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Strong upwelling conditions drive differences in species abundance and community composition along the Atlantic coasts of Morocco and Western Sahara


Upwelling strongly influences the composition and dynamics of coastal communities by affecting species abundances, recruitment, dispersal and distribution. Coastal upwelling areas are key model regions to study the responses of coastal species to climate change because they are characterized by cooler water conditions and experience lower warming rates than adjacent regions, making them effective ‘control’ or refuge sites. This is particularly true for the benthic species of rocky shores in upwelling areas because they are sedentary, inhabit the interface between marine and terrestrial habitats, are exposed to extremely severe and variable environmental conditions and often live near their tolerance limits. We sampled roughly 2000 km of the Atlantic coast of Morocco and Western Sahara to assess the influence of upwelling cells on patterns of diversity and abundance of northern African rocky shore species. We recorded 186 taxa, providing clarification of the distribution of 141 algae and documenting nine new species records for Morocco and Western Sahara. The results emphasize the influence of upwelling on the abundance and distribution of these organisms. The contrast between non-upwelling and upwelling areas highlights the direct and indirect importance of water temperature in shaping these communities, pointing to the consequences of large-scale warming. Such warming is likely to threaten intertidal species that already live close to their thermal tolerance limits and are not buffered by the effects of upwelling.

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We thank I. Bárbara, A.M. Tavares and M. Namora for assistance with species identification. We thank R. Jacinto, M. Silva and P. Madeira for field assistance. We thank Stephen Hawkins and an anonymous referee for their comments.


This research was funded by projects UID/Multi/04326/2019 and IF/01413/2014/CP1217/CT0004 from the Fundação para a Ciência e Tecnologia (FCT-MEC, Portugal) and based upon research supported by the National Research Foundation of South Africa (Grant number 64801). LAK was supported by a PhD scholarship from the Programme Science without Borders (Conselho Nacional de Desenvolvimento Científico e Tecnológico, of the Ministry of Science, Technology and Innovation of Brazil—237998/2012-2).

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Correspondence to Gerardo I. Zardi.

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The authors declare that they have no conflict of interest.

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This article does not contain any studies with animals performed by any of the authors.

Sampling and field studies

No specific permits were necessary as the species collected were not endangered or legally protected, and all sampling sites had free public access and were not protected or privately owned.

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The datasets generated during and/or analysed during the current study are available from the corresponding author on reasonable request.

Author contribution

GIZ and KRN conceived, designed research conducted survey. CDM conducted survey and contributed to statistical analyses. KRN and CRL conducted statistical analyses. CRL conducted field survey. LAK contributed to analyses. CRL, KRN and GIZ wrote the manuscript. All authors read and approved the manuscript.

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S1 Table

Sampling sites. List of sampling sites along the Atlantic coasts of Morocco and Western Sahara. (DOCX 13 kb)

S2 Table

Results of DistLM model. Environmental variables were analysed individually (marginal tests) and sequentially using the ‘all specified’ selection procedure and the Akaike (AIC) selection criterion. Prop (%) is the proportion of variance in taxon richness, abundance of functional groups, taxa from quadrats and from transects explained by each variable. Significant (p < 0.05) values are indicated in bold. (DOCX 17 kb)

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Lourenço, C.R., Nicastro, K.R., McQuaid, C.D. et al. Strong upwelling conditions drive differences in species abundance and community composition along the Atlantic coasts of Morocco and Western Sahara. Mar. Biodivers. 50, 15 (2020).

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  • Biogeography
  • Intertidal
  • Biodiversity
  • Distributional shift