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The influence of palaeogeography and tectonic events on trilobite distributions in Morocco and northwestern Algeria

Published online by Cambridge University Press:  25 January 2022

Valentin Bault Open the ORCID record for Valentin Bault [Opens in a new window] ,
Catherine Crônier  and
Arnaud Bignon
Valentin Bault*
Affiliation:
Univ. Lille, CNRS, UMR 8198 – Evo-Eco-Paleo, F-59000Lille, France
Catherine Crônier
Affiliation:
Univ. Lille, CNRS, UMR 8198 – Evo-Eco-Paleo, F-59000Lille, France
Arnaud Bignon
Affiliation:
Centro de Investigaciones en Ciencias de la Tierra: Consejo Nacional de Investigaciones Científicas y Técnicas–Universidad Nacional de Córdoba, Córdoba (X5016GCA), Argentina
*
Author for correspondence: Valentin Bault, Email: [email protected]
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Abstract

The effects of sea-level changes and anoxia on Devonian trilobites have been studied for a long time, but the importance of palaeogeographic and tectonic events in this key-period is still not well understood. In the Devonian Period, trilobites invaded many different marine environments and areas in North Africa where important palaeogeographic changes occurred. Distribution patterns of trilobites through time and space have been analysed using a hierarchical cluster analysis and diversity indices. Our examination of the literature suggests that trilobites were scarce during Lochkovian time before an important diversification in Pragian time. Trilobites flourished in many North African regions without there being important taxonomic differences between basins, because of free migration in relatively flat palaeo-topography and homogeneous environments. During Middle Devonian time, early Variscan tectonic movements transformed the eastern Anti-Atlas area into a basin with a platform topography. Geographical barriers such as deep basins prevented trilobite migrations. At the beginning of Eifelian time, the reduction in migration between the different regions of this area coincided with a decrease in diversity. Consequently, tectonic events played an important role in the decline of trilobites during Middle Devonian time, especially when these were combined with sea-level changes and anoxic/hypoxic events. A recovery occurred in Famennian time involving only new genera. As at the global scale, cyrtosymbolines developed in shallow seas, whereas phacopids evolved in deeper environments. The basin and platform system still hampered migrations, although sea-level variations led to episodic exchanges. The late Famennian regression reduced trilobite diversity dramatically in the study area.

Keywords

arthropodsAnti-Atlasassociationsdiversityfaunal successiontectonicpalaeobiogeography
Type
Original Article
Information
Geological Magazine , Volume 159 , Issue 5 , May 2022 , pp. 707 - 729
Copyright
© The Author(s), 2022. Published by Cambridge University Press

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References

Abbache, A, Ouali Mehadji, A, Crônier, C, Allaire, N and Monnet, C (2019) Le Dévonien supérieur du Sahara nord-occidental (Algérie): faciès, environnements et signification géodynamique des calcaires griottes. Carnets de Géologie 19, 71–96.Google Scholar
Abe, FR and Lieberman, BS (2012) Quantifying morphological change during an evolutionary radiation of Devonian trilobites. Paleobiology 38, 292–307.CrossRefGoogle Scholar
Adrain, JM (2008) A global species database of Trilobita: progress, results, and revision of the Treatise. In Advances in Trilobite Research (eds Rabano, I, Gozalo, R and Garcia-Bellido, D), pp. 27–8. Cuadernos del Museo Geominero 9. Madrid: Instituto Geológico y Minero de España. Google Scholar
Alberti, GKB (1964) Neue Trilobiten aus dem marokkanischen und deutschen Unter- und Mitteldevon. Senckenbergiana lethaea 45, 115–33.Google Scholar
Alberti, GKB (1966a) Über einige neue Trilobiten aus dem Silurium und Devon, besonders von Marokko. Senckenbergiana lethaea 47, 111–21.Google Scholar
Alberti, GKB (1966b) Zur taxonomie und verbreitung der Trilobiten-Gattung Kolihapeltis Prantl & Přibyl 1947 im Unter-Devon. Paläontologische Zeitschrift 40, 192–204.CrossRefGoogle Scholar
Alberti, GKB (1967a) Neue obersilurische sowie unter- und mitteldevonische Trilobiten aus Marokko, Deutschland und einigen anderen europäischen Gebieten. 1. Senckenbergiana lethaea 48, 463–79.Google Scholar
Alberti, GKB (1967b) Neue obersilurische sowie unter- und mitteldevonische Trilobiten aus Marokko, Deutschland und einigen anderen europäischen Gebieten. 2. Senckenbergiana lethaea 48, 481–509.Google Scholar
Alberti, GKB (1969) Trilobiten des jüngeren Siluriums sowie des Unter- und Mitteldevons. Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft 520, 1–692.Google Scholar
Alberti, GKB (1970) Trilobiten des jüngeren Siluriums sowie des Unter-und-Mitteldevons. II. Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft 525, 1–233.Google Scholar
Alberti, GKB (1981a) Scutelluidae (Trilobita) aus dem Unterdevon des Hamar Laghdad (Tafilalt, SE-Marokko) und das Alter der ‘mud mounds’ (Ober-Zlichovium bis tiefstes Dalejum). Senckenbergiana lethaea 62, 193–204.Google Scholar
Alberti, GKB (1981b) Trilobiten des jüngeren Siluriums sowie des Unter- und Mitteldevons. III. Senckenbergiana lethaea 62, 1–75.Google Scholar
Alberti, GKB (1982) Der Hamar Laghdad (Tafilalt, SE Marokko), eine bedeutende Fundstätte devonischer Trilobiten. Natur und Museum 112, 172–82.Google Scholar
Alberti, GKB (1983) Trilobiten des jüngeren Siluriums sowie des Unter- und Mitteldevons. IV. Senckenbergiana lethaea 64, 1–87.Google Scholar
Alberti, H (1972) Ontogenie des Trilobiten Phacops accipitrinus . Neues Jahrbuch für Geologie und Paläontologie Abhandlungen 141, 1–36.Google Scholar
Alberti, H (1973) Neue Trilobiten (Cyrtosymbolen) aus dem Ober-Devon IV bis VI (Nord-Afrika und Mittel-Europa). Beitrag 1. Neues Jahrbuch für Geologie und Paläontologie Abhandlungen 144, 143–80.Google Scholar
Alberti, H (1974) Neue Trilobiten (Chaunoproetiden, Mirabolen) aus dem Ober-Devon IV-VI (Nord-Afrika und Mittel-Europa). Beitrag 2. Neues Jahrbuch für Geologie und Paläontologie Abhandlungen 146, 221–61.Google Scholar
Alberti, H (1975a) Neue Trilobiten (Waribole) aus dem Ober-Devon IV-VI (Nord-Afrika und Mittel-Europa). Beitrag 3. Neues Jahrbuch für Geologie und Paläontologie Abhandlungen 149, 180–210.Google Scholar
Alberti, H (1975b) Neue Trilobiten (Waribole) aus dem Ober-Devon IV-V (Nord-Afrika und Mittel-Europa). Beitrag 4. Nebst Revision einiger Typus-Spezies der Proetidae Salter, 1864, aus dem Ober-Devon II-V. Neues Jahrbuch für Geologie und Paläontologie Abhandlungen 150, 207–26.Google Scholar
Alberti, H (1976a) Neue Trilobiten (Cyrtosymbole) aus dem Ober-Devon II (Nord-Afrika und Mittel-Europa). Beitrag 5. Neues Jahrbuch für Geologie und Paläontologie Abhandlungen 152, 207–26.Google Scholar
Alberti, H (1976b) Neue Trilobiten (Richteraspis und Vittaella n. g.) aus dem Ober-Devon III (Nord-Afrika und Mittel-Europa). Beitrag 7. Neues Jahrbuch für Geologie und Paläontologie Abhandlungen 152, 19–50.Google Scholar
Allaire, N, Abbache, A, Crônier, C and Monnet, C (2020) Famennian (Late Devonian) ammonoids from the Ouarourout section (Saoura Valley, Algeria). Annales de Paléontologie 106, 102363. doi: 10.1016/j.annpal.2019.08.002.CrossRefGoogle Scholar
Baidder, L, Raddi, Y, Tahiri, M and Michard, A (2008) Devonian extension of the Panafrican crust north of the West African Craton, and its bearing on the Variscan foreland deformation: evidence from eastern Anti-Atlas (Morocco). In The Boundaries of the West African Craton (eds Ennih, N and Liégeois, JP), pp. 453–65. Geological Society of London, Special Publication no. 297.Google Scholar
Baselga, A (2010) Partitioning the turnover and nestedness components of beta diversity. Global Ecology and Biogeography 19, 134–43.CrossRefGoogle Scholar
Baselga, A and Orme, CDL (2012) Betapart: an R package for the study of beta diversity. Methods in Ecology and Evolution 3, 808–12.CrossRefGoogle Scholar
Bault, V, Crônier, C, Allaire, N and Monnet, C (2021) Trilobite biodiversity trends in the Devonian of North Africa. Palaeogeography, Palaeoclimatology, Palaeoecology 565, 110208. doi: 10.1016/j.palaeo.2020.110208.CrossRefGoogle Scholar
Becker, RT, Aboussalam, ZS, Helling, S, Afhüppe, L, Baidder, L and El Hassani, A (2018) The world-famous Devonian mudmounds at Hamar Laghdad and overlying cephalopod-rich strata. Münstersche Forschungen zur Geologie und Paläontologie 110, 188–213.Google Scholar
Becker, RT, Jansen, U, Plodowski, G, Schindler, E, Aboussalam, ZS and Weddige, K (2004) Devonian litho-and biostratigraphy of the Dra Valley area – an overview. Document de l’Institut Scientifique, Rabat 19, 3–18.Google Scholar
Becker, RT, Königshof, P and Brett, CE (2016) Devonian climate, sea level and evolutionary events: an introduction. In Devonian Climate, Sea Level and Evolutionary Events (eds Becker, RT, Königshof, P and Brett, CE), pp. 1–10. Geological Society of London, Special Publication no. 423.Google Scholar
Belka, Z, Klug, C, Kaufmann, B, Korn, D, Döring, S, Feist, R and Wendt, J (1999) Devonian conodont and ammonoid succession of the eastern Tafilalt (Ouidane Chebbi section), Anti-Atlas, Morocco. Acta Geologica Polonica 49, 1–23.Google Scholar
Bendella, M and Ouali Mehadji, A (2015) Depositional environment and ichnology (Nereites ichnofacies) of the Late Devonian Sahara region (SW Algeria). Arabian Journal of Geosciences 8, 5303–16.CrossRefGoogle Scholar
Benhamou, M, Abbache, A, Elmi, S, Mekahli, L, Racheboeuf, P, Ouali-Mehadji, A and Boumendjel, K (2004) Les calcaires griottes et faciès associés du Dévonien supérieur des environs de Beni-Abbes au Djebel Heche (Saoura, Algérie): environnements et implications paléogéographiques. Bulletin du Service géologique de l’Algérie 15, 27–49.Google Scholar
Bignon, A, Corbacho, J and López-Soriano, FJ (2014) A revision of the first Asteropyginae (Trilobita, Devonian). Geobios 47, 281–91.CrossRefGoogle Scholar
Bignon, A and Crônier, C (2015) Trilobite faunal dynamics on the Devonian continental shelves of the Ardenne Massif and Boulonnais (France, Belgium). Acta Palaeontologica Polonica 60, 949–62.Google Scholar
Bonelli, JR Jr and Patzkowsky, ME (2008) How are global patterns of faunal turnover expressed at regional scales? Evidence from the Upper Mississippian (Chesterian Series), Illinois Basin, USA. Palaios 23, 760–772.CrossRefGoogle Scholar
Boumendjel, K, Brice, D, Copper, P, Gourvennec, R, Jahnke, H, Lardeux, H, Le Menn, J, Melou, M, Morzadec, P, Paris, F, Plusquellec, Y and Racheboeuf, P (1997) Les faunes du Dévonien de l’Ougarta (Sahara occidental, Algérie). Annales Société géologique du Nord 5, 89–116.Google Scholar
Brachert, TC, Buggisch, W, Flügel, E, Hüssner, HM, Joachimski, MM, Tourneur, F and Walliser, OH (1992) Controls of mud mound formation: the Early Devonian Kess-Kess carbonates of the Hamar Laghdad, Antiatlas, Morocco. Geologische Rundschau 81, 15–44.CrossRefGoogle Scholar
Brenchley, PJ and Harper, DAT (1998) Palaeoecology: Ecosystems, Environments and Evolution. Oxford: Routledge, 402 pp.Google Scholar
Brett, CE, Ivany, LC and Schopf, KM (1996) Coordinated stasis: an overview. Palaeogeography, Palaeoclimatology, Palaeoecology 127, 1–20.CrossRefGoogle Scholar
Bultynck, P (1985) Lower Devonian (Emsian)–Middle Devonian (Eifelian and lowermost Givetian) conodont successions from the Ma’der and the Tafilalt, southern Morocco. Courier Forschungsinstitut Senckenberg 75, 261–86.Google Scholar
Bultynck, P and Walliser, OH (2000) Devonian boundaries in the Moroccan Anti-Atlas. Courier Forschungsinstitut Senckenberg 225, 211–24.Google Scholar
Cecca, F (2002) Palaeobiogeography of Marine Fossil Invertebrates: Concepts and Methods. London: Taylor and Francis, 273 pp.Google Scholar
Chatterton, BDE, Fortey, R, Brett, K, Gibb, S and McKellar, R (2006) Trilobites from the upper Lower to Middle Devonian Timrhanrhart Formation, Jebelgara el Zguilma, southern Morocco. Palaeontographica Canadiana 25, 1–177.Google Scholar
Chatterton, BDE and Gibb, S (2010) Latest Early to Early Middle Devonian trilobites from the Erbenochile Bed, Jbel Issoumour, Southeastern Morocco. Journal of Paleontology 84, 1188–205.CrossRefGoogle Scholar
Chatterton, BDE, Gibb, S and McKellar, RC (2020) Species of the Devonian aulacopleurid trilobite Cyphaspides from southeastern Morocco. Journal of Paleontology 94, 99–114.CrossRefGoogle Scholar
Chatterton, BDE, Siveter, D, Edgecombe, G and Hunt, A (1990) Larvae and relationships of the Calymenina (Trilobita). Journal of Paleontology 64, 255–77.CrossRefGoogle Scholar
Chatterton, BDE and Speyer, SE (1997) Ontogeny. In Treatise on Invertebrate Paleontology, Part O, Arthropoda 1, Trilobita, Revised (ed. Kaessler, RL), pp. 173–247. Boulder: Geological Society of America and Lawrence: University of Kansas Press.Google Scholar
Chlupáč, I (1977) The phacopid trilobites of the Silurian and Devonian of Czechoslovakia. Rozpravy Ústředního ústavu geologického 43, 1–172.Google Scholar
Chlupáč, I (1994) Devonian trilobites—evolution and events. Geobios 27, 487–505.CrossRefGoogle Scholar
Chlupáč, I, Luke, ÐP, Paris, F and Schönlaub, HP (1985) The Lochkovian–Pragian boundary in the Lower Devonian of the Barrandian area (Czechoslovakia). Jahrbuch der Geologischen Bundesanstalt–A 128, 9–41.Google Scholar
Clarke, KR (1993) Non-parametric multivariate analyses of changes in community structure. Australian Journal of Ecology 18, 117–43.CrossRefGoogle Scholar
Cohen, KM, Finney, SC, Gibbard, PL and Fan, J-X (2013) (updated) The ICS International Chronostratigraphic Chart. Episodes 36, 199–204.CrossRefGoogle Scholar
Crônier, C, Abbache, A, Khaldi, AY, Oudot, M, Maillet, S and Ouali Mehadji, A (2018a) Middle Devonian trilobites of the Saoura Valley, Algeria: insights into their biodiversity and Moroccan affinities. Geological Magazine 155, 811–40.CrossRefGoogle Scholar
Crônier, C, Bignon, A and François, A (2011) Morphological and ontogenetic criteria for defining a trilobite species: the example of Siluro-Devonian Phacopidae. Comptes Rendus Palevol 10, 143–53.CrossRefGoogle Scholar
Crônier, C and François, A (2014) Distribution patterns of Upper Devonian phacopid trilobites: paleobiogeographical and paleoenvironmental significance. Palaeogeography, Palaeoclimatology, Palaeoecology 404, 12–23.CrossRefGoogle Scholar
Crônier, C, Malti, FZ, Francois, A, Benyoucef, M and Brice, D (2013) First occurrence of a phacopid trilobite faunule from the Upper Devonian of Saoura Valley, Algeria and biodiversity fluctuations. Geological Magazine 150, 1002–21.CrossRefGoogle Scholar
Crônier, C, Oudot, M, Klug, C and de Baets, K (2018b) Trilobites from the Red Fauna (latest Emsian, Devonian) of Hamar Laghdad, Morocco and their biodiversity. Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen 290, 241–76.CrossRefGoogle Scholar
Crônier, C and van Viersen, A (2007) Trilobite palaeobiodiversity during the Devonian in the Ardennes Massif. Bulletin de la Société géologique de France 178, 473–83.CrossRefGoogle Scholar
Davis, JC (1986) Statistics and Data Analysis in Geology, 2nd Edition. New York: John Wiley and Sons, pp. 563–5.Google Scholar
de Baets, K, Klug, C and Plusquellec, Y (2010) Zlíchovian faunas with early ammonoids from Morocco and their use for the correlation between the eastern Anti-Atlas and the western Dra Valley. Bulletin of Geosciences 85, 317–52.CrossRefGoogle Scholar
Emmrich, HF (1839) De Trilobitis. Dissertatio Petrefactologica quam Consensu et Auctoritate Amplissimi Philosophorum Ordinis in Alma Litterarum Universitate Friderica Guilelma pro Summis in Philosophia Honoribus. Berlin: Berolini, 56 pp.Google Scholar
Feist, R (1991) The late Devonian trilobite crises. Historical Biology 5, 197–214.CrossRefGoogle Scholar
Feist, R (2002) Trilobites from the latest Frasnian Kellwasser Crisis in North Africa (Mrirt central Morocan Meseta). Acta Palaeontologica Polonica 47, 203–10.Google Scholar
Feist, R and Belka, Z (2018) Late Emsian (Devonian) trilobite communities from the Kess-Kess mounds, Hamar Laghdad (Anti-Atlas, Morocco). Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen 290, 277–90.CrossRefGoogle Scholar
Feist, R, Mahboubi, A and Girard, C (2016) New Late Devonian phacopid trilobites from Marhouma, SW Algerian Sahara. Bulletin of Geosciences 91, 243–59.CrossRefGoogle Scholar
Feist, R and Orth, B (2000) Trilobites de la limite Eifélien/Givétien de la région stratotypique (Tafilalet, Maider, Maroc). In Proceedings of the Subcommission on Devonian Stratigraphy (SDS)–IGCP 421 Morocco Meeting, pp. 78–91. Travaux de l’Institut Scientifique Rabat, Série Géologie & Géographie Physique 20.Google Scholar
Feist, R and Weyer, D (2018) The proetid trilobite Perliproetus, a marker of the late Famennian in Central Europe and North Africa. Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen 287, 195–206.CrossRefGoogle Scholar
Frey, L, Naglik, C, Hofmann, R, Schemm-Gregory, M, Frýda, J, Kröger, B, Taylor, PD, Wilson, MA and Klug, C (2014) Diversity and palaeoecology of invertebrate associations of the Early Devonian in the Tafilalt (Morocco, Anti-Atlas). Bulletin of Geoscience 89, 75–112.Google Scholar
Frey, L, Pohle, A, Rücklin, M and Klug, C (2019) Fossil-Lagerstätten, palaeoecology and preservation of invertebrates and vertebrates from the Devonian in the eastern Anti-Atlas (Morocco). Lethaia 53, 242–66.CrossRefGoogle Scholar
Gaston, KJ, Blackburn, TM, Greenwood, JJ, Gregory, RD, Quinn, RM and Lawton, JH (2000) Abundance–occupancy relationships. Journal of Applied Ecology 37, 39–59.CrossRefGoogle Scholar
Gibb, S and Chatterton, BD (2010) Gerastos (Order Proetida; Class Trilobita) from the Lower to Middle Devonian of the southern Moroccan Anti-Atlas region. Palaeontographica Canadiana 30, 1–89.Google Scholar
Gleason, HA (1927) Further views on the succession-concept. Ecology 8, 299–326.CrossRefGoogle Scholar
Greenacre, MJ (1984) Theory and Applications of Correspondence Analysis. London: Academic Press.Google Scholar
Hammer, Ø and Harper, DA (2008) Paleontological Data Analysis. Oxford: Blackwell Publishing.Google Scholar
Hammer, Ø, Harper, DA and Ryan, PD (2001) PAST: paleontological statistics software package for education and data analysis. Palaeontologia Electronica 4, 1–9.Google Scholar
Harnik, PG (2009) Unveiling rare diversity by integrating museum, literature, and field data. Paleobiology 35, 190–208.CrossRefGoogle Scholar
Harper, DAT (ed.) (1999) Numerical Palaeobiology: Computer Based Modelling and Analysis of Fossils and their Distributions. New York: John Wiley and Sons.Google Scholar
Holland, SM, Miller, AI, Meyer, DL and Dattilo, BF (2001) The detection and importance of subtle biofacies within a single lithofacies: the Upper Ordovician Kope Formation of the Cincinnati, Ohio region. Palaios 16, 205–17.2.0.CO;2>CrossRefGoogle Scholar
Hollard, H (1963) Un tableau stratigraphique du Dévonien du Sud de l’Anti-Atlas. Notes du Service géologique du Maroc 23, 131–6.Google Scholar
Hollard, H (1967) Le Dévonien du Maroc et du Sahara nord-occidental. In International Symposium on the Devonian System, Calgary, 1967, Volume 1 (ed. Oswald, DH), pp. 203–44. Alberta Society of Petroleum Geologists.Google Scholar
Hollard, H (1974) Recherches sur la stratigraphie des formations du Dévonien moyen, de l’Emsien superieur au Frasnien, dans le sud du Tafilalt et dans le Ma’der (Anti-Atlas Oriental). Notes du Service géologique du Maroc 36, 7–68.Google Scholar
Hollard, H (1981) Principaux caractères des formations dévoniennes de l’Anti-Atlas. Tableaux de correlation du Silurien et du Dévonien de l’Anti-Atlas. Notes du Service géologique du Maroc 42, 15–23.Google Scholar
House, MR (1985) Correlation of mid-Palaeozoic ammonoid evolutionary events with global sedimentary perturbations. Nature 313, 17–22.CrossRefGoogle Scholar
House, MR (2002) Strength, timing, setting and cause of mid-Palaeozoic extinctions. Palaeogeography, Palaeoclimatology, Palaeoecology 181, 5–25.CrossRefGoogle Scholar
Hüneke, H (2006) Erosion and deposition from bottom currents during the Givetian and Frasnian: response to intensified oceanic circulation between Gondwana and Laurussia. Palaeogeography, Palaeoclimatology, Palaeoecology 234, 146–67.CrossRefGoogle Scholar
Jansen, U, Becker, G, Plodowski, G, Schindler, E, Vogel, O and Weddige, K (2004) The Emsian to Eifelian near Foum Zguid (NE Dra Valley, Morocco). In Devonian Neritic–Pelagic Correlation and Events in the Dra Valley (Western Anti-Atlas, Morocco). Subcommission on Devonian Stratigraphy, International Meeting on Stratigraphy, Rabat, 1–10 March 2004 (ed. El Hassani, A), pp. 19–28. Document de l’Institut Scientifique, Rabat 19.Google Scholar
Johnson, RG and Fortey, RA (2012) Proetid Trilobites from the Lower Devonian (Pragian) Ihandar Formation, Anti-Atlas, Morocco. Journal of Paleontology 86, 1032–50.CrossRefGoogle Scholar
Johnson, JG, Klapper, G and Sandberg, CA (1985) Devonian eustatic fluctuations in Euramerica. Geological Society of America Bulletin 96, 567–87.2.0.CO;2>CrossRefGoogle Scholar
Jongman, E and Jongman, SRR (1995) Data Analysis in Community and Landscape Ecology. Cambridge: Cambridge University Press.CrossRefGoogle Scholar
Kaiser, SI, Becker, RT, Steuber, T and Aboussalam, ZS (2011) Climate-controlled mass extinctions, facies, and sea-level changes around the Devonian–Carboniferous boundary in the eastern Anti-Atlas (SE Morocco). Palaeogeography, Palaeoclimatology, Palaeoecology 310, 340–64.CrossRefGoogle Scholar
Kaufmann, B (1998) Facies, stratigraphy and diagenesis of Middle Devonian reef-and mud-mounds in the Mader (eastern Anti-Atlas, Morocco). Acta Geologica Polonica 48, 43–106.Google Scholar
Khaldi, AY, Crônier, C, Hainaut, G, Abbache, A and Ouali Mehadji, A (2016) A trilobite faunule from the Lower Devonian of the Saoura Valley, Algeria: biodiversity, morphological variability and palaeobiogeographical affinities. Geological Magazine 153, 357–87.CrossRefGoogle Scholar
Klug, C, Frey, L, Korn, D, Jattiot, R and Rücklin, M (2016) The oldest Gondwanan cephalopod mandibles (Hangenberg Black Shale, Late Devonian) and the mid-Palaeozoic rise of jaws. Palaeontology 59, 611–29.CrossRefGoogle Scholar
Klug, C, Korn, D, Naglik, C, Frey, L and de Baets, K (2013) The Lochkovian to Eifelian succession of the Amessoui Syncline (southern Tafilalt). In Excursion Guidebook, International Field Symposium “The Devonian and Lower Carboniferous of Northern Gondwana”, Morocco 2013 (eds Becker, RT, El Hassani, A and Tahiri, A), pp. 51–60. Document de l‘Institut Scientifique, Rabat 27.Google Scholar
Klug, C, Kröger, B, Korn, D, Ruecklin, M, Schemm-Gregory, M, de Baets, K and Mapes, RH (2008) Ecological change during the early Emsian (Devonian) in the Tafilalt (Morocco), the origin of the Ammonoidea, and the first African pyrgocystid edrioasteroids, machaerids and phyllocarids. Palaeontographica A 283, 1–94.Google Scholar
Klug, C, Samankassou, E, Pohle, A, de Baets, K, Franchi, F and Korn, D (2018) Oases of biodiversity: Early Devonian palaeoecology at Hamar Laghdad, Morocco. In Palaeontology of the Devonian of Hamar Laghdad (Tafilalt, Morocco). Special Volume Honouring Jobst Wendt (eds Klug, C and Korn, D). Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen 290, 9–48.Google Scholar
Korn, D and Bockwinkel, J (2017) The genus Gonioclymenia (Ammonoidea; Late Devonian) in the Anti-Atlas of Morocco. Neues Jahrbuch für Geologie und Paläontologie Abhandlungen 285, 97–115.CrossRefGoogle Scholar
Le Maitre, D (1952) La faune du dévonien inférieur et moyen de la Saoura et des abords de l’Erg el Djemal (Sud-Oranais). Matériaux pour la carte géologique de l’Algérie, Paléontologie 12, 1–170.Google Scholar
Legendre, P and Legendre, L (1998) Numerical Ecology, 2nd Edition. Developments in Environmental Modelling 20. Amsterdam: Elsevier Science, 870 pp.Google Scholar
Lerosey-Aubril, R and Feist, R (2005) Ontogeny of a new cyrtosymboline trilobite from the Famennian of Morocco. Acta Palaeontologica Polonica 50, 449–64.Google Scholar
Lerosey-Aubril, R and Feist, R (2006) Late ontogeny and hypostomal condition of a new cyrtosymboline trilobite from the Famennian of Morocco. Palaeontology 49, 1053–68.CrossRefGoogle Scholar
Lerosey-Aubril, R and Feist, R (2012) Quantitative approach to diversity and decline in Late Palaeozoic trilobites. In Global Biodiversity, Extinction Intervals and Biogeographic Perturbations through Time (ed. Talent, JA), pp. 535–55. Berlin: Springer Science.CrossRefGoogle Scholar
Limam, H, Mehadji, AO and Bendella, M (2021) The Chefar El Ahmar Formation (late Emsian–Frasnian Ib) in the Marhouma section (Saoura, Algerian Sahara): insights from ichnology and sedimentology. Arabian Journal of Geosciences 14, 274. doi: 0.1007/s12517-021-06669-4.CrossRefGoogle Scholar
Lubeseder, S, Carr, ID and Redfern, J (2003) A third-order sequence stratigraphic framework for the Devonian of Morocco: its implications for enhanced regional correlation of the Devonian in North Africa. In AAPG Hedberg Conference, Paleozoic and Triassic Petroleum Systems in North Africa, 18–20 February 2003, Algiers, Algeria, pp. 1–4.Google Scholar
Lubeseder, S, Rath, J, Rücklin, M and Messbacher, R (2010) Controls on Devonian hemi-pelagic limestone deposition analyzed on cephalopod ridge to slope sections, Eastern Anti-Atlas, Morocco. Facies 56, 295–315.CrossRefGoogle Scholar
Maillet, S, Tanjaoui-Arif, R, Milhau, B, Nicollin, J-P and Ouali Mehadji, A (2013) Ostracodes of the Chefar el Ahmar Formation (Middle Devonian) in the Km 30 section (Beni-Abbes area, Saoura, Ougarta Range, Algeria). Revue de Paléobiologie, Genève 32, 465–80.Google Scholar
Mángano, MG, Buatois, LA, Waisfeld, BG, Muñoz, DF, Vaccari, NE and Astini, RA (2021) Were all trilobites fully marine? Trilobite expansion into brackish water during the early Palaeozoic. Proceedings of the Royal Society B 288, 20202263. doi: 10.1098/rspb.2020.2263.CrossRefGoogle ScholarPubMed
Massa, D (1965) Observations sur les Séries Siluro-Dévoniennes des confins Algéro-Marocains du sud (1954–1955). Paris: Notes et Mémoires Compagnie Française des Pétroles 8, 187 pp.Google Scholar
McKellar, RC and Chatterton, BD (2009) Early and Middle Devonian Phacopidae (Trilobita) of southern Morocco. Palaeontographica Canadiana 28, 1–110.Google Scholar
Michard, A, Saddiqi, O, Chalouan, A and de Lamotte, DF (2008) Continental Evolution: The Geology of Morocco: Structure, Stratigraphy, and Tectonics of the Africa-Atlantic-Mediterranean Triple Junction. Berlin, Heidelberg: Springer-Verlag, pp. 65–132.CrossRefGoogle Scholar
Morzadec, P (1988) Le genre Psychopyge (Trilobita) dans le Dévonien Inférieur du Nord de l’Afrique et l’Ouest de l’Europe. Palaeontographica, Abteilung A 200, 153–61.Google Scholar
Morzadec, P (1992) Evolution des Asteropyginae (Trilobita) et variations eustatiques au Dévonien. Lethaia 25, 85–96.CrossRefGoogle Scholar
Morzadec, P (1997) Asteropyginae trilobites from the Devonian of the Ougarta (Algeria) Palaeontographica, Abteilung A 244, 143–58.CrossRefGoogle Scholar
Morzadec, P (2001) Les Trilobites Asteropyginae du Dévonien de l’Anti-Atlas (Maroc). Palaeontographica, Abteilung A 262, 53–85.CrossRefGoogle Scholar
Mottequin, B, Malti, FZ, Benyoucef, M, Crônier, C, Samar, L, Randon, C and Brice, D (2015) Famennian rhynchonellides (Brachiopoda) from deep-water facies of the Ougarta Basin (Saoura Valley, Algeria). Geological Magazine 152, 1009–24.CrossRefGoogle Scholar
Ouali Mehadji, A, Atif, KFT, Bouterfa, B, Nicollin, J-P and Besseghier, FZ (2011) Environnements sédimentaires de la Saoura-Ougarta (Sahara Nord-Ouest, Algérie) au Dévonien inférieur (Lochkovien supérieur pro parte-Emsien). Geodiversitas 33, 553–80.CrossRefGoogle Scholar
Ouali Mehadji, A, Bendella, M, Atif, K, Belkacemi, K and Abbache, A (2012) Estimation de l’épaisseur réelle et des environnements de depots d’âge famennien (Dévonien supérieur) de la region de Marhouma (Saoura, Sahara nord-Ouest, Algérie). Bulletin du Service géologique national 23, 3–27.Google Scholar
Ouanaimi, H and Lazreq, N (2008) The ‘Rich’ group of the Draa Basin (Lower Devonian, Anti-Atlas, Morocco): an integrated sedimentary and tectonic approach. In The Boundaries of the West African Craton (eds Ennih, N and Liégeois, JP), pp. 467–82. Geological Society of London, Special Publication no. 297.Google Scholar
R Core Team (2020) R: A Language and Environment for Statistical Computing. Vienna, Austria: R Foundation for Statistical Computing. http://www. Rproject.org/Google Scholar
Qian, H, Ricklefs, RE and White, PS (2005) Beta diversity of angiosperms in temperate floras of eastern Asia and eastern North America. Ecology Letters 8, 15–22.CrossRefGoogle Scholar
Schraut, G and Feist, R (2004) The Devonian styginid trilobite Paralejurus, with new data from Spain and Morocco. Journal of Paleontology 78, 709–22.2.0.CO;2>CrossRefGoogle Scholar
Sørensen, TJ (1948) A method of establishing groups of equal amplitude in plant sociology based on similarity of species content and its application to analyses of the vegetation on Danish commons. Det Kongelige Danske Videnskabers Selskabs Skrivter 5, 1–34.Google Scholar
Speyer, SE and Chatterton, BDE (1989) Trilobite larvae and larval ecology. Historical Biology 3, 27–60.CrossRefGoogle Scholar
Termier, G and Termier, H (1950) Paléontologie marocaine, t. II, Invertébrés de l’Ere primaire: Annélides, Arthropodes, Echinodermes, Conularides et Graptolithes. Notes et Mémoires du service géologique du Maroc 79, 1–279.Google Scholar
Thompson, WL (2004) Sampling Rare or Elusive Species. Washington D.C.: Island Press, 429 pp.Google Scholar
Toto, EA, Kaabouben, F, Zouhri, L, Belarbi, M, Benammi, M, Hafid, M and Boutib, L (2008) Geological evolution and structural style of the Palaeozoic Tafilalt sub-basin, eastern Anti-Atlas (Morocco, North Africa). Geological Journal 43, 59–73.CrossRefGoogle Scholar
Ulrich, W and Gotelli, NJ (2007) Null model analysis of species nestedness patterns. Ecology 88, 1824–31.CrossRefGoogle ScholarPubMed
van Viersen, AP and Holland, D (2016) Morphological trends and new species of Cyphaspis (Trilobita, Otarioninae) in the Devonian of Morocco, Turkey, Germany and Belgium. Geologica Belgica 19, 251–71.CrossRefGoogle Scholar
van Viersen, AP and Lerouge, F (2019) Cornuproetine (proetide) trilobites with nine thorax segments from the Devonian of Morocco, Germany and the Czech Republic. Paläontologische Zeitschrift 94, 227–54.CrossRefGoogle Scholar
Wendt, J (1985) Disintegration of the continental margin of northwestern Gondwana: Late Devonian of the eastern Anti-Atlas (Morocco). Geology 13, 815–8.2.0.CO;2>CrossRefGoogle Scholar
Wendt, J (1988) Facies pattern and paleogeography of the Middle and Late Devonian in the eastern Anti-Atlas (Morocco). Canadian Society of Petroleum Geologists, Memoir 14, 467–80.Google Scholar
Wendt, J (1993) Steep-sided carbonate mud mounds in the Middle Devonian of the eastern Anti-Atlas, Morocco. Geological Magazine 130, 69–83.CrossRefGoogle Scholar
Wendt, J (1995) Shell directions as a tool in palaeocurrent analysis. Sedimentary Geology 95, 161–86.CrossRefGoogle Scholar
Wendt, J (2021a) Middle and Late Devonian paleogeography of the eastern Anti-Atlas (Morocco). International Journal of Earth Sciences 110, 1531–44.CrossRefGoogle Scholar
Wendt, J (2021b) Middle and Late Devonian sea-level changes and synsedimentary tectonics in the eastern Anti-Atlas (Morocco). Journal of African Earth Sciences 182, 1–16.CrossRefGoogle Scholar
Wendt, J, Aigner, T and Neugebauer, J (1984) Cephalopod limestone deposition on a shallow pelagic ridge: the Tafilalt Platform (upper Devonian, eastern Anti-Atlas, Morocco). Sedimentology 31, 601–25.CrossRefGoogle Scholar
Wendt, J and Belka, Z (1991) Age and depositional environment of Upper Devonian (early Frasnian to early Famennian) black shales and limestones (Kellwasser facies) in the eastern Anti-Atlas, Morocco. Facies 25, 51–89.CrossRefGoogle Scholar
Wendt, J, Kaufmann, B, Belka, Z, Klug, C and Lubeseder, S (2006) Sedimentary evolution of a Palaeozoic basin and ridge system: the Middle and Upper Devonian of the Ahnet and Mouydir (Algerian Sahara). Geological Magazine 143, 269–99.CrossRefGoogle Scholar
Whittaker, RH (1960) Vegetation of the Siskiyou Mountains, Oregon and California. Ecological Monographs 30, 279–338.CrossRefGoogle Scholar
Wright, DH and Reeves, JH (1992) On the meaning and measurement of nestedness of species assemblages. Oecologia 92, 416–28.CrossRefGoogle ScholarPubMed
Zapalski, MK and Klug, K (2018) Trilobite sclerites as attachment surface for Emsian tabulate corals of Hamar Laghdad (Anti-Atlas, Morocco). Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen 290, 111–15.CrossRefGoogle Scholar
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