 2017 in relation to the year of the cost estimation. Each entry also includes the timespan linked with the recorded costs, facilitated by the expandYearlyCosts function from the invacost R package [ 42 ]. This function distributes the total reported cost evenly across probable start and end years, generating an expanded dataset where each entry represents a cost estimate for a specific year. Within the InvaCost database, each publication served as a distinct point of reference for reported costs, although the duration over which costs were estimated differed among references. Complete details regarding the compilation and standardization of data stored in InvaCost are provided in Diagne et al. [ 17 ]. First, we calculated the total economic cost reported in InvaCost for each of these 14 IAS. Then, to ascertain a relative annual cost per species, we calculated the annual amount per spatial unit, whether it be as large as a country or as small as a local site, utilizing the ‘Spatial_scale’ column. A site refers to an area at the intra-country level, while a unit denotes a well-defined surface area or entity, such as a hectare or a river basin. This adjustment was necessary due to the non-uniformity in the number of reported costs and the spatial scale of these costs in InvaCost across various species. We took into account the costs incurred by different impacted sectors, as indicated in the ‘Impacted_sector’ column, and the cost typology (i.e., management and damages), as specified in the ‘Type_of_cost_merged’ column. We then extrapolated the economic costs accounting for the distribution of each species, by multiplying the annual cost per site, obtained from the reported data in InvaCost , by the number of occurrences sourced from GBIF. An estimate of the total cost incurred by invasive species within Morocco was computed as the sum of extrapolated annual costs per site for all species. Finally, we created a map that included the sum of the extrapolated annual costs per site for all the species in order to understand the spatial distribution of potential economic costs in Morocco and highlight the places in the country that are predicted to cause the highest economic costs of invasive alien species. For the two species that have costs at the country level ( C. livia and S. elaeagnifolium ), we first calculated a cost per site by dividing the country cost by the number of occurrences in Morocco. Declarations Data availability Statement Data supporting the results reported in this article can be found in: Global Invasive Species Database (GISD) https://www.iucngisd.org/gisd/ Global Register of Introduced and Invasive Species (GRIIS) https://griis.org/ Standardising and Integrating Alien Species workflow (SInAS) https://doi.org/10.3897/neobiota.59.53578 Terrestrial Invasive Alien Species in the Arab Maghreb https://www.cbd.int/invasive/doc/arab-maghreb-eee-fr.pdf Elements for Reflection on Primary Invasive Alien Species in Morocco https://doi.org/10.1002/9781119607045.ch7 InvaCost database https://doi.org/10.6084/m9.figshare.12668570.v5 Global Biodiversity Information Facility (GBIF) https://www.gbif.org/fr/ Acknowledgments We would like to thank the Direction of Climate and the Biological Diversity, Ministry of Energy Transition and Sustainable Development, Morocco Author contributions: EA and AT contributed to the study conception and design. Data collection was performed by CD, EA, FC, and LBM, the methodology was agreed by JEJ, LBM, EA and AT and analysis were performed by JEJ, CD, EA and LBM. Supervision was carried out by EA and AT and funding was spent by FC, EA and DA. The first draft of the manuscript was written by JEJ and visualization items were created by JEJ, AT and EA. All authors commented on previous versions of the manuscript and all authors read and approved the final manuscript. Additional Information Funding Funding was partially given to Elena Angulo by Junta de Andalucía, Consejería de Universidad, Investigación e Innovación, PROYEXCEL_00688 within the PAIDI 2020. The InvaCost project was funded by the French National Research Agency (ANR-14-CE02-0021), the BNP-Paribas Foundation Climate Initiative, the AXA Research Fund Chair of Invasion Biology of University Paris Saclay and by the BiodivERsA and Belmont-Forum call 2018 on biodiversity scenarios (AlienScenarios; BMBF/PT DLR 01LC1807C). 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Supplementary Files Supplementarymaterial.docx Cite Share Download PDF Status: Published Journal Publication published 15 Jan, 2026 Read the published version in Scientific Reports → Version 1 posted Editorial decision: Revision requested 17 Apr, 2025 Reviews received at journal 27 Mar, 2025 Reviews received at journal 23 Mar, 2025 Reviewers agreed at journal 23 Mar, 2025 Reviewers agreed at journal 20 Mar, 2025 Reviewers agreed at journal 19 Mar, 2025 Reviewers invited by journal 18 Mar, 2025 Editor assigned by journal 24 Oct, 2024 Editor invited by journal 13 Aug, 2024 Submission checks completed at journal 08 Aug, 2024 First submitted to journal 12 Jul, 2024 You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. As a division of Research Square Company, we’re committed to making research communication faster, fairer, and more useful. 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Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-4731421","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Article","associatedPublications":[],"authors":[{"id":348361632,"identity":"44729f5e-b82a-4ecd-acf5-8a86b8a04631","order_by":0,"name":"Jazila El Jamaai","email":"data:image/png;base64,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","orcid":"","institution":"Chouaïb Doukkali University","correspondingAuthor":true,"prefix":"","firstName":"Jazila","middleName":"El","lastName":"Jamaai","suffix":""},{"id":348361633,"identity":"be594010-6993-4368-869a-7a332786407e","order_by":1,"name":"Ahmed Taheri","email":"","orcid":"","institution":"Chouaïb Doukkali University","correspondingAuthor":false,"prefix":"","firstName":"Ahmed","middleName":"","lastName":"Taheri","suffix":""},{"id":348361634,"identity":"fbf3a28a-483f-48cb-b322-4adb8007e78c","order_by":2,"name":"Liliana Ballesteros-Mejia","email":"","orcid":"","institution":"Muséum d’Histoire Naturelle, CNRS, Université des Antilles","correspondingAuthor":false,"prefix":"","firstName":"Liliana","middleName":"","lastName":"Ballesteros-Mejia","suffix":""},{"id":348361635,"identity":"66c76551-9d1d-4c51-b079-a0883b677a16","order_by":3,"name":"Danish Ahmed","email":"","orcid":"","institution":"CAMB, Gulf University for Science and Technology","correspondingAuthor":false,"prefix":"","firstName":"Danish","middleName":"","lastName":"Ahmed","suffix":""},{"id":348361636,"identity":"4f5373d5-dce0-45c4-82ed-a46e5f60de86","order_by":4,"name":"Alok Bang","email":"","orcid":"","institution":"Azim Premji University","correspondingAuthor":false,"prefix":"","firstName":"Alok","middleName":"","lastName":"Bang","suffix":""},{"id":348361637,"identity":"6fcdc206-a91b-4e4b-8d89-2cd560783f38","order_by":5,"name":"Christophe Diagne","email":"","orcid":"","institution":"CBGP, IRD, Univ Montpellier, CIRAD, INRAE, Institut Agro","correspondingAuthor":false,"prefix":"","firstName":"Christophe","middleName":"","lastName":"Diagne","suffix":""},{"id":348361638,"identity":"2a3deee4-2b19-4d70-b576-ddbfed8beb15","order_by":6,"name":"Franck Courchamp","email":"","orcid":"","institution":"CNRS, Université Paris-Saclay","correspondingAuthor":false,"prefix":"","firstName":"Franck","middleName":"","lastName":"Courchamp","suffix":""},{"id":348361639,"identity":"b0ce6eb3-dcb1-4c2c-ad9d-063ec1a3ad02","order_by":7,"name":"Elena Angulo","email":"","orcid":"","institution":"Estación Biológica de Doñana (CSIC)","correspondingAuthor":false,"prefix":"","firstName":"Elena","middleName":"","lastName":"Angulo","suffix":""}],"badges":[],"createdAt":"2024-07-12 15:53:29","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-4731421/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-4731421/v1","draftVersion":[],"editorialEvents":[{"content":"https://doi.org/10.1038/s41598-025-31767-8","type":"published","date":"2026-01-15T16:30:33+00:00"}],"editorialNote":"","failedWorkflow":false,"files":[{"id":64165690,"identity":"d0f2cc36-a032-4d0e-842d-2239c78faaec","added_by":"auto","created_at":"2024-09-09 09:16:27","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":419453,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003eThe number of locations of invasive alien species in Morocco\u003c/strong\u003e is classified based on whether they have costs listed in InvaCost outside Morocco or lack such costs. The distribution of these locations is categorized taxonomically by their phylum (a), and, the distribution of estimated costs of invasive alien species in Morocco classified taxonomically by phylum and species (b)\u003c/p\u003e\n\u003cp\u003eDiagram created using SankeyMATIC (https://sankeymatic.com/)\u003c/p\u003e","description":"","filename":"1.png","url":"https://assets-eu.researchsquare.com/files/rs-4731421/v1/fd2e65a0804fc48de7f9de0f.png"},{"id":64165691,"identity":"a137e88c-13c1-43d3-9f01-5f46bb4c65b6","added_by":"auto","created_at":"2024-09-09 09:16:27","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":466294,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003eThe number of locations for the invasive alien vertebrate species with more than 50 occurrences in Morocco\u003c/strong\u003e categorized by whether they incur costs in InvaCost (a) or not (b), alongside the count of locations for invasive alien plant species with over 50 occurrences in Morocco, incurring costs (c) or not (d)\u003c/p\u003e","description":"","filename":"2.png","url":"https://assets-eu.researchsquare.com/files/rs-4731421/v1/8b9914413619de0346590f2d.png"},{"id":64165693,"identity":"73c59e63-dba0-44fd-9a1e-3184432fc2bf","added_by":"auto","created_at":"2024-09-09 09:16:27","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":822678,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003eEstimated costs associated with the 12 identified invasive alien species in Morocco\u003c/strong\u003e, categorized by type of cost (red and purple) and affected economic sector (green and yellow)\u003c/p\u003e\n\u003cp\u003eDiagram created using R version 4.4.0 (https://cran.r-project.org/bin/windows/base/)\u003c/p\u003e","description":"","filename":"3.png","url":"https://assets-eu.researchsquare.com/files/rs-4731421/v1/f2e4966d8f4207e462f35fa4.png"},{"id":64166252,"identity":"8661e835-c5fa-46f5-98ac-d61ddfd18a9c","added_by":"auto","created_at":"2024-09-09 09:32:27","extension":"png","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":2154214,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003eOccurrences of the twelve invasive alien species having reported economic costs and their costs in Dírham (MAD).\u003c/strong\u003e The volume of the yellow circle represents the relative amount of their costs on a log scale. The species are ordered by the number of occurrences retrieved from the Global Biodiversity Information Facility (https://www.gbif.org). Photos obtained in iNaturalist with creative commons license.\u003c/p\u003e\n\u003cp\u003eMaps created using Arcgis 10.8 (https://desktop.arcgis.com)\u003c/p\u003e","description":"","filename":"4.png","url":"https://assets-eu.researchsquare.com/files/rs-4731421/v1/bdc813c0f66afbe8b5915d4a.png"},{"id":64166034,"identity":"0fec94ac-4775-4d82-9779-6382a4b9800f","added_by":"auto","created_at":"2024-09-09 09:24:27","extension":"png","order_by":5,"title":"Figure 5","display":"","copyAsset":false,"role":"figure","size":832654,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003eGeographic distribution of estimated costs in Morocco.\u003c/strong\u003eAbbreviations of the regions on the map are: BK Beni Mellal-Khenifra; CS Casablanca-Settat; DO Dakhla-Oued Eddahab; DT Draa-Tafilalet; FM Fez-Meknes; GO Guelmim-Oued Noun; LS Laayoune-Sakia El Hamra; MS Marrakech-Safi; OR Oriental; RK Rabat-Sale-Kenitra; SM Souss-Massa; TTA Tangier-Tetouan-Al Hoceima\u003c/p\u003e\n\u003cp\u003eMaps created using Arcgis Desktop 10.8 (https://desktop.arcgis.com)\u003c/p\u003e","description":"","filename":"5.png","url":"https://assets-eu.researchsquare.com/files/rs-4731421/v1/42e4c0143b8e2668fb950486.png"},{"id":100616350,"identity":"3019e035-508d-435d-ac15-ca0cbed79585","added_by":"auto","created_at":"2026-01-19 17:42:26","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":7063493,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-4731421/v1/acf4f8b4-f600-47b7-893c-edadc15939b3.pdf"},{"id":64165695,"identity":"5b1a4f60-fb71-4253-845c-075d8160485b","added_by":"auto","created_at":"2024-09-09 09:16:27","extension":"docx","order_by":2,"title":"","display":"","copyAsset":false,"role":"supplement","size":15423,"visible":true,"origin":"","legend":"","description":"","filename":"Supplementarymaterial.docx","url":"https://assets-eu.researchsquare.com/files/rs-4731421/v1/0bcd40dc840d98bcd7763c14.docx"}],"financialInterests":"No competing interests reported.","formattedTitle":"Biological invasions and their potential economic costs in Morocco","fulltext":[{"header":"Introduction","content":"\u003cp\u003eGlobalization has led to unprecedented growth in the number, reach, and impact of biological invasions, with adverse effects on ecological and socioeconomic systems worldwide [\u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e1\u003c/span\u003e, \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2\u003c/span\u003e]. The increase in biological invasions is primarily due to the intensification of international trade and transport networks, resulting in the proliferation of pathways for the introduction and establishment of invasive alien species [\u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e3\u003c/span\u003e, \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e4\u003c/span\u003e, \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e5\u003c/span\u003e]. Biological invasions are considered as a major driver of global change, along with land-use and climate changes, which threaten ecosystems by altering biodiversity and functioning of ecosystems [\u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e6\u003c/span\u003e, \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e7\u003c/span\u003e], resulting in severe negative impacts on ecosystem services, human health, food security, and national economies [\u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e1\u003c/span\u003e]. Despite the substantial threat, the ability to prevent and effectively mitigate biological invasions has remained inadequate in numerous countries [\u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e8\u003c/span\u003e, \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e9\u003c/span\u003e]. For most taxonomic groups, the rate of first records has been steadily increasing annually, with no sign of saturation in the long term [\u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e10\u003c/span\u003e]. Future projections indicate that this upward trend will persist, and often accelerate, at least for the next three decades [\u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e11\u003c/span\u003e]. Countries with a high level of economic activity face increased vulnerability to harm from biological invasions, and more often document the presence of alien species [\u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e12\u003c/span\u003e, \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e13\u003c/span\u003e]. However, they also possess greater potential to mitigate such damage, rendering them at lower risk than nations with more limited resources and lower import levels. Consequently, a nation's economic capability partially determines the efficacy of investing in the detection, control and prevention of invasive species. Developing economies such as those in the Africa continent, with fewer resources directed towards tackling ecological crises, will likely experience more dire impacts of biological invasions [\u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e14\u003c/span\u003e, \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e15\u003c/span\u003e].\u003c/p\u003e \u003cp\u003eIntense research efforts in invasion science over the last few decades have provided a more comprehensive understanding of the environmental impacts of invasive alien species [\u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e16\u003c/span\u003e], but only recently have there been major developments in the evaluation of their economic impacts [\u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e17\u003c/span\u003e, \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e18\u003c/span\u003e]. Invasive species are estimated to have cost more than US \u003cspan\u003e$\u003c/span\u003e423\u0026nbsp;billion annually globally and have been increasing four-fold every decade since the 1970s [\u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e9\u003c/span\u003e]. With much progress in cost collection and standardization, economic analyses have provided cost assessments at various geographical scales, for different countries and taxonomic groups, various habitats, and several economic sectors [\u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e18\u003c/span\u003e, \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e19\u003c/span\u003e, \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e20\u003c/span\u003e, \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e21\u003c/span\u003e, \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e22\u003c/span\u003e, \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e23\u003c/span\u003e]. Moreover, numerous studies have shown that allocating greater investments toward prevention and other management measures during the initial stages of biological invasions increases the chances of attaining overall net benefits [\u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e24\u003c/span\u003e, \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e25\u003c/span\u003e, \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e26\u003c/span\u003e, \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e27\u003c/span\u003e, \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e28\u003c/span\u003e]. Given that prevention outweighs the efficacy of post-invasion mitigation and restoration, management strategies should prioritize early intervention in the invasion process [\u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e15\u003c/span\u003e, \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e27\u003c/span\u003e]. Such an intervention would involve rapid response, risk assessments, managing pathways and vectors, and early detection [\u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e1\u003c/span\u003e]. Building global collaborations and evidence-based innovations for effectively mitigating the ecological and economic impacts of invasive species is a necessity. To that end, resources for minimizing the negative effects of invasives should be allocated based on comprehensive assessments of their distribution and economic impact, a practice currently deficient in most countries worldwide [\u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e29\u003c/span\u003e]. However, to fully realize the benefits of effective management, more effort must be made to improve data availability, particularly in emerging countries integrated into global trade [\u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e26\u003c/span\u003e].\u003c/p\u003e \u003cp\u003eMorocco, situated in the northwest corner of Africa, is just 15 kilometers from Europe across the Strait of Gibraltar. It boasts a rich history as an international commercial hub and is one of Europe\u0026rsquo;s largest trading partners in Africa. Indeed, it serves as a vital transportation node and commercial intermediary for numerous African nations, particularly those located in the hinterland. The country\u0026rsquo;s geographical diversity is evident in its extensive coastline, stretching 3500 kilometers along the Atlantic and the Mediterranean oceans. Morocco features three major topographical zones, each with its unique characteristics. The verdant plains in the north thrive as agricultural centers, while the Rif region in the extreme north showcases a blend of plains and mountains. Dominating the landscape, the Atlas Mountains serve as the country\u0026rsquo;s backbone. The climate across Morocco exhibits significant variation, transitioning from a Mediterranean climate in the northern coastal regions to mountainous areas with winter snow, and extending to extremely arid deserts in the south. This diversity in ecosystems, alongside high arrival rates of incoming alien species via trans-Mediterranean and trans-Atlantic routes of trade, as well as transport and tourism, makes the country increasingly susceptible to the introduction of many alien species, some of which may become invasive. However, there are currently no studies estimating the distribution and economic impacts of these species in Morocco, nor identifying the regions and sectors most adversely affected. From there, more research effort is required, given, for example, the presence of invasive species with considerable negative economic impacts [\u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e30\u003c/span\u003e], such as the tomato leafminer [\u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e31\u003c/span\u003e] and the silver nightshade, which pose a threat to various crops in Morocco [\u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e32\u003c/span\u003e]. Furthermore, the most recent evaluation of the cost of environmental degradation to Moroccan society, estimated at approximately US\u003cspan\u003e$\u003c/span\u003e 3.9\u0026nbsp;billion or 3.5% of the country\u0026rsquo;s GDP in 2014, did not include an assessment of biological invasions [\u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e33\u003c/span\u003e].\u003c/p\u003e \u003cp\u003eCurrently, the management of biological invasions in Morocco and the wider Arab Maghreb region primarily revolves around implementing restrictive measures, preventing introductions, and controlling and combating harmful exotic species. However, these efforts are across various legal texts from different institutional structures and enforced by various competent services, often lacking comprehensive knowledge of invasive species. Better coordination among these services is required, leading to less fragmented governance and more effective collaboration. Additionally, the scarcity of data on invasive species, including inventories, prioritization, introduction pathways, propagation dynamics, and their responses to environmental pressures, hampers integrated research efforts. As a result, the effective management of these species remains challenging, making it difficult to accurately estimate the economic and biodiversity costs [\u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e34\u003c/span\u003e].\u003c/p\u003e \u003cp\u003eThe \u003cem\u003eInvaCost\u003c/em\u003e database provides the most current, comprehensive, and standardized compilation of global economic costs linked to biological invasions [\u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e17\u003c/span\u003e]. Through various analyses, this database has facilitated descriptive studies on the economic impacts of invasive species across different regions and countries [\u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e35\u003c/span\u003e, \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e36\u003c/span\u003e, \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e37\u003c/span\u003e, \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e38\u003c/span\u003e, \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e39\u003c/span\u003e]. In particular, the costs of invasive species in Africa have been examined [\u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e14\u003c/span\u003e], revealing lower costs than in other continents such as North America [\u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e39\u003c/span\u003e], Europe [\u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e36\u003c/span\u003e], or Asia [\u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e38\u003c/span\u003e]. Also, there is a notable geographic imbalance in research efforts and financial resources, with a significant portion of African costs originating from South Africa [\u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e40\u003c/span\u003e]. Despite concerted efforts, the availability of cost data specific to Morocco within the \u003cem\u003eInvaCost\u003c/em\u003e database remains limited, even following thorough searches in non-English languages [\u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e19\u003c/span\u003e]. Such unavailability of data underscores the need for compiling comprehensive lists of invasive species and mapping their distribution across Morocco to understand their potential economic impacts better. Here, we provide a projected cost assessment, drawing upon a global analysis of invasive species currently established within the country.\u003c/p\u003e"},{"header":"Results","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e \u003ch2\u003eObserved invasive alien species in Morocco\u003c/h2\u003e \u003cp\u003eAfter reviewing biodiversity databases, published papers, and other materials, we identified a total of 551 invasive species established in Morocco. Of these, 141 species were documented in the \u003cem\u003eInvaCost\u003c/em\u003e database, whereas 410 lacked cost entries. Also, out of the total 551 species, only 188 had location data in the GBIF database (Table \u003cspan refid=\"MOESM1\" class=\"InternalRef\"\u003eS1\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eAmong the 141 invasive species with recorded costs, 107 had documented occurrences. These occurrences were primarily associated with vertebrates (76.0%, 25 species), plants (23.5%, 68 species), and arthropods (0.4%, 11 species). In contrast, among the 410 invasive species with no recorded costs in \u003cem\u003eInvaCost\u003c/em\u003e, 81 had documented occurrences in Morocco. These occurrences were mainly comprised of vertebrates (90.1%, ten species), followed by plants (9.3%, 56 species), red algae (0.3%, two species), and arthropods (0.2%, ten species). The taxonomic distribution of these species, ranked by the number of locations in Morocco across both lists, is illustrated in Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003ea.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eConsidering the invasive alien species established in Morocco with over 50 recorded locations, vertebrates emerged as the predominant taxonomic group (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). Among these, three avian species, namely the Eurasian collared-dove (\u003cem\u003eStreptopelia decaocto\u003c/em\u003e), the cattle egret (\u003cem\u003eBubulcus ibis\u003c/em\u003e), and the pigeon (\u003cem\u003eColumba livia\u003c/em\u003e), were observed at over 8,000 locations collectively, making them notably the most extensively distributed invasive alien species in the country. Additionally, two other bird species, the mallard (\u003cem\u003eAnas platyrhynchos\u003c/em\u003e) and the purple swamphen (\u003cem\u003ePorphyrio porphyrio\u003c/em\u003e), each exceeded 500 occurrences. Among these species, only the pigeon had documented economic costs (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003ea,b).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eAmong the invasive alien plants, the oleander (\u003cem\u003eNerium oleander\u003c/em\u003e), tree tobacco (\u003cem\u003eNicotiana glauca\u003c/em\u003e), castor bean (\u003cem\u003eRicinus communis\u003c/em\u003e) and common storksbill (\u003cem\u003eErodium cicutarium\u003c/em\u003e) were notably widespread, each ranging between 200 and 350 locations in Morocco, all with reported costs outside Morocco (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003ec). Also, numerous other plant species had fewer than 200 locations, with most lacking reported costs in \u003cem\u003eInvaCost\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003ed).\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec4\" class=\"Section2\"\u003e \u003ch2\u003eCost breakdowns by taxonomy, typology and impacted sector\u003c/h2\u003e \u003cdiv id=\"Sec5\" class=\"Section3\"\u003e \u003ch2\u003eTaxonomic distribution of the costs\u003c/h2\u003e \u003cp\u003eOut of the 14 most widespread invasive species in Morocco that incur economic costs outside the country (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003ea,c), two had reported costs not only for the target species but also for multiple species collectively. This was observed for the false yellowhead (\u003cem\u003eDittrichia viscosa\u003c/em\u003e) and the feral cat (\u003cem\u003eFelis catus\u003c/em\u003e), whose management costs were documented alongside those of similar species (in the case of the former, with only two cost records), or alongside all invasive species in the region (in the case of the latter, with 26 cost records). Despite their inclusion at the bottom of Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e, these two species were excluded from the subsequent calculations. Considering annual costs in \u003cem\u003eInvaCost\u003c/em\u003e, the remaining 12 species accounted for 480 entries. The cumulative global economic cost for these species amounted to US\u003cspan\u003e$\u003c/span\u003e 4.26\u0026nbsp;billion. The annual costs of these species in Morocco were estimated by multiplying the annual cost per site (or per country in the cases of country-level costs) by the number of locations in Morocco. Thus, we estimated a total economic impact of US\u003cspan\u003e$\u003c/span\u003e 1.61\u0026nbsp;billion (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab1\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 1\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003e \u003cb\u003eCost estimates in InvaCost for the 14 invasive alien species in Morocco, using the expanded dataset.\u003c/b\u003e Species are ordered by their total economic impact reported in InvaCost (US\u003cspan\u003e$\u003c/span\u003e in 2017), except for two species with costs that include other species (the false yellowhead, Dittrichia viscosa, and the feral cat, Felis catus). We also provide the number of cost entries in InvaCost for each species, the type of costs (D: damage, M: management, DM: both), the spatial scale of the costs reported and the countries in which these costs incurred, and the annual costs estimated for Morocco. Annual costs for species within Morocco were estimated by multiplying the annual costs by the number of sites, for all species except those with country as spatial scale. Labels for the spatial scale: C: country level, S: site, within country level, R: river basin, Ha: hectare, G: global. Labels for the countries: Sp: Spain, Po: Portugal, Fr: France, UK: United Kingdom; Mx: Mexico; SA: South Africa; US: United States of America; Au: Australia. Costs estimated in Morocco are annual costs in US\u003cspan\u003e$\u003c/span\u003e 2017; b: billion, m: million, t: thousand.\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"7\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eSpecies\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eAnnual cost\u003c/p\u003e \u003cp\u003ein \u003cem\u003eInvaCost\u003c/em\u003e\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eNumber of entries\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eCost\u003c/p\u003e \u003cp\u003etype\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eSpatial Scale\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003eCountry\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c7\"\u003e \u003cp\u003eCost in Morocco\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eColumba livia\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e1 106 745 250.67\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eDM\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eC\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eUS, UK\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e1.107b\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eSolanum elaeagnifolium\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e456 122 075.40\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eD\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eC\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eRussia\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e456.12m\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eErodium cicutarium\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e96 165.07\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e12\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eD\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eS\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eAu\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e20.58m\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eArundo donax\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e65 126.88\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e229\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eM\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eS, R, Ha\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eFr, Po, Mx, SA, Sp\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e4.77m\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eRicinus communis\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e58 091.49\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e44\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eM\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eS\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eSA, Sp\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e10.90m\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eArctotheca calendula\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e42 738.66\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e115\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eDM\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eC, S, R\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eSp, Au\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e4.41m\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eOxalis pes-caprae\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e35 085.28\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e12\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eM\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eS, G\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eSp, Po\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e5.72m\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eNicotiana glauca\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e1 105.68\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e28\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eM\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eS\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eSp\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e323.96t\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eGambusia holbrooki\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e918.90\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e10\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eM\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eS\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eSp\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e51.46t\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eNerium oleander\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e149.28\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e20\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eM\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eS\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eSA\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e50.16t\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eSolanum linnaeanum\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e128.60\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eM\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eS\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eSp\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e14.27t\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eSchinus molle\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e29.37\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eM\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eS\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eSp\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e3.64t\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eSubtotal\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003e4.256b\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e\u003cb\u003e450\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cb\u003e1.610b\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eFelis catus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e49 107.52\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eM\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eS\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eSp\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e2.60m\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eDittrichia viscosa\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e6 258.37\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e26\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eM\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eS\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eSp\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e657.13t\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eTotal\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003e4.257b\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e\u003cb\u003e495\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cb\u003e1.613b\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003eSpecies with only country-level cost estimates emerged as the most financially burdensome when annual costs were calculated across various spatial scales. Notably, the pigeon (\u003cem\u003eColumba livia\u003c/em\u003e), which experienced a relatively large range expansion in Morocco, incurred the highest economic toll. The reported expenses of pigeon invasions included two instances from the United States (management) and one from the United Kingdom (damage), all documented in the early 2000s. Similarly, the silverleaf nightshade (\u003cem\u003eSolanum elaeagnifolium\u003c/em\u003e), for which country-level cost estimates were available in Russia, were extrapolated to represent potential damages should the species invade the country, likely contributing to the considerable costs reported. Furthermore, costs associated with capeweed (\u003cem\u003eArctotheca calendula\u003c/em\u003e) were reported across various spatial scales, including country (e.g., natural areas in Spain), site, and river levels. While expenses were relatively modest at the country level, ranging around US\u003cspan\u003e$\u003c/span\u003e 2,173.11 annually (N\u0026thinsp;=\u0026thinsp;11), costs were substantially higher at lower spatial scales in Spain and South Africa, reaching US\u003cspan\u003e$\u003c/span\u003e 47,439.19 at the site level (N\u0026thinsp;=\u0026thinsp;103) and US\u003cspan\u003e$\u003c/span\u003e 4,805.56 at the river scale (N\u0026thinsp;=\u0026thinsp;1). As a result, capeweed was classified as having costs predominantly at the site level (all costs were averaged independently of the spatial scale).\u003c/p\u003e \u003cp\u003eWhile the pigeon and the silverleaf nightshade collectively incurred a projected annual cost exceeding US\u003cspan\u003e$\u003c/span\u003e 1.5\u0026nbsp;billion in Morocco, the remaining invasive alien species accounted for an annual cost of US\u003cspan\u003e$\u003c/span\u003e 49.91\u0026nbsp;million in the same region during the same period (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003eb) The lower expenses associated with the remaining species can be due to costs primarily comprising management expenses rather than direct damages. The taxonomic breakdown of estimated annual costs for Morocco is as follows: US\u003cspan\u003e$\u003c/span\u003e 20.58\u0026nbsp;million from 12 management costs reported for the common storksbill (\u003cem\u003eErodium cicutarium\u003c/em\u003e), across three different states in Australia; US\u003cspan\u003e$\u003c/span\u003e 10.90\u0026nbsp;million from 44 management costs reported for the castor bean (\u003cem\u003eRicinus communis\u003c/em\u003e) across various villages in Spain and Kruger National Park in South Africa; US\u003cspan\u003e$\u003c/span\u003e 5.72\u0026nbsp;million from 12 management costs reported for the Bermuda buttercup (\u003cem\u003eOxalis pes-caprae\u003c/em\u003e) with expenditures on research projects conducted in Spain and Portugal; US\u003cspan\u003e$\u003c/span\u003e 4.76\u0026nbsp;million from 229 management costs reported for the giant reed (\u003cem\u003eArundo donax\u003c/em\u003e), covering various regions in Spain, Portugal, and France, as well as specific protected areas in Mexico and South Africa; and US\u003cspan\u003e$\u003c/span\u003e 4.41\u0026nbsp;million from 71 management costs reported for the capeweed (\u003cem\u003eArctotheca calendula\u003c/em\u003e), across different sites in Spain and various states in Australia, along with 44 cost entries accounting for damage costs across different states in Australia.\u003c/p\u003e \u003cp\u003eThe projected annual expenses for the remaining invasive species were less than US\u003cspan\u003e$\u003c/span\u003e 1\u0026nbsp;million. Specifically, this included US\u003cspan\u003e$\u003c/span\u003e 323.96 thousand coming from 28 management expenditures associated with the tree tobacco (\u003cem\u003eNicotiana glauca\u003c/em\u003e), across various locations in Spain; US\u003cspan\u003e$\u003c/span\u003e 51.46 thousand from 10 management costs linked to the mosquitofish (\u003cem\u003eGambusia holbrooki\u003c/em\u003e), from different sites in Spain; and US\u003cspan\u003e$\u003c/span\u003e 50.16 thousand from 20 management costs for the oleander (\u003cem\u003eNerium oleander\u003c/em\u003e), situated in the Kruger National Park in South Africa. The apple of Sodom (\u003cem\u003eSolanum linnaeanum\u003c/em\u003e) with four management costs, and the false pepper tree (\u003cem\u003eSchinus molle\u003c/em\u003e) with a single management cost reported, both at sites in Spain, represented the remaining cost entries totaling US\u003cspan\u003e$\u003c/span\u003e20.70 thousand and US\u003cspan\u003e$\u003c/span\u003e15.95 thousand.\u003c/p\u003e \u003c/div\u003e \u003c/div\u003e \u003cdiv id=\"Sec6\" class=\"Section2\"\u003e \u003ch2\u003eManagement and damage costs\u003c/h2\u003e \u003cp\u003eIn terms of cost type (either damage or management), only four species have reported damage costs: pigeons (\u003cem\u003eColumba livia\u003c/em\u003e, N\u0026thinsp;=\u0026thinsp;1), capeweed (\u003cem\u003eArctotheca calendula\u003c/em\u003e, N\u0026thinsp;=\u0026thinsp;44), the silverleaf nightshade (\u003cem\u003eSolanum elaeagnifolium\u003c/em\u003e, N\u0026thinsp;=\u0026thinsp;2) and the common storksbill (\u003cem\u003eErodium cicutarium\u003c/em\u003e, N\u0026thinsp;=\u0026thinsp;12). By extrapolating, we estimate an annual damage cost of US\u003cspan\u003e$\u003c/span\u003e 856.80\u0026nbsp;million in Morocco, presuming that all occurrences of pigeons and capeweed resulted in direct damages. Based on the annual management costs for pigeons and capeweed, we estimate a potential annual expenditure amounting to US\u003cspan\u003e$\u003c/span\u003e 1.5\u0026nbsp;billion (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e). Notably, projected management expenses surpass damage costs, and this difference is mainly influenced by the reported costs associated with pigeons. If we remove pigeons, damage costs surpass management costs (US\u003cspan\u003e$\u003c/span\u003e 483.03m vs US\u003cspan\u003e$\u003c/span\u003e 27.09m).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec7\" class=\"Section2\"\u003e \u003ch2\u003eSectoral distribution of cost\u003c/h2\u003e \u003cp\u003eAll damage costs were reported for agriculture, except for pigeons, where agriculture and public/social welfare were jointly reported. Regarding management costs, the majority were borne by the authorities and/or stakeholders, including governmental services and official organizations such as conservation agencies, forest services, or associations involved in the broad management of biological invasions, e.g., control programs, eradication campaigns, research funding (Table S2). Two exceptions were noted, capeweed (\u003cem\u003eArctotheca calendula\u003c/em\u003e) and giant reed (\u003cem\u003eArundo donax\u003c/em\u003e), which also reported management costs in agriculture (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003eb). On assessing the annual costs per site for the five invasive species affecting the agricultural sector outside of Morocco, and extrapolating these figures based on occurrences in Morocco, the estimated annual cost impacting agriculture totaled US\u003cspan\u003e$\u003c/span\u003e 859.10\u0026nbsp;million. Similarly, in this context, the economic impact on authorities and stakeholders amounted to US\u003cspan\u003e$\u003c/span\u003e 1.49\u0026nbsp;billion annually.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec8\" class=\"Section2\"\u003e \u003ch2\u003eGeographic distribution of cost\u003c/h2\u003e \u003cp\u003eThe geographic distribution of costs is dependent on the locations of the occurrences of the twelve invasive alien species considered (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e). Besides, it is also dependent on the scale of the cost that these species have.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eSumming the extrapolated annual costs per site for all the species, we observe that the potential economic costs in Morocco are unevenly distributed across the country. Higher costs are concentrated in more densely populated regions, such as Casablanca, Rabat, and Marrakech, as well as along the Atlantic coast, including Casablanca and Rabat.\u003c/p\u003e \u003c/div\u003e"},{"header":"Discussion","content":"\u003cp\u003eThis study presents the first comprehensive global analysis of invasive species in Morocco, offering insights into the potential economic costs associated with biological invasions in the country and the geographic distribution of the estimated costs. Urgent research was prompted by the necessity to identify invasive alien species and map their distribution across Morocco, an essential step for assessing their economic impacts. A total of 551 invasive alien species were identified, representing 1.76% of the nation's plant and animal biodiversity. The absence of cost data for Morocco in the \u003cem\u003eInvaCost\u003c/em\u003e database highlighted the need for promoting cost reporting, and based on extrapolation of costs from other countries, we estimated a potential annual cost amounting to at least US\u003cspan\u003e$\u003c/span\u003e 1.61\u0026nbsp;billion.\u003c/p\u003e \u003cp\u003eCompared to the estimated annual mean cost for the entire African continent, estimated between US\u003cspan\u003e$\u003c/span\u003e2.6 and US\u003cspan\u003e$\u003c/span\u003e 8.6\u0026nbsp;billion in 2019 [\u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e1\u003c/span\u003e, \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2\u003c/span\u003e], the potential economic toll of invasive species in Morocco is, therefore, substantial. However, this estimate is likely conservative, as they were predominantly based on data from a single country, with South Africa accounting for approximately 86% of the total number of cost entries [\u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e14\u003c/span\u003e]. Besides, Eschen et al. [\u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e43\u003c/span\u003e] projected the annual cost of invasive alien species in the Moroccan agricultural sector alone to be US\u003cspan\u003e$\u003c/span\u003e 1.66\u0026nbsp;billion, drawn from literature data and stakeholder interviews. Another example of the underestimation of the costs from Africa generally, is the descriptions of the costs caused by biological invasions in the Mediterranean basin, which were available for only 15 out of 26 countries, with most reports arising from Western European nations, i.e., France, Spain, and Italy, and only 11 records from African countries such as Libya (US\u003cspan\u003e$\u003c/span\u003e 593,04\u0026nbsp;million, N\u0026thinsp;=\u0026thinsp;8) and Egypt (US\u003cspan\u003e$\u003c/span\u003e 147.16\u0026nbsp;million, N\u0026thinsp;=\u0026thinsp;3) [\u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e35\u003c/span\u003e]. While both studies relied on the \u003cem\u003eInvaCost\u003c/em\u003e database, note that none of them incorporated cost data for Morocco. Thus, using known cost estimates for documented species appears to be the only practical and straightforward approach for estimating missing costs in other countries affected by invasions [\u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e44\u003c/span\u003e].\u003c/p\u003e \u003cp\u003eWe estimated a potential annual cost of US\u003cspan\u003e$\u003c/span\u003e 1.61\u0026nbsp;billion in Morocco, and as a first assessment for the country, it should help develop effective national policies and global management initiatives to tackle this issue. Our assessment is essential for informed decision-making at both national and international levels. Birds are the most prevalent invasive alien species nationwide; however, only the pigeon (\u003cem\u003eC. livia\u003c/em\u003e) has recorded economic costs. The lack of cost data for most exotic bird species may stem from their occurrence in regions with limited resources for researching their impacts or in remote areas where tracking their effects proves challenging [\u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e29\u003c/span\u003e]. Alternatively, it could be attributed to research prioritizing well-known taxa [\u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e1\u003c/span\u003e, \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2\u003c/span\u003e] or the most threatening alien species [\u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e36\u003c/span\u003e, \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e46\u003c/span\u003e], given that exotic birds often have minimal environmental and socioeconomic impacts [\u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e47\u003c/span\u003e, \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e48\u003c/span\u003e]. Despite this, pigeons accounted for over half of all exotic bird costs globally [\u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e49\u003c/span\u003e], reflecting their significant economic burden - with an estimated cost of US\u003cspan\u003e$\u003c/span\u003e 1.11\u0026nbsp;billion in Morocco.\u003c/p\u003e \u003cp\u003eIn contrast, invasive alien plants such as the oleander (\u003cem\u003eN. oleander\u003c/em\u003e), tree tobacco (\u003cem\u003eN. glauca\u003c/em\u003e), castor bean (\u003cem\u003eR. communis\u003c/em\u003e), and common storksbill (\u003cem\u003eE. cicutarium\u003c/em\u003e), were the most widespread, each observed ranging between 200 and 350 locations across Morocco. While some authors consider oleander and the common storksbill to be native to Morocco [\u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e50\u003c/span\u003e], projected costs for those species are still very conservative. The tree tobacco is native to northwest Bolivia and Argentina, and is recognized as an invasive plant in Morocco [\u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e51\u003c/span\u003e], whereas the castor bean is native to Northeastern Africa; both probably introduced before 1931 [\u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e52\u003c/span\u003e]. Notably, the silverleaf nightshade, a global threat and the second most expensive plant species in Morocco, is estimated to cost approximately US\u003cspan\u003e$\u003c/span\u003e 456.12\u0026nbsp;million. This plant, introduced unintentionally around 1949, profoundly impacts agriculture, biodiversity, and the environment. For example, losses of up to 64% in maize production have been recorded in Morocco [\u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e53\u003c/span\u003e]. While costs for the silverleaf nightshade were not directly assessed in Morocco, estimates were derived from economic costs reported in Russia, with extrapolations made to predict potential damages if the species spreads there [\u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e54\u003c/span\u003e]. Even though it is challenging to standardize costs across different economies, using exchange rates provides a practical method, albeit with limitations due to the lack of data on Purchasing Power Parity (PPP).\u003c/p\u003e \u003cp\u003eWhile the pigeon and silverleaf nightshade incurred a combined annual cost surpassing US\u003cspan\u003e$\u003c/span\u003e 1.5\u0026nbsp;billion, the other invasive alien species accrued an estimated annual cost of US\u003cspan\u003e$\u003c/span\u003e 49.91\u0026nbsp;million. The comparatively lower costs associated with the remaining species can be attributed to the fact that most of these expenses are primarily related to management rather than direct damages. This aligns with earlier findings regarding the costs of invasive alien species, where reported management expenses typically fall well below reported damages [\u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e28\u003c/span\u003e]. It must be considered that the costs for Morocco are estimated costs using the costs from other countries. Thus, whether costs are typified as management or damage corresponds to the availability of costs in \u003cem\u003eInvaCost\u003c/em\u003e for the species invading Morocco. Thus, as many of the costs come from Spain (many of the invasive species are the same, very likely because of the proximity), where management costs surpass damages [\u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e19\u003c/span\u003e], this could be one of the reasons why most invasive species have management costs available for Morocco. Given that at the global scale, management costs are about ten times lower than damage costs [\u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e28\u003c/span\u003e, \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e29\u003c/span\u003e], the actual costs for Morocco might be much higher than estimated here.\u003c/p\u003e \u003cp\u003eThe uneven geographic distribution of potential economic costs from invasive species in Morocco can be attributed to several factors. Firstly, more densely populated regions like Casablanca, Rabat, and Marrakech have higher economic activity, infrastructure, and assets at risk, leading to greater potential costs when invasive species cause damage. These urban centers serve as major hubs of human and commercial activity, facilitating the introduction and proliferation of invasive species through increased transportation, trade, and tourism. Secondly, the Atlantic coast, where Casablanca and Rabat are located, features ecosystems that are particularly vulnerable to certain invasive species (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e), which can thrive in these environments and cause significant economic harm. For example, \u003cem\u003eSolanum elaeagnifolium\u003c/em\u003e was introduced to Morocco in the mid-20th century, first establishing itself in the Tadla region (Beni Mellal-Khenifra region) [\u003cspan citationid=\"CR55\" class=\"CitationRef\"\u003e55\u003c/span\u003e] and spreading widely by the 1990s. The estimated potential cost incurred thus far stands at US\u003cspan\u003e$\u003c/span\u003e 0.45\u0026nbsp;billion annually (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e), emphasising the significant economic consequences of its spread. Currently, all six geographical zones in Morocco are infested, with the least impact in the Saharan and mountainous areas. Around 70% of infestation sites are in major agricultural districts, and more than 15% along the Rabat-Casablanca axis. Moreover, this species is rapidly adapting and spreading to new regions [\u003cspan citationid=\"CR56\" class=\"CitationRef\"\u003e56\u003c/span\u003e, \u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e57\u003c/span\u003e, \u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e58\u003c/span\u003e]. Coastal regions often host diverse and economically significant industries such as fisheries, tourism, and agriculture that can be severely impacted by invasive species, thereby driving up the costs, such as the Marrakech-Safi region, with an estimated potential cost of US\u003cspan\u003e$\u003c/span\u003e 361.62\u0026nbsp;million (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003e). As the primary tourist destination in the kingdom, Marrakech is home to Morocco\u0026rsquo;s busiest airport, which served over 6.3\u0026nbsp;million passengers in 2019, and includes Safi, the country\u0026rsquo;s oldest port. Additionally, the region accounts for 22% of Morocco\u0026rsquo;s useful agricultural area. Lastly, urban areas and economically vibrant regions tend to have more comprehensive monitoring and reporting systems in place, which can result in better documentation and more frequent records of invasive species presence, compared to less populated and economically active areas.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eGiven that the failure to act on invasive species management can lead to substantial avoidable costs [\u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e27\u003c/span\u003e], Morocco must establish proactive measures aimed at preventing the introduction of new invasive alien species. To achieve this, future studies should focus on identifying and cataloging these potential invasive species. With a better understanding of which species pose a threat and early detection of possible introduction pathways, the country will be better positioned to enable policymakers and environmental managers to develop targeted strategies for interception and control. By taking timely action, Morocco can effectively reduce the economic costs linked to the impacts of invasive species in the future.\u003c/p\u003e \u003cp\u003eOn synthesis, although the estimated annual economic impact of invasive species in Morocco is US\u003cspan\u003e$\u003c/span\u003e 1.61\u0026nbsp;billion, this figure likely underestimates the actual economic burden for several reasons: it is derived from data on only 2% of the invasive species present in Morocco, there is a lack of Morocco-specific cost data, and many invasive species worldwide have relatively low reported costs, including the 12 invasive alien species for which we found cost data. The disproportionate focus on specific taxa, such as pigeons, highlights the need for a more comprehensive approach to invasive species management that considers the full spectrum of ecological and economic impacts. Improved cost reporting, early detection, and effective management strategies are essential for minimizing the economic and environmental consequences of biological invasions. Enhanced collaboration between researchers, policymakers, and stakeholders is necessary to develop targeted laws and interventions, and allocate resources efficiently. By addressing these challenges, we can better protect ecosystems, safeguard biodiversity, and mitigate the economic risks of invasive species in Morocco and the wider region.\u003c/p\u003e"},{"header":"Methods","content":"\u003cdiv id=\"Sec11\" class=\"Section2\"\u003e \u003ch2\u003eData collection, compilation and filtering\u003c/h2\u003e \u003cp\u003eTo assess the economic costs associated with invasive alien species in Morocco, we first identified existing species within the country using comprehensive databases such as GISD, GRIIS, and the \u0026lsquo;Standardising and Integrating Alien Species workflow\u0026rsquo; [\u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e41\u003c/span\u003e]. Additionally, we utilized resource lists such as \u0026lsquo;Terrestrial Invasive Alien Species in the Arab Maghreb\u0026rsquo; and \u0026lsquo;Elements for Reflection on Primary Invasive Alien Species in Morocco\u0026rsquo;. Supplementary systematic searches in online repositories (Web of Science, Google Scholar and Google search engine), and gray literature were performed. We further engaged with national experts and stakeholders to gather additional information, e.g., official national managers or researchers that could provide cost data. We removed duplicates and considered only those that were taxonomically ranked at the species level (i.e., discarding cases with only genus information).\u003c/p\u003e \u003cp\u003eWe then cross-checked the \u003cem\u003eInvaCost\u003c/em\u003e v4.1 database [\u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e17\u003c/span\u003e] (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.6084/m9.figshare.12668570.v5\u003c/span\u003e\u003cspan address=\"10.6084/m9.figshare.12668570.v5\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e) to determine whether any of these species incurred economic costs somewhere in the world. Although \u003cem\u003eInvaCost\u003c/em\u003e compiles cost information obtained from non-English sources, including French and Arabic [\u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e19\u003c/span\u003e], the database still did not feature any reported costs for Morocco. Comparing the list of invasive alien species detected in Morocco with the invasive species having costs in \u003cem\u003eInvaCost\u003c/em\u003e, we separated this information into two categories: invasive alien species detected in Morocco lacking cost information in \u003cem\u003eInvaCost\u003c/em\u003e, and species observed in Morocco with documented costs in \u003cem\u003eInvaCost\u003c/em\u003e, but arising from cost occurrences outside of Morocco (Table \u003cspan refid=\"MOESM1\" class=\"InternalRef\"\u003eS1\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eOccurrence records detailing their current invasive distribution in the country were retrieved from the Global Biodiversity Information Facility (GBIF; \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://www.gbif.org\u003c/span\u003e\u003cspan address=\"https://www.gbif.org\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e) for all species listed in both datasets. In order to have a reliable dataset, we removed duplicated geographic coordinates, occurrences without GPS coordinates, and those with erroneous coordinates falling outside terrestrial borders that could not be georeferenced from the information provided. Taxonomic descriptions were provided for species in both lists, and those with more than 50 occurrences (N\u0026thinsp;=\u0026thinsp;14 species) were selected.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec12\" class=\"Section2\"\u003e \u003ch2\u003eEstimating costs of invasive alien species in Morocco\u003c/h2\u003e \u003cp\u003eBy filtering the \u0026lsquo;Species\u0026rsquo; column in the \u003cem\u003eInvaCost\u003c/em\u003e database, we obtained the reported costs for the 14 invasive alien species. We considered only high-reliability cost entries from the \u0026lsquo;Reliability\u0026rsquo; column, i.e., cost estimates sourced from official peer-reviewed material and/or material with reproducible methods [\u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e17\u003c/span\u003e]. Costs were extracted from the \u0026lsquo;Cost_estimate_per_year_2017_USD_exchange_rate\u0026rsquo; column, such that all cost entries were adjusted to a common currency and year for comparability, i.e., in 2017 US dollars. This adjustment also factored in inflation, referencing the Consumer Price Index of 2017 in relation to the year of the cost estimation. Each entry also includes the timespan linked with the recorded costs, facilitated by the \u003cem\u003eexpandYearlyCosts\u003c/em\u003e function from the \u003cem\u003einvacost\u003c/em\u003e R package [\u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e42\u003c/span\u003e]. This function distributes the total reported cost evenly across probable start and end years, generating an expanded dataset where each entry represents a cost estimate for a specific year. Within the \u003cem\u003eInvaCost\u003c/em\u003e database, each publication served as a distinct point of reference for reported costs, although the duration over which costs were estimated differed among references. Complete details regarding the compilation and standardization of data stored in \u003cem\u003eInvaCost\u003c/em\u003e are provided in Diagne et al. [\u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e17\u003c/span\u003e].\u003c/p\u003e \u003cp\u003eFirst, we calculated the total economic cost reported in \u003cem\u003eInvaCost\u003c/em\u003e for each of these 14 IAS. Then, to ascertain a relative annual cost per species, we calculated the annual amount per spatial unit, whether it be as large as a country or as small as a local site, utilizing the \u0026lsquo;Spatial_scale\u0026rsquo; column. A site refers to an area at the intra-country level, while a unit denotes a well-defined surface area or entity, such as a hectare or a river basin. This adjustment was necessary due to the non-uniformity in the number of reported costs and the spatial scale of these costs in \u003cem\u003eInvaCost\u003c/em\u003e across various species. We took into account the costs incurred by different impacted sectors, as indicated in the \u0026lsquo;Impacted_sector\u0026rsquo; column, and the cost typology (i.e., management and damages), as specified in the \u0026lsquo;Type_of_cost_merged\u0026rsquo; column.\u003c/p\u003e \u003cp\u003eWe then extrapolated the economic costs accounting for the distribution of each species, by multiplying the annual cost per site, obtained from the reported data in \u003cem\u003eInvaCost\u003c/em\u003e, by the number of occurrences sourced from GBIF. An estimate of the total cost incurred by invasive species within Morocco was computed as the sum of extrapolated annual costs per site for all species. Finally, we created a map that included the sum of the extrapolated annual costs per site for all the species in order to understand the spatial distribution of potential economic costs in Morocco and highlight the places in the country that are predicted to cause the highest economic costs of invasive alien species. For the two species that have costs at the country level (\u003cem\u003eC. livia\u003c/em\u003e and \u003cem\u003eS. elaeagnifolium\u003c/em\u003e), we first calculated a cost per site by dividing the country cost by the number of occurrences in Morocco.\u003c/p\u003e \u003c/div\u003e"},{"header":"Declarations","content":"\u003cp\u003e\u003cstrong\u003eData availability Statement\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eData supporting the results reported in this article can be found in:\u003c/p\u003e\n\u003cul\u003e\n \u003cli\u003eGlobal Invasive Species Database (GISD) https://www.iucngisd.org/gisd/\u003c/li\u003e\n \u003cli\u003eGlobal Register of Introduced and Invasive Species (GRIIS) https://griis.org/\u003c/li\u003e\n \u003cli\u003eStandardising and Integrating Alien Species workflow (SInAS) https://doi.org/10.3897/neobiota.59.53578\u003c/li\u003e\n \u003cli\u003eTerrestrial Invasive Alien Species in the Arab Maghreb https://www.cbd.int/invasive/doc/arab-maghreb-eee-fr.pdf\u003c/li\u003e\n \u003cli\u003eElements for Reflection on Primary Invasive Alien Species in Morocco https://doi.org/10.1002/9781119607045.ch7\u003c/li\u003e\n \u003cli\u003e\u003cem\u003eInvaCost\u003c/em\u003e database https://doi.org/10.6084/m9.figshare.12668570.v5\u003c/li\u003e\n \u003cli\u003eGlobal Biodiversity Information Facility (GBIF) https://www.gbif.org/fr/\u003c/li\u003e\n\u003c/ul\u003e\n\u003cp\u003e\u0026nbsp;\u003cstrong\u003eAcknowledgments\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eWe would like to thank the Direction of Climate and the Biological Diversity, Ministry of Energy Transition and Sustainable Development, Morocco\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAuthor contributions:\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eEA and AT contributed to the study conception and design. Data collection was performed by CD, EA, FC, and LBM, the methodology was agreed by JEJ, LBM, EA and AT and analysis were performed by JEJ, CD, EA and LBM. Supervision was carried out by EA and AT and funding was spent by FC, EA and DA. The first draft of the manuscript was written by JEJ and visualization items were created by JEJ, AT and EA. All authors commented on previous versions of the manuscript and all authors read and approved the final\u0026nbsp;manuscript.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAdditional Information\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eFunding\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eFunding was partially given to Elena Angulo by Junta de Andaluc\u0026iacute;a, Consejer\u0026iacute;a de Universidad, Investigaci\u0026oacute;n e Innovaci\u0026oacute;n, PROYEXCEL_00688 within\u0026nbsp;the\u0026nbsp;PAIDI\u0026nbsp;2020. The InvaCost project was funded by the French National Research Agency (ANR-14-CE02-0021), the BNP-Paribas Foundation Climate Initiative, the AXA Research Fund Chair of Invasion Biology of University Paris Saclay and by the BiodivERsA and Belmont-Forum call 2018 on biodiversity scenarios (AlienScenarios; BMBF/PT\u0026nbsp;DLR\u0026nbsp;01LC1807C).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eCompeting Interests\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe authors have no conflicts of interest to disclose.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\n\u003cli\u003ePy\u0026scaron;ek, P., Richardson, D. M. 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Infestation by Silverleaf Nightshade Solanum elaeagnifolium (Cav.) in the eastern region of Morocco. \u003cem\u003eRevue Marocaine de Protection des Plantes\u003c/em\u003e, \u003cstrong\u003e4\u003c/strong\u003e:59-66 (2013)\u003c/li\u003e\n\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":true,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"scientific-reports","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"scirep","sideBox":"Learn more about [Scientific Reports](http://www.nature.com/srep/)","snPcode":"","submissionUrl":"","title":"Scientific Reports","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"stoa","reportingPortfolio":"Scientific Reports","inReviewEnabled":true,"inReviewRevisionsEnabled":true},"keywords":"Invasive species, Economic impacts, Management strategies, Morocco, Biological invasions, InvaCost","lastPublishedDoi":"10.21203/rs.3.rs-4731421/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-4731421/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003e Biological invasions pose substantial economic threats globally, yet detailed cost assessments for many Global South nations, especially in Africa, remain scarce. This study presents the first comprehensive breakdown of the potential costs of biological invasions in Morocco. We identified 551 invasive alien species, comprising approximately 1.76% of Morocco's biodiversity. Using the \u003cem\u003eInvaCost\u003c/em\u003e database, we gathered cost data for the 12 most frequent invasive species with available data. Calculating the mean annual cost for each species and extrapolating based on their prevalence in Morocco, we estimated the potential annual economic impact to be US\u003cspan\u003e$\u003c/span\u003e 1.61\u0026nbsp;billion. Urban species management and damage, especially pigeons, accounted for a substantial portion of this impact. Invasive plant species also emerged as prevalent and costly. Annual management costs exceeded damage costs (US\u003cspan\u003e$\u003c/span\u003e 1.50\u0026nbsp;billion vs. 856.80\u0026nbsp;million), driven mainly by pigeon management. Costs by authorities and stakeholders outweighed agricultural costs (US\u003cspan\u003e$\u003c/span\u003e 1.49\u0026nbsp;billion vs. 859.10\u0026nbsp;million). Despite challenges in extrapolating cost data from other regions, this study underscores the urgent need for targeted management and policy interventions to minimize the spread of invasive species and reduce their economic toll. Morocco can implement proactive management measures and foster international collaborations to tackle this socio-ecological crisis, ensuring long-term sustainability and prosperity.\u003c/p\u003e","manuscriptTitle":"Biological invasions and their potential economic costs in Morocco","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2024-09-09 09:16:23","doi":"10.21203/rs.3.rs-4731421/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"decision","content":"Revision requested","date":"2025-04-17T06:14:09+00:00","index":"","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2025-03-27T10:46:48+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2025-03-23T18:49:48+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"40806093071461913790582026132428618314","date":"2025-03-23T11:02:46+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"164961587425545823470562008043569109380","date":"2025-03-20T09:30:59+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"288444016046852267935173854942010734761","date":"2025-03-19T09:31:26+00:00","index":"hide","fulltext":""},{"type":"reviewersInvited","content":"","date":"2025-03-18T08:18:33+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2024-10-24T04:13:23+00:00","index":"","fulltext":""},{"type":"editorInvited","content":"","date":"2024-08-13T17:57:59+00:00","index":"","fulltext":""},{"type":"checksComplete","content":"","date":"2024-08-08T06:47:13+00:00","index":"","fulltext":""},{"type":"submitted","content":"Scientific Reports","date":"2024-07-12T15:44:24+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"scientific-reports","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"scirep","sideBox":"Learn more about [Scientific Reports](http://www.nature.com/srep/)","snPcode":"","submissionUrl":"","title":"Scientific Reports","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"stoa","reportingPortfolio":"Scientific Reports","inReviewEnabled":true,"inReviewRevisionsEnabled":true}}],"origin":"","ownerIdentity":"d177648f-b353-416e-b476-424ef65beb20","owner":[],"postedDate":"September 9th, 2024","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"published-in-journal","subjectAreas":[],"tags":[],"updatedAt":"2026-01-19T17:06:55+00:00","versionOfRecord":{"articleIdentity":"rs-4731421","link":"https://doi.org/10.1038/s41598-025-31767-8","journal":{"identity":"scientific-reports","isVorOnly":false,"title":"Scientific Reports"},"publishedOn":"2026-01-15 16:30:33","publishedOnDateReadable":"January 15th, 2026"},"versionCreatedAt":"2024-09-09 09:16:23","video":"","vorDoi":"10.1038/s41598-025-31767-8","vorDoiUrl":"https://doi.org/10.1038/s41598-025-31767-8","workflowStages":[]},"version":"v1","identity":"rs-4731421","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-4731421","identity":"rs-4731421","version":["v1"]},"buildId":"qtupq5eGEP_6zYnWcrvyt","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}