Diversity and palaeoecology of Early Devonian invertebrate associations in the Tafilalt (Anti-Atlas, Morocco)


Authors: Frey L, Naglik C, Hofmann R, Schemm-Gregory M, Frýda J, Kröger B, Taylor PD, Wilson MA, Klug C

Published in: Bulletin of Geosciences, volume 89, issue 1; pages: 75 - 112; Received 13 June 2013; Accepted in revised form 29 October 2013; Online 12 December 2013

Keywords: Early Devonian, alpha diversity, palaeoecology, Cephalopoda, Gastropoda, Anti-Atlas, Morocco,

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Quantitative analyses of the taxonomic composition and palaeoecology of five Early Devonian faunules (earliest Lochkovian to early Emsian) collected from the locality Jebel Ouaoufilal in the Tafilalt (Morocco) were conducted. We examined 3376 specimens belonging to 158 species and their stratigraphic distribution. The quantitative data sets were analysed for alpha diversity and ecospace utilization. Macrofossils of every faunule were identified, counted and grouped according to ecological categories of tiering, motility and feeding behaviour. Based on these data, we noted (i) a strong increase in species richness, especially in benthic species, from the lowermost Lochkovian to the Pragian stage and a following subtle decrease in the early Emsian and (ii) a considerable expansion of ecospace use from the earliest Lochkovian to the early Emsian. These general trends are considered to be a result of favourable living conditions due to a growing oxygen content at the sea floor, which correlates with a regional and potentially a global regression from the Lochkovian to the Pragian. A transgression in the early Zlíchovian probably reduced the species richness by decreasing the oxygenation of bottom waters. Additionally, we described and figured three new genera and six new species of the diverse Pragian faunule (86 species): the hederelloid Filihernodia buccina gen. et sp. nov., the crinoid Hexawacrinus claudiakurtae gen. et sp. nov., the gastropods Oriomphalus multiornatus sp. nov., Eohormotomina restisevoluta gen. et sp. nov., and the cephalopods Tenuitheoceras secretum gen. et sp. nov. and Arionoceras kennethdebaetsi sp. nov. We record the first occurrence of Tiaracrinus moravicus from Africa.


Alberti , G.K.B. 1969. Trilobiten des jüngeren Siluriums sowie des Unter- und Mitteldevons; I. Mit einigen Beiträgen zur Silur-Devon Stratigraphie einiger Gebiete Marokkos und Oberfrankens. Abhandlungen der Seckenbergischen naturforschenden Gesellschaft 520, 1-692.

Alberti, G.K.B. 1980. Neue Daten zur Grenze Unter-/Mittel-Devon, vornehmlich aufgrund der Tentaculiten und Trilobiten im Tafilalt (Marrokko). Neues Jahrbuch für Geologie und Paläontologie, Monatshefte 1980, 581-594.View article

Alberti, G.K.B. 1981. Daten zur stratigraphischen Verbreitung der Nowakiidae (Dacryoconarida) im Devon von NW-Afrika (Marokko, Algerien). Senckenbergiana lethaea 62, 205-216.

Alberti, G.K.B. 1983. Trilobiten des jüngeren Siluriums sowie des Unter- und Mittel-Devons; IV. Senckenbergiana lethaea 61(1), 1-87.

Ausich, W.I. & Bottjer, D.J. 1982. Tiering in suspension-feeding communities on soft substrata throughout the Phanerozoic. Science New Series 216(4542), 173-174.

Austin, T. & Austin, T. Jr. 1843. A Description of several new genera and species of Crinoidea. Annals and Magazine of Natural History (series 1) 11(69), 195-207.View article

Barrande, J. 1846. Notice préliminaire sur le systeme silurien et les trilobites de Boheme. 86 pp. Hirschfeld, Leipzig.View article

Barrande, J. 1866. Systeme silurien du centre de la Boheme, I. Partie, Vol. II: Céphalopodes 2. Pl. 108-244. Privately published, Prague & Paris.

Barrande, J. 1867. Systeme silurien du centre de la Boheme, I. Partie, Vol. II: Céphalopodes 1. 712 pp. Privately published, Prague & Paris.

Barrande, J. 1868. Systeme silurien du centre de la Boheme, I. Partie, Vol. II: Céphalopodes 3. Pl. 245-350. Privately published, Prague & Paris.

Barrande, J. 1870. Systeme silurien du centre de la Boheme, I. Partie, Vol. II: Céphalopodes 4. Pl. 351-460. Privately published, Prague & Paris.

Barrande, J. 1881. Systeme silurien du centre de la Boheme. Classe des Mollusques, ordre des Acéphalés 6. 342 pp. Privately published, Prague & Paris.

Barskov, I.S. 1963. System and phylogeny of the Pseudorthoceratids. Byulleten Moskovskogo obshchevestva ispytatelei prirody, Otdel geologicheskii 38(4), 149-150.

Barskov, I.S. 1966. Cephalopods of the Late Ordovician and Silurian of Kazakhstan and Middle Asia. Autoreferat dissertatcii na soiskanie utchenoyi stepeni kandidata geologicheskii mineralogicheskii nauka. 200 pp. Izdatel’stvo Moskovskogo Universiteta, Moscow.

Bassler, R.S. 1935. Bryozoa (generum et gentyporum index et bibliographia). Fossilium Catalogus. I: Animalia Pars 67, 1-229.

Bassler, R.S. 1939. The Hederelloidea, a suborder of Paleozoic cyclostomatous Bryozoa. Proceedings of the United States National Museum 87, 25-91.View article

Bassler, R.S. 1953. Bryozoa, G1-G253. In Moore, R.C. (ed.) Treatise on Invertebrate Paleontology, Part G. Geological Society of America and University of Kansas Press, Lawrence.

Becker, R.T. & House, M.R. 1994. International Devonian goniatite zonation, Emsian to Givetian, with new records from Morocco. Courier Forschungsinstitut Senckenberg 169, 79-135.

Becker, R.T. & House, M.R. 2000. Emsian and Eifelian ammonoid succession at Bou Tchrafine (Tafilalt platform, Anti-Atlas, Morocco), 21-26. In El Hassani, A. & Tahiri, A. (eds) Moroccan meeting of the Subcommision on Devonian Stratigraphy (SDS) - IGCP 421 Excursion Guidebook, Notes et Mémoirs du Service Géologique 399.

BELKA, Z., KAUFMANN, B. & BULTYNCK, P. 1997. Conodont-based quantitative biostratigraphy for the Eifelian of the eastern Anti-Atlas, Morocco. Geological Society of American Bulletin 109, 643-651. View article

BELKA, Z., KLUG, C., KAUFMANN, B., KORN, D., DÖRING, S., FEIST, R. & WENDT, J. 1999. Devonian conodont and ammonoid succession of the eastern Tafilalt (Ouidane Chebbi section), Anti-Atlas, Morocco. Acta Geologica Polonica 49(1), 1-23.

BLODGETT, R.B., FRÝDA, J. & RACHEBOEUF, P.R. 1999. Upper Middle Devonian (Givetian) gastropods from the Kersiadou Formation, Brittany, France. Journal of Paleontology 73(6), 1081-1100.

BLODGETT, R.B., ROHR, D.M. & BOUCOT, A.J. 1988. Lower Devonian gastropod biogeography of the Western Hemisphere, 281-294. In MCMILLAN, N.J., EMBRY, A.F. & GLASS, D.J. (eds) Devonian of the World. Canadian Society of Petroleum Geologists Memoir 14(3).

BLODGETT, R.B., ROHR, D.M. & BOUCOT, A.J. 1990. Early and Middle Devonian biogeography, 277-284. In MCKERROW, W.S. &. SCOTESE, C.D. (eds) Palaeozoic Palaeogeography and Biogeography. Geological Society Memoir 12.

BOTTJER, D.J. & AUSICH, W.I. 1986. Phanerozoic development of tiering in soft substrata suspension-feeding communities. Paleobiology 12(4), 400-420.View article

BOUCHET, P., ROCROI, J. P., FRÝDA, J., HAUSDORF, B., PONDER, W., VALDES, A. & WARÉN, A. 2005. Classification and nomenclator of gastropod families. Malacologia 47, 1-368.

BRACHERT, T.C., BUGGISCH, W., FLÜGEL, E., HÜSSNER, H., JOACHIMSKI, M.M., TOURNEUR, F. & WALLISER, O.H. 1992. Controls of mud mound formation: the Early Devonian Kess-Kess carbonates of the Hamar Laghdad, Anti-Atlas, Morocco. Geologische Rundschau 81, 15-44.View article

BRONGNIART, A. 1822. Les Trilobites, 1-65. In BRONGNIART, A. & DESMAREST, A.G. (eds) Histoire naturelle des crustacés fossiles, sous les rapports zoologiques et géologiques. F.G. Levrault, Paris & Strasbourg.View article

BULTYNCK, P. & WALLISER, O.H. 2000a. Emsian to Middle Frasnian sections in the northern Tafilalt, 11-20. In EL HASSANI, A. & TAHIRI, A. (eds) Moroccan meeting of the Subcommision on Devonian Stratigraphy (SDS) - IGCP 421 Excursion Guidebook, Notes et Mémoires du Service Géologique 399.

BULTYNCK, P. & WALLISER, O.H. 2000b. Devonian boundaries in the Moroccan Anti-Atlas. Courier Forschungsinstitut Senckenberg 225, 211-226.

BUSH, A.M., BAMBACH, R.K. & DALEY, G.M. 2007. Changes in theoretical ecospace utilization in marine fossil assemblages between the mid-Paleozoic and late Cenozoic. Paleobiology 33(1), 76-97.View article

CHLUPÁČ, I., GALLE, A., HLADIL, J. & KALVODA, J. 2000. Series and stage boundaries in the Devonian of the Czech Republic. Courier Forschungsinstitut Senckenberg 225, 159-172.

CHLUPÁČ, I. & KUKAL, Z. 1988. Possible global events and the stratigraphy of the Palaeozoic of the Barrandian (Cambrian-Middle Devonian, Czechoslovakia). Sborník geologických věd, Geologie 43, 83-146.

CHLUPÁČ, I. & KUKAL, Z. 1996. Reflection of possible global Devonian events in the Barrandian area, C.S.S.R, 169-179. In WALLISER, O.H. (ed.) Global Bio-events, a critical approach. Lecture Notes on Earth Sciences 8.

CLARIOND, L. 1934a. Sur le Dévonien du Tafilalet et du Maroc. Compte rendu sommaire des séances de la Société Géologique de France 1934, 5-11.

CLARIOND, L. 1934b. A propos d’une coupe de la région d’Erfoud. Compte rendu sommaire des séances de la Société Géologique de France 1934, 223-224.

CORRIGA, M.G., CORRADINI, C., HAUDE, R. & WALLISER, O.H. 2013. Upper Silurian and Lower Devonian conodonts and crinoids from the scyphocrinoid beds of southeastern Morocco, 71-72. In LINDSKOG, A. & MEHLQVIST, K. (eds) Proceedings of the 3rd IGCP 591 Annual Meeting, Lund, Sweden.

COX, L.R. 1960. Gastropoda. General characteristics of Gastropoda, I84-I169. In MOORE, R.C. (ed.) Treatise on Invertebrate Paleontology, Part I, Mollusca 1. Geological Society of America & University of Kansas Press, Boulder & Lawrence.

CUVIER, G. 1797. Tableau élémentaire de l’histoire naturelle des animaux. 710 pp. Baudouin, Paris.View article

DANA, J.D. 1846. Structure and classification of zoophytes: U.S. Exploring Expedition during the years 1838-1842 under the command of Charles Wilkes, U.S.N. 7. 740 pp. Lea & Blanchard, Philadelphia.

DE BAETS, K., KLUG, C. & MONNET, C. 2013. Intraspecific variability through ontogeny in early ammonoids. Paleobiology 39(1), 75-94.View article

DE BAETS, K., KLUG, C. & 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(2), 317-352.View article

DONOVAN, S.K. & WEBSTER, G.D. 2013. Platyceratid gastropod infestations of Neoplatycrinus Wanner (Crinoidea) from the Permian of West Timor: speculations on thecal modifications. Proceedings of the Geologists’ Association. View article

DÖRING, A. 2002. Sedimentological Evolution of the late Emsian to early Givetian carbonate ramp in the Mader (eastern Anti-Atlas, SE-Morocco). 80 pp. Ph.D. thesis, Eberhard-Karls-Universität, Tübingen, Germany.

DZIK, J. 1984. Phylogeny of the Nautiloidea. Palaeontologia Polonica 45, 1-203.

ELDREDGE, N. 1971. Patterns in cephalic musculature in the Phacopina (Trilobita) and their phylogenetic significance. Journal of Palaeontology 45, 52-67.

ETHERIDGE, R. 1890. Descriptions of Upper Silurian fossils from the Lilydale Limestone , Upper Yarra District, Victoria. Records of the Australian Museum 1(1), 60-67.View article

ETHERIDGE, R. 1891. Further descriptions of the Upper Silurian fossils from the Lilydale Limestone, Upper Yarra District, Victoria. Records of the Australian Museum 1(7), 125-130.View article

FERRETI, A., GNOLI, M. & VAI, G.B. 1999. Silurian to Lower Devonian communities of Sardinia, 271-281. In BOUCOT, A.J. & LAWSON, J.D. (eds) Paleocommunities - a case study from the Silurian and Lower Devonian. Cambridge University Press, Cambridge.

FISCHER, P. 1882. Manuel de conchyliologie et de paléontologie conchyliologique ou histoire naturelle des mollusques vivants et fossiles, I. Partie: Synopsis des genres, 327-418. Savy, Paris.

FLOWER , R.H. & CASTER, K.E. 1935. The cephalopod fauna of the Conewango Series of the Upper Devonian in New York and Pennsylvania. Bulletins of American Paleontology 22(75), 1-74.

FLOWER, R.H. & KUMMEL, B. 1950. A classification of the Nautiloidea. Journal of Paleontology 24, 604-616.

FORTEY, R.A. & OWENS, R.M. 1999. Feeding habits in trilobites. Palaeontology 42(3), 429-465.View article

FRANCHI, F., SCHEMM-GREGORY, M. & KLUG, C. 2012. A new species of Ivdelinia Andronov, 1961 and its palaeoecological and palaeobiogeographical implications (Morocco, Givetian). Bulletin of Geosciences 87(1), 1-11.View article

FRÖHLICH, S. 2004. Evolution of a Devonian carbonate shelf at the northern margin of Gondwana (Jebel Rheris, eastern Anti-Atlas, Morocco). 71 pp. Ph.D. thesis, Eberhard-Karls-Universität, Tübingen, Germany.

FRÝDA, J. 1993. Oldest representative of the family Palaeozygopleuridae (Gastropoda) with notes on its higher taxonomy. Journal of Paleontology 67, 822-827. View article

FRÝDA, J. 1997. Oldest representatives of the superfamily Cirroidea (Vetigastropoda) with notes on their early phylogeny. Journal of Paleontology 71(5), 839-847.View article

FRÝDA, J. 1998a. Some new and better recognized Devonian gastropods from the Prague Basin (Bohemia). Bulletin of the Czech Geological Survey 73(1), 41-49.

FRÝDA, J. 1998b. Some new and better recognized Devonian gastropods from the Prague Basin (Bohemia): part II. Bulletin of the Czech Geological Survey 73(4), 355-363.

FRÝDA, J. 1999. Further new gastropods from the Early Devonian Boucotonotus - Palaeozygopleura Community of the Prague Basin. Journal of the Czech Geological Society 44, 317-325.

FRÝDA, J. & BANDEL, K. 1997. New Early Devonian gastropods from the Plectonotus (Boucotonotus)-Palaeozygopleura Community in the Prague Basin (Bohemia). Mitteilungen aus dem Geologisch-Paläontologischen Institut der Universität Hamburg 80, 1-58.

FRÝDA, J. & BLODGETT, R.B. 1998. Two new cirroidean genera (Vetigastropoda, Archaeogastropoda) from the Emsian (late Early Devonian) of Alaska with notes on the early phylogeny of Cirroidea. Journal of Paleontology 72, 265-273.View article

FRÝDA, J. & BLODGETT, R.B. 2004. New Emsian (late Early Devonian) gastropods from Limestone Mountain, Medfra B-4 quadrangle, west-central Alaska (Farewell terrane), and their paleobiogeographic affinities and evolutionary significance. Journal of Paleontology 78(1), 111-132.View article

FRÝDA, J., FERROVÁ, L., BERKYOVÁ, S. & FRÝDOVÁ, B. 2008. A new Early Devonian palaeozygopleurid gastropod from the Prague Basin (Bohemia) with notes on the phylogeny of the Loxonematoidea. Bulletin of Geosciences 83(1), 93-100.View article

FRÝDA, J., FERROVÁ, L. & FRÝDOVÁ, B. 2013. Review of palaeozygopleurid gastropods (Palaeozygopleuridae, Gastropoda) from Devonian strata of the Perunica microplate (Bohemia), with a re-evaluation of their stratigraphic distribution, notes on their ontogeny, and descriptions of new taxa. Zootaxa 3669(4), 469-489.View article

FRÝDA, J. & MANDA, Š. 1997. A gastropod faunule from the Monograptus uniformis graptolite Biozone (Early Lochkovian, Early Devonian) in Bohemia. Mitteilungen aus dem Geologisch-Paläontologischen Institut der Universität Hamburg 80, 59-122.

GAHN, F.J. & BAUMILLER, T.K. 2003. Infestation of Middle Devonian (Givetian) camerate crinoids by platyceratid gastropods and its implications for the nature of their biotic interaction. Lethaia 36, 71-82.View article

GAHN, F.J. & BAUMILLER, T.K. 2006. Using platyceratid gastropod behaviour to test functional morphology. Historical Biology 18, 397-404.View article

GARCÍA-ALCADE, J., TRUYÓLS-MASSONI, M., PARDO-ALONSO, M., BULTYNCK, P. & CARLS, P. 2000. Devonian chronostratigraphy of Spain. Courier Forschungsinstitut Senckenberg 225, 131-144.

GRABAU, A.W. & SHIMER, H.W. 1909. North American index fossils: Invertebrates 1. 853 pp. New York. View article

GUBANOV, A.P. & YOCHELSON, E.L. 1994. A Wenlockian (Silurian) gastropod shell and operculum from Siberia. Journal of Paleontology 68, 486-481.View article

HALL, J. 1852. Containing descriptions of the organic remains of the lower middle division of the New York system. Paleontology of New York 2, 1-362.

HALL, J. 1861. Descriptions of new species of fossils from the Upper Helderberg, Hamilton and Chemung Groups; with observations upon previously described species. Fourteenth Annual Report of the Regents of the University of the State of New York on the condition of the State Cabinent of Natural History and the Historical and Antiquarian Collection annexed thereto, Albany, 99-109.

HALL, J. 1881. Bryozoans of the Upper Helderberg and Hamilton groups. Transactions of the Albany Institute 10, 145-197.

HALL, J. 1884. Lamellibranchiata, I. Monomyaria of the Upper Helderberg, Hamilton, Portage and Chemung Groups . New York Geological Survey, Paleontology 5(1), 1-268.

HAMMER, O., HARPER, D.A.T. & RYAN, P.D. 2001. PAST: Palaeontological statistics software package for education and data analysis. Palaeontologia Electronica 4(1), 1-9.

HAQ, B.U. & SCHUTTER, S.R. 2008. A Chronology of Paleozoic Sea-Level Changes. Science 322(5898), 64-68.View article

HAUDE, R. 1992. Scyphocrinoiden, die Bojen-Seelilien im hohen Silur-tiefen Devon. Palaeontographica, Abteilung A 222, 141-187.

HAUDE, R. & WALLISER, O.H. 1998. Conodont-based Upper Silurian-Lower Devonian range of scyphocrinoids in SE Morocco, 94-96. In GUTIÉRREZ-MARCO, J.C. & RÁBANO, I. (eds) Proceedings of the sixth international graptolite conference of the GWG (IPA) and the SW Iberia field meeting 1998 of the International Subcommission on Silurian Stratigraphy (ICS-IUGS). Instituto Tecnológico Geominero de Espana, Madrid.

HAUSER, J. 2008. Revision von Tiaracrinus (Crinoidea, Cladida) aus dem Paläozoikum der Eifel und dem Sauerland (Rheinische Schiefergebirge), Massive Armoricain (Frankreich), Böhmen (Tschechien) und Nord-Afrika (Algerien). Crinoiden des Devon, 1-8. http://www.devon-crinoiden.de/tiaracrinus.pdf

HOLLARD, H. 1967. Le Dévonien du Maroc et du Sahara nord occidental. International Symposium on the Devonian System, Calgary 1, Canadian Society of Petroleum Geologists 1, 203-244.

HOLLARD, H. 1974. Recherches sur la stratigraphie des formations du Dévonien moyen, de l’Emsien supérieur au Frasnien, dans le Sud du Tafilalt et dans le Ma’der (Anti-Atlas oriental, Maroc). Notes du Service géologique du Maroc 36(264), 7-68.

HOLLARD, H. 1981.Tableaux de Corrélation du Silurien et Dévonien de l’Anti-Atlas. Notes du Service géologique du Maroc 42(308), 23-46.

HORNÝ, R.J. 1955. Palaeozygopleuridae nov. fam. (Gastropoda) ze středočeského devonu. Sborník Ústředního ústavu geologického, Oddíl paleontologický 21(1), 17-74.

HORNÝ, R.J. 1963. Lower Paleozoic Bellerophontina (Gastropoda) of Bohemia. Sborník geologických věd, Paleontologie 2, 57-164.

HORNÝ, R. 1992a. New Lower Devonian gastropod genera of Bohemia (Mollusca). Časopis Národního muzea, Řada přírodovědná 158(1-4), 105-107.

HORNÝ, R. 1992b. Lytospira Koken a Murchisonia (Hormotomina) Grabau et Shimer v českém spodním devonu. [Lytospira Koken and Murchisonia (Hormotomina) Grabau et Shimer in the Lower Devonian of Bohemia.] Časopis Národního muzea, Řada přírodovědná 160(1-4), 55-56.

HORNÝ, R.J. 1998. Two additional, isolated, paucispiral gastropod opercula from the Lower Devonian Koněprusy Limestone (Bohemia, Barrandian area). Journal of the National Museum (Prague), Natural History Series 167 (1-4), 91-94.

JELL, P.A. & JELL, J.S. 1999. Crinoids, a blastoid and a cyclocystoid from the Upper Devonian reef complex of the Canning Basin, Western Australia. Memoirs of The Queensland Museum 43, 201-236.

JOHNSON, J.G., KLAPPER, G. & ELRICK, M. 1996. Devonian transgressive-regressive cycles and biostratigraphy, Northern Antelope Range, Nevada: Establishment of reference horizons for global cycles. Palaios 11, 3-14.View article

JOHNSON, J.G., KLAPPER, G. & SANDBERG, C.A. 1985. Devonian eustatic fluctuations in Euramerica. Geological Society of America Bulletin 96, 567-587.View article

KAUFMANN, B. 1998. Facies analysis, stratigraphy and diagenesis of Middle Devonian reef- and mud-mounds in the Mader (eastern Anti-Atlas, Morocco). Acta Geologica Polonica 48(1), 43-106.

KLEIN, J.T. 1734. Naturalis dispositio Echinodermatum: Accesseit Lucubratiuncula de aculeis echinorum marinorum, cum Spicilegio de belemnitis. Typis Thom. Joh. Schreiberi.View article

KLUG, C. 2001. Early Emsian ammonoids from the eastern Anti-Atlas (Morocco) and their succession. Paläontologische Zeitschrift 74(4), 479-515.View article

KLUG, C. 2002. Quantitative stratigraphy and taxonomy of late Emsian and Eifelian ammonoids of the eastern Anti-Atlas (Morocco). Courier Forschungsinstitut Senckenberg 238, 1-109.

KLUG, C. & KORN, D. 2002. Occluded umbilicus in the Pinacitinae (Devonian) and its palaeoecological implications. The Palaeontological Association 45(5), 917-931.View article

KLUG, C., KORN, D., NAGLIK, C., FREY, L. & DE BAETS, K. 2013. The Lochkovian to Eifelian succession of the Amessoui Syncline (southern Tafilalt), 51-60. In BECKER, R.T., EL HASSANI, A. & TAHIRI, A. (eds) International Field Symposium “The Devonian and Lower Carboniferous of northern Gondwana”, Morocco 2013.

KLUG, C., KORN, D. & REISDORF, A. 2000. Ammonoid and conodont stratigraphy of the late Emsian to early Eifelian (Devonian) at the Jebel Ouaoufilal (near Taouz, Tafilalt, Morocco), 45-56. In TAHIRI, A. & EL HASSANI, A. (eds) Travaux de l’Institut Scientifique, Série Géologie & Géographie Physique 20. Proceedings of the Subcommission on Devonian Stratigraphy (SDS) - IGCP 421 Morocco Meeting.

KLUG, C., KRÖGER, B., KIESSLING, W., MULLINS, G. L., SERVAIS, T., FRÝDA, J., KORN, D. & TURNER, S. 2010. The Devonian nekton revolution. Lethaia 43, 465-477.View article

KLUG, C., KRÖGER, B., KORN, D., RÜCKLIN, M., SCHEMM-GREGORY, M., DE BAETS, K. & MAPES, R.H. 2008a. 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, Abteilung A 283(4-6), 83-176.

KLUG, C., MEYER, E., RICHTER, U. & KORN, D. 2008b. Soft-tissue imprints in fossil and Recent cephalopod septa and septum formation. Lethaia 41, 477-492.View article

KNIGHT, J.B. 1941. Paleozoic gastropod genotypes. Geological Society of America, Special Paper 32, 1-510.

KNIGHT, J.B. 1956. New families of Gastropoda. Washington Academy of Sciences Journal 46, 41-42.

KOKEN, E. 1896. Die Leitfossilien. 848 pp. Chr. Herm. Trachnitz, Leipzig.

KOKEN, E. 1889. Über die Entwicklung der Gastropoden vom Kambrium bis zur Trias. Neues Jahrbuch für Mineralogie, Geologie und Paläontologie, Beilage Band 6, 305-484.

KONINICK, L.G. DE 1881. Faune du calcaire carbonifere de Belgique, Pt. 3, Gastéropodes. Annales du Musée royal d’Histoire naturelle de Belgique, paleontological series 6, 1-170.

KŘÍŽ, J. 1998. Recurrent Silurian-lowest Devonian cephalopod limestone of Gondwanan Europe and Perunica, 183-198. In LANDING, E. & JOHNSON, M.E. (eds) Silurian cycles; linkages of dynamic stratigraphy with atmospheric, oceanic, and tectonic changes. New York State Museum Bulletin 491.

KŘÍŽ, J. 2000. Lochkovian bivalves of Bohemian type from the eastern Anti-Atlas (Lower Devonian, Morocco). Senckenbergiana lethaea 80(2), 485-523.View article

KRÖGER, B. 2008. Nautiloids before and during the origin of ammonoids in Siluro-Devonian section in the Tafilalt, Anti-Atlas, Morocco. Special Papers in Palaeontology 79, 1-110.

KUHN, O. 1940. Paläozoologie in Tabellen. 50 pp. Fischer Verlag, Jena.

LEVEILLE, C. 1835. Aperçu géologique de quelques localités tres riches en coquilles sur les frontieres de France et de Belgique. Mémoires de la Société géologique de France 2(1), 29-40.

LINDSTRÖM, G. 1884. On the Silurian gastropoda and Pteropoda of Gotland. Kongliga Svenska Vetenskaps Akademiens Handlingar 19(6), 1-250.

LINNAEUS, C. 1758. Systema Naturae per Regna Tria Naturae, secundum Classes, Ordines, Genera, Species, cum Characteribus, Differentiis, Synonymis, Locis. Tomus I. 823 pp. Laurentii Salvii, Holmiae (Stockholm).View article

LUBESEDER, S., REDFERN, J. & BOUTIB, L. 2009. Mixed siliciclastic-carbonate shelf sedimentation-Lower Devonian sequences of the SW Anti-Atlas, Morocco. Sedimentary Geology 215(1-4), 13-32.View article

MASSA, D., COMBAZ, A. & MANDERSCHEID, G. 1965. Observations sur les séries siluro-dévoniennes des confins algéro-marocains du Sud. Notes et Mémoires, Compagnie Française des Pétroles 8, 1-188.

M’COY, F. 1844. A synopsis of the characters of the Carboniferous Limestone fossils of Ireland. 274 pp. University Press, Dublin.View article

M’COY, F. 1851. On some new Silurian Mollusca. Annals and Magazine of History, including Zoology, Botany, and Geology, 2nd series 7, 45-63.

MENEGHINI, G. 1857. Paléontologie de l’Ile Sardaigne, 53-144. In LA MARMORA, A. (ed.) Voyage en Sardaigne. Imprimerie Royal, Turin.

MILNE-EDWARDS, H. & HAIME, J. 1851. Monographie des Polypiers fossiles des terrains palaeozoiques, précédée d’un tableau général de la classification des polypes. 502 pp. Gide et J. Baudry, Paris.

MILLER, J.S. 1821. A natural history of the Crinoidea or lily-shaped animals, with observation on the genera Astria, Euryale, Comatula, and Marsupites. 150 pp. Bryan & Co., Bristol.

MILLER, J. 1976. The sensory fields and life mode of Phacops rana (Green, 1832). Transactions of the Royal Society of Edinburgh 69, 337-367.View article

MILLER, S.A. 1892. North American geology and palaeontology. First appendix, 665-718. Western Methodist Book Concern, Cincinnati.

MOORE, R.C. & LAUDON, L.R. 1943. Evolution and classification of Paleozoic crinoids. Geological Society of America, Special Paper 46, 1-153.View article

MUNIER-CHALMAS, E. 1876. Mollusques nouveaux des terrains paléozoiques des environs de Rennes. Journal de Conchyliologie, 3e series 16, 102-109.

NEYMAN, A.A. 1967. Limits to the application of the ‘trophic group’ concept in benthic studies. Oceanology, Academy of Sciences of the USSR 7, 49-155.

OEHLERT, D. 1881. Documents pour servir a l’étude des faunes dévoniennes dans l’ouest de la France. Mémoires de la Société Géologique de France, 3e série 2, 1-38.

PEEL, J. 1984. Faunal succession and mode of life of Silurian gastropods in the Arisaig Group, Nova Scotia. Palaeontology 21, 258-306.

PERNER, J. 1903. Gastéropodes, 1-164. In BARRANDE, J. (ed.) Systeme silurien du centre de la Boheme 4(1). Prague.

PERNER, J. 1907. Gastéropodes, 1-380. In BARRANDE, J. (ed.) Systeme silurien du centre de la Boheme 4(2). Prague.

PIQUÉ, A. & MICHARD, A. 1989. Moroccan Hercynides: A synopsis. The Paleozoic sedimentary and tectonic evolution at the northern margin of west Africa. American Journal of Science 289, 286-330.View article

RACHEBOEUF, P.R. 1990. Les Brachiopodes Chonetacés dans les assemblages benthiques siluriens et dévoniens. Palaeogeography, Palaeoclimatology, Palaeoecology 81, 141-171.View article

ŘÍha, A. 1938. Příspěvek k poznání nových plžů ze středočeského paleozoika. Věstník Královské České společnosti nauk, Třída matematicko-přírodovědecká, 1-12.

Roch, E. 1934. Sur des phénomenes remarquables observés dans la région d’Erfoud (confins algéro-marocains du Sud). Association Études de la Géologie Méditerranéenne Occidentale 5, 1-10.

Rohr, D.M. & Smith, R.E. 1978. Lower Devonian Gastropoda from the Canadian Arctic Islands. Canadian Journal of Earth Sciences 15, 1228-1241.View article

Sanders, H.L. 1968. Benthic diversity: A comparative study. American Society of Naturalists 102(925), 243-282.View article

Saunders, W.B. & Ward, P.D. 1987. Ecology, distribution and population characteristics of Nautilus, 135-162. In Saunders, W.B. & Landman, N.H. (eds) The Biology and Paleobiology of a Living Fossil. Reprinted with additions (2009). Springer, Berlin.

Schönlaub, H.P. 1996. Significant geological events in the Paleozoic record of the Southern Alps (Austrian part), 163-168. In Walliser, O.H. (ed.) Global events and event stratigraphy in the Phanerozoic. Springer, Berlin.

Schultze, L. 1866. Monographie der Echinodermen des Eifler-Kalkes. Denkschrift

Königlichen Akademie der Wissenschaften., mathematisch-naturwissenschaftlichen Klasse 26, 113-230.

Scotese, C.R. 1997. Paleogeographic Atlas. 37 pp. PALEOMAP Progress Report 90-0497, Department of Geology, University of Texas at Arlington.

Scotese, C.R. & McKerrow, W.S. 1990. Revised world maps and introduction, 1-21. In McKerrow, W.S. & Scotese, C.R. (eds) Palaeozoic Palaeogeography and Biogeography. Geological Society of London Memoirs 12.

Shimizu, S. & Obata, T. 1935. New genera of Gotlandian and Ordovician nautiloids. Journal of the Shanghai Science Institute, Section 2, 1-10.

Simpson, G.B. 1897. A handbook of the genera of the North American Paleozoic Bryozoa. Fourteenth Annual Report of the State Geologist, New York 1894, 407-608.

Simroth, H. 1906. Versuch einer neuen Deutung der Bellerophontiden. Sitzungsberichte der Naturforschenden Gesellschaft zu Leipzig 1905, 3-8.

Solle, G. 1952. Neue Untergattungen und Arten der Bryozoen-Gattung Hederella und eine Hernodia im rheinischen Unterdevon. Notizblatt des Hessischen Landesamtes für Bodenforschung zu Wiesbaden 6(3), 35-55.

Solle, G. 1968. Hederelloidea (Cyclostomata) und einige ctenostome Bryozoen aus dem rheinischen Devon. Abhandlungen des Hessischen Landesamtes für Bodenforschung 54, 1-40.

Südkamp, W.H. & Burrow, C.J. 2007. The acanthodian Machaeracanthus from the Lower Devonian Hunsrück Slate of the Hunsrück region (Germany). Systematic palaeontology. Paläontologische Zeitschrift 81, 97-104.View article

Talent, J.A., Mawson, R., Andrew, A.S., Hamilton, J. & Whiteford, D.J. 1993. Middle Palaeozoic extinction events: faunal and isotope data. Palaeogeography, Palaeoclimatology, Palaeoecology 104, 139-152.View article

Talent, J.A. & Yolkin, E.A. 1987. Transgression-regression patterns for the Devonian of Australia and southern West Siberia. Courier Forschungs-Institut Senckenberg 92, 235-249.

Tassell, C.B. 1980. Further gastropods from the Early Devonian Lilydale Limestone, Victoria. Records of the Queen Victoria Museum 69, 1-27.

Tassell, C.B. 1982. Gastropods from the Early Devonian “Receptaculites” Limestone, Taemas, New South Wales. Records of the Queen Victoria Museum 77, 1-59.

Taylor, P.D. & Wilson, M.A. 2008. Morphology and affinities of hederelloid “bryozoans”, 301-309. In Hageman, S.J., Key, M.M., Jr. & Winston, J.E. (eds) Bryozoan Studies 2007: Proceedings of the 14th International Bryozoology Conference, Boone, North Carolina, July 1-8, 2007. Virginia Museum of Natural History, Special Publication 15.

Teichert, C., Kummel, B., Sweet, W.C., Stenzel, H.B., Furnish, W.M., Glenister, B.F., Erben, H.K., Moore, R.C. & Zeller, D.E.N. 1964. Endoceratoidea-Actinoceratoidea-Nautiloidea, K13-K124. In Moore, R.C. (ed.) Treatise on Invertebrate Paleontology, Part K, Mollusca 3, Cephalopoda. Geological Society of America and University of Kansas Pres, Boulder & Lawrence.

Termier, G. & Termier, H. 1950. Paléontologie Marocaine. II. Invertébrés de l’ére Primaire. Fascicule III. Mollusques. Notes et Mémoires du Service Géologique, Rabat 78, 1-246.

Toto, E.A., Kaabouben, F., Zouhri, L., Belarbi, M., Benammi, M., Hafid, M. & 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.View article

Ubaghs, G. 1978. Part T Echinodermata 2, T403-T812. In Moore, R.C. (ed.) Treatise on Invertebrate Paleontology, Part T. Geological Society of America and University of Kansas Press 2, Lawrence.

Ubaghs, G. & Bouček, B. 1962. Sur la présence du genre Tiaracrinus Schultze (Crinoidea) dans le Dévonien inférieur de Moravie (Tiaracrinus moravicus n. sp.). Sborník Ústředního ústavu geologického, Oddíl paleontologický 27, 41-50.

Van Geldern, R., Joachimski, M.M., Day, J., Jansen, U., Alvarez, F., Yolkin, E.A. & Ma, X.-P. 2006. Carbon, oxygen and strontium isotope records of Devonian brachiopod shell calcite. Palaeogeography, Palaeoclimatology, Palaeoecology 240, 47-67.View article

Verneuil, N. de 1850. Notes sur les fossiles Dévoniens du district de Sabéro (León). Bulletin de la Société Géologique de France 7(2), 155-186.

Wachsmuth, C. & Springer, F. 1885. Revision of the Paleocrinoidea, Part III: Discussion and classification of the brachiate crinoids, and conclusion of the generic description. Proceedings of the Academy ofNatural Sciences of Philadelphia, 223-364.

Walker, K.R. & Bambach, R.K. 1971. The significance of fossil assemblages from fine grained sediments: Time-averaged communities. Geological Society of America Abstracts with Programs 3, 783-784.

Walliser, O.H. 1984. Global events, event-stratigraphy and “chronostratigraphy” within the Phanerozoic, 208. In Bogdanov, N.A. (ed.) 27th International Geological Congress. Abstracts, section 1.

Walliser, O.H. 1985. Natural boundaries and Commission boundaries in the Devonian. Courier Forschungsinstitut Senckenberg 75, 401-408.

Webster, G.D. & Donovan, S.K. 2012. Before the extinction - Permian platyceratid gastropods attached to platycrinitid crinoids and an abnormal four-rayed Platycrinites wachsmuthi (Wanner) from West Timor. Palaeoworld 21(3-4), 153-159.View article

Wendt, J. 1985. Disintegration of the continental margin of northwestern Gondwana: Late Devonian of the eastern Anti-Atlas (Morocco). Geology 13, 815-818.View article

Wendt, J. 1988. Facies pattern and paleogeography of the Middle and Late Devonian in the eastern Anti-Atlas (Morocco), 467-479. In McMillan, J., Embry, A.F. & Glass, D.J. (eds) Devonian of the World, volume I. Canadian Society of Petroleum Geologists, Calgary.

Wendt, J., Aigner, T. & Neugebauer, J. 1984. Cephalopod limestone deposition on a shallow pelagic ridge: the Tafilalt Platform (upper Devonian, eastern Anti-Atlas, Morocco). Sedimentology 31(5), 601-625.View article

Wenz, W. 1938. Gastropoda, Teil I. In Schindewolf, O.H. (ed.) Handbuch der Paläozoologie 6. 1639 pp. Borntraeger, Berlin.

Westermann, G.E.G. 1999. Life of habits of nautiloids. In Savazzi, E. (ed.) Functional Morphology of the Invertebrate Skeleton. John Wiley & Sons, New York.

Westermann, G.E.G. & Tsujita, C.J. 1999. Life habits of ammonoids, 299-325. In Savazzi, E. (ed.) Functional Morphology of the Invertebrate Skeleton. John Wiley & Sons, New York.

Whittaker, R.H. 1960. Vegetation of the Siskiyou Mountains, Oregon and California. Ecological Monographs 30, 279-338.View article

Whittaker, R.H. 1972. Evolution and measurements of species diversity. Taxon 21(2-3), 213-251.View article

Whittington, H.B. 1988. Hypostomes of post-Cambrian trilobites. Memoirs of the New Mexico Bureau of Mines and Mineral Resources 44, 321-339.

Williams, A., Bruton, C.H.C. & Carlson, S.J. 2000. Brachiopoda (revised), 1-920. In Kaesler, R.S. (ed.) Treatise on Invertebrate Paleontology, Part H (2-3). Geological Society of America & University of Kansas Press, Lawrence.

Yochelson, E.L. & Linsley, R.M. 1972. Opercula of two gasteropods from the Lilydale Limestone (Early Devonian) of Victoria, Australia. National Museum of Victoria, Memoir 33, 1-14.View article

Zhuravleva, F.A. 1959. On the family Michelinoceratidae. Materialy k Osnovam paleontologii 3, 47-48.

Ziegler, B. 1972. Allgemeine Paläontologie: Einführung in die Paläontologie, Teil 1. 181 pp. Schweizerbart’sche Verlagsbuchhandlung, Stuttgart.