Description of two new species of freshwater Gammarus Fabricius, 1775 (Amphipoda: Gammaridae) from the Middle Atlas of Morocco
Abstract
This study describes, both morphologically and molecularly, two new species, Gammarus latispinus sp. nov. and G. tazekkaensis sp. nov. from the Middle Atlas of Morocco. Both species are similar to Gammarus acalceolatus, but differ by having setae as long as, or longer than, the diameter of the article on the second antenna, lacking calceoli, and by the absence of long setae on merus and carpus of pereopods 5–7. Gammarus latispinus sp. nov. is characterized by the presence of spinules on the outer face of epimeral plates 2 and 3, a pereopod 5 much shorter than pereopods 6 and 7, and telson lobes with dorsal setae but without lateral spines. Gammarus tazekkaensis sp. nov. is also characterized by the presence of a single lateral spine on telson lobes. Full descriptions of the new species are provided in this paper together with molecular data (genes COI and 28S) that corroborate the molecular delimitation. The two new species are phylogenetically close to G. acalceolatus, followed by G. marmouchensis and G. rifatlensis, all of which inhabit the Middle Atlas Mountains in Morocco.
References
Arfianti T., Wilson S. & Costello M.J. 2018. Progress in the discovery of amphipod crustaceans. PeerJ 6: e5187. https://doi.org/10.7717/peerj.5187
Aspin T.W.H., Khamis K., Matthews T.J., Milner A.M., O’Callaghan M.J., Trimmer M., Woodward G. & Ledger M.E. 2019. Extreme drought pushes stream invertebrate communities over functional thresholds. Global Change Biology 25 (1): 230–244. https://doi.org/10.1111/gcb.14495
Ayati K., Hadjab R., Khammar H., Dhaouadi S., Piscart C. & Mahmoudi E. 2019. Origin, diversity and distribution of freshwater epigean amphipods in Maghreb. Annales de Limnologie – International Journal of Limnology 55: 13. https://doi.org/10.1051/limn/2019012
Bespalaya Y.V., Vinarski M.V., Aksenova O.V. et al. 2024. Phylogeny, taxonomy, and biogeography of the Sphaeriinae (Bivalvia: Sphaeriidae). Zoological Journal of the Linnean Society 201 (2): 305–338. https://doi.org/10.1093/zoolinnean/zlad139
Castresana J. 2000. Selection of conserved blocks from multiple alignments for their use in phylogenetic analysis. Molecular Biology and Evolution 17 (4): 540–552. https://doi.org/10.1093/oxfordjournals.molbev.a026334
Dehedin A., Dole-Olivier M.J., Piscart C., Mimoun, D. & Marmonier P. 2013a. Global change and drying modality affect interstitial assemblages of alluvial wetlands. Wetlands 33: 537–550
Dehedin, A., Maazouzi S., Puijalon S., Marmonier P. & Piscart C. 2013b. Combined effects of the water level reduction and the increase in ammonia concentrations on organic matter processing by key freshwater shredders in alluvial wetlands. Global Change Biology 19: 763–774
Edgar R.C. 2004. MUSCLE: multiple sequence alignment with high accuracy and high throughput. Nucleic Acids Research 32 (5): 1792–1797. https://doi.org/10.1093/nar/gkh340
El Caidi S., Taybi A.F. & Mabrouki Y. 2025. Re-discovery of the invasive guppy Poecilia reticulata (Peters, 1859) (Poeciliidae) in Morocco. Natura Croatici: Periodicum Musei Historiae Naturalis Croatici 34 (1): 330237. https://doi.org/10.20302/NC.2025.34.6
El Morabet I., Taybi A., Mabrouki Y., Bouhaddioui A. & Bahhou J. 2021. Assessment of physico-chemical and biological parameters of Aguelmam Sidi Ali Lake, a threatened Ramsar Site, Morocco. AACL Bioflux 15 (1): 575–583.
Fadil F. & Dakki M. 2006. Deux espèces nouvelles du genre Gammarus (Crustacés, Amphipodes) du Maroc. Bulletin de l’Institut Scientifique, Rabat, Section Sciences de la Vie (28): 13–24.
Felsenstein J. 1985. Confidence limits on phylogenies: An approach using the bootstrap. Evolution 39: 783–791. https://doi.org/10.2307/2408678
Folmer O., Black M., Hoeh W., Lutz R. & Vrijenhoek R. 1994. DNA primers for amplification of mitochondrial cytochrome c oxidase subunit I from diverse metazoan invertebrates. Molecular Marine Biology and Biotechnology 3 (5): 294–299.
Glöer P., Mabrouki Y. & Taybi A.F. 2020. Two new valvatoid genera (Gastropoda, Hydrobiidae) from Morocco. Ecologica Montenegrina 30: 124–128. https://doi.org/10.37828/em.2020.30.12
Gouy M., Guindon S. & Gascuel O. 2010. SeaView version 4: A multiplatform graphical user interface for sequence alignment and phylogenetic tree building. Molecular Biology and Evolution 27 (2): 221–224. https://doi.org/10.1093/molbev/msp259
Hadjab R., Ayati K. & Piscart C. 2021. A new species of freshwater amphipods Echinogammarus (Amphipoda, Gammaridae) from Algeria. Taxonomy 1 (1): 36–47. https://doi.org/10.3390/taxonomy1010005
Hasegawa M., Kishino H. & Yano T.A. 1985. Dating of the human-ape splitting by a molecular clock of mitochondrial DNA. Journal of Molecular Evolution 22: 160–174. ttps://doi.org/10.1007/BF02101694
Hou Z., Fu J. & Li S. 2007. A molecular phylogeny of the genus Gammarus (Crustacea: Amphipoda) based on mitochondrial and nuclear gene sequences. Molecular Phylogenetics and Evolution 45 (2): 596–611. https://doi.org/10.1016/j.ympev.2007.06.006
Ibrahimi H., Mabrouki Y. & Taybi A.F. 2021. Tinodes atlasensis Ibrahimi, Mabrouki & Taybi, sp. nov. (Trichoptera: Psychomyiidae), a new species from Atlas Mountains, Morocco. Natura Croatica: Periodicum Musei Historiae Naturalis Croatici 30 (2): 367–375. https://doi.org/10.20302/NC.2021.30.23
Karaman G. & Pinkster S. 1977. Freshwater Gammarus species from Europe, North-Africa and adjacent regions of Asia (Crustacea-Amphipoda). 1. Gammarus pulex-group and related species. Bijdragen tot de Dierkunde 47 (1): 1–97.
Maasri A. & Bonada N. 2024. Chapter 2 – Ecology of Mediterranean freshwater ecosystems. In: Maasri A. & Thorp J.H. (eds) Identification and Ecology of Freshwater Arthropods in the Mediterranean Basin: 5–16. Elsevier, Amsterdam.
Mabrouki Y., Terec A., Taybi F., Dénes A. & Keresztes L. 2023. Taxonomic notes and key to the West Palearctic Antocha (Antocha) Osten Sacken, 1860 (Diptera, Limoniidae) with description of a new species from Morocco. Biodiversity Data Journal 11: e103849. https://doi.org/10.3897/BDJ.11.e103849
Mabrouki Y., Taybi F.A., Shrestkha M., Sidorovskyi S., Utevsky A. & Utevsky S. 2025. Do wild and aquacultural alien leeches of the genus Barbronia (Hirudinea: Salifidae) belong to Barbronia weberi? Evidence from Morocco and Ukraine. Ecologica Montenegrina 84: 161–173. https://doi.org/10.37828/em.2025.84.12
Piscart C., Camacho A.I., Coineau N., Christodoulou M., Messana G. & Wittmann K.J. 2024. Chapter 7 – Class Malacostraca (subclass Eumalacostraca). In: Maasri A. & Thorp J.H. (eds) Identification and Ecology of Freshwater Arthropods in the Mediterranean Basin: 157–223. Elsevier, Amsterdam.
Puillandre N., Brouillet S. & Achaz G. 2021. ASAP: assemble species by automatic partitioning. Molecular Ecology Resources 21 (2): 609–620. https://doi.org/10.1111/1755-0998.13281
Saitou N. & Nei M. 1987. The neighbor-joining method: a new method for reconstructing phylogenetic trees. Molecular Biology and Evolution 4: 406–425.
Schneider A., Jost A., Coulon C., Silvestre M., Théry S. & Ducharne A. 2017. Global-scale river network extraction based on high-resolution topography and constrained by lithology, climate, slope, and observed drainage density. Geophysical Research Letters 44 (6): 2773–2781. https://doi.org/10.1002/2016GL071844
Stock J.H. & Von Vaupel Klein J.C. 1996. Mounting Media Revisited: The suitability of Reyne’s fluid for small Crustaceans on JSTOR. Crustaceana 69 (6): 794–798.
Taybi A.F., Glöer P. & Mabrouki Y. 2022. Four new species of the genus Pseudamnicola Paulucci, 1878 from Morocco (Gastropoda: Hydrobiidae). Invertebrate Zoology 19 (2): 191–198. https://doi.org/10.15298/invertzool.19.2.07
Taybi A.F., Mabrouki Y. & Piscart C. 2023. Distribution of freshwater alien animal species in Morocco: current knowledge and management issues. Diversity 15 (2): 169. https://doi.org/10.3390/d15020169
Taybi A.F., Mabrouki Y., Glöer P. & Piscart C. 2024. Factors explaining the distribution of Physella acuta (Draparnaud, 1805) in freshwaters of Morocco. Water 16 (6): 803. https://doi.org/10.3390/w16060803
Tierno de Figueroa J.M., López-Rodríguez M.J., Fenoglio S., Sánchez-Castillo P. & Fochetti R. 2013. Freshwater biodiversity in the rivers of the Mediterranean Basin. Hydrobiologia 719 (1): 137–186. https://doi.org/10.1007/s10750-012-1281-z
Tuekam Kayo R.P., Marmonier P., Zebaze Togouet S.H., Nola M. & Piscart C. 2012. An annotated checklist of freshwater stygobiotic crustaceans of Africa and Madagascar. Crustaceana 85 (12–13): 1613–1631. https://doi.org/10.1163/15685403-00003134
Vences M., Patmanidis S., Fedosov A., Miralles A. & Puillandre N. 2024. iTaxoTools 1.0: Improved DNA barcode exploration with TaxI2. In: DeSalle R. (ed.) DNA Barcoding: Methods and Protocols: 281–296. Springer US, New York, USA. https://doi.org/10.1007/978-1-0716-3581-0_18
Verovnik R., Sket B. & Trontelj P.P. 2005. The colonization of Europe by the freshwater crustacean Asellus aquaticus (Crustacea: Isopoda) proceeded from ancient refugia and was directed by habitat connectivity. Molecular Ecology 14 (14): 4355–4369. https://doi.org/10.1111/j.1365-294X.2005.02745.x
Zhang J., Kapli P., Pavlidis P. & Stamatakis A. 2013. A general species delimitation method with applications to phylogenetic placements. Bioinformatics 29 (22): 2869–2876. https://doi.org/10.1093/bioinformatics/btt499
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