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Reply to Comment on: Álvaro J.J., Esteve, J. & Zamora, S. 2019. Morphological assessment of the earliest paradoxidid trilobites (Cambrian Series 3) from Morocco and Spain [Geological Magazine] by Geyer G, Nowicki J, Żylińska A & Landing E

Published online by Cambridge University Press:  13 April 2020

J Javier Álvaro Open the ORCID record for J Javier Álvaro [Opens in a new window]  and
Jorge Esteve Open the ORCID record for Jorge Esteve [Opens in a new window]
J Javier Álvaro*
Affiliation:
Instituto de Geociencias (CSIC-UCM), Dr Severo Ochoa 7, 28040Madrid, Spain
Jorge Esteve
Affiliation:
Facultad de Ciencias, Dpto. de Geociencias, Universidad de los Andes, Cr 1 18A-12, Bogotá, Colombia
*
Author for correspondence: J Javier Álvaro, Email: jj.alvaro@csic.es
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Abstract

Two research teams have different opinions regarding the number of paradoxidine species close to the base of the Miaolingian in Morocco and Spain. Geyer & Vincent (2015) recognized five species based on a mosaic of overlapping characters that include some relative length and width ratios, qualitative descriptions of cranidial and pygidial outlines, and relief observations. In contrast, Álvaro et al. (2018) conducted a morphometric analysis of all these specimens from the Cambropallas telesto acme level of the Assemame quarry and concluded that the Moroccan material had been taxonomically over-split and was represented by only two species, Acadoparadoxides mureroensis and A. nobilis. Subsequently, Geyer et al. (2019) commented that we had not analysed some diagnostic characters, and applied the stratigraphic setting of their morphospecies as a diagnostic character for taxonomic identification. After sampling the Cambropallas telesto acme level in the Taroucht quarry, where the other team purchased its collection, and re-analysing the diagnostic characters claimed by Geyer et al. (2019), our conclusions are maintained: (i) morphometric values of several paratype specimens fall outside the purported ranges of diagnostic characters; (ii) all 2D morphometric analyses incorporating the diagnostic characters of the morphospecies are unable to detect interspecific differences, except for A. nobilis; and (iii) the slight concavity displayed by some pygidia, strongly controlled by preservation, is in need of 3D biometrical analysis. Hence, our proposed synonymy should be maintained until 3D statistical analyses are available on material preserved in carbonate or concretions.

Keywords

trilobiteCambrianmorphometricsWest Gondwana
Type
Discussion - Reply
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Geological Magazine , Volume 157 , Issue 12 , December 2020 , pp. 1971 - 1982
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© Cambridge University Press 2020

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References

Algouti, A, Chabane, A, Dal Piaz, GV, El Boukhari, A, Ellero, A, Feroni, AC, Ghiselli, F, Kamal, TE, Malusà, MG, Massironi, M, Musumeci, G, Ottria, G, Ouanaimi, H, Pertusati, PC and Schiavo, A (2007) Carte géologique du Maric au 1/50,000. Feuille Taroucht. Notes et Mémoires du Service Géologique du Maroc 520, 1–78.Google Scholar
Álvaro, JJ (2014) Rift, pull-apart rift, and continental drift crossword puzzles across the lower–middle Cambrian transition of Iberia and Morocco. GFF 136, 2–5.CrossRefGoogle Scholar
Álvaro, JJ and Clausen, S (2005) Major geodynamic and sedimentary constraints on the chronostratigraphic correlation of the Lower–Middle Cambrian transition in the western Mediterranean region. Geosciences Journal 9, 145–60.CrossRefGoogle Scholar
Álvaro, JJ and Clausen, S (2006) Microbial crusts as indicators of stratigraphic diastems in the Cambrian Micmacca Breccia, Moroccan Atlas. Sedimentary Geology 185, 255–65.CrossRefGoogle Scholar
Álvaro, JJ and Clausen, S (2008) Paleoenvironmental significance of Cambrian hiatal shell accumulations in an aborted intra-cratonic rift, Atlas Mountains, Morocco. In Dynamics of Epeiric Seas (eds Pratt, BR and Holmden, C), pp. 39–54. Geological Association of Canada, Special Paper no. 48. St John’s, NF.Google Scholar
Álvaro, JJ and Vizcaïno, D (2018) The Furongian break-up (rift-drift) unconformity in the Anti-Atlas, Morocco. Journal of Iberian Geology 44, 567–87.CrossRefGoogle Scholar
Álvaro, JJ, Benziane, F, Thomas, AR, Walsh, GJ and Yazidi, A (2014) Neoproterozoic–Cambrian stratigraphic framework of the Anti-Atlas and Ouzellagh promontory (High Atlas), Morocco. Journal of African Earth Sciences 98, 19–33.CrossRefGoogle Scholar
Álvaro, JJ, Ezzouhairi, H, Clausen, S, Ribeiro, ML and Solá, R (2015) Syn-rift unconformities punctuating the lower–middle Cambrian transition of the Atlas Rift, Morocco. International Journal of Earth Sciences 104, 752–73.CrossRefGoogle Scholar
Álvaro, JJ, Esteve, J and Zamora, S (2018) Morphological assessment of the earliest paradoxidid trilobites (Cambrian Series 3) from Morocco and Spain. Geological Magazine 155, 1566–95.CrossRefGoogle Scholar
Angielczyk, KD and Sheets, HD (2007) Investigation of simulated tectonic deformation in fossils using geometric morphometrics. Paleobiology 33, 125–48.CrossRefGoogle Scholar
Benton, MJ (2015) Palaeodiversity and formation counts: redundancy or bias? Palaeontology 58, 1003–29.CrossRefGoogle Scholar
Boudda, A, Choubert, G and Faure-Muret, A (1979) Essai de stratigraphie de la couverture sédimentaire de l’Anti-Atlas: Adoudounien, Cambrien inférieur. Notes et Mémoires du Service Géologique du Maroc 271, 1–96.Google Scholar
Buggisch, W, Marzela, C and Hügel, P (1978) Die fazielle und paläogeographische Entiwocklung der infrakambrischen bis ordovizischen Sedimente im Mittleren Antiatlas um Agdz (S-Marokko). Geologische Rundschau 68, 195–224.CrossRefGoogle Scholar
Catuneanu, O, Galloway, WE, Kendall, CGStC, Miall, AD, Posamentier, HW, Strasser, A and Tucker, ME (2011) Sequence stratigraphy: methodology and nomenclature. Newsletters on Stratigraphy 44, 173–245.CrossRefGoogle Scholar
Cisne, JL, Molenock, J and Rabe, BD (1980) Evolution in a cline: the trilobite Triarthus along an Ordovician depth gradient. Lethaia 13, 47–59.CrossRefGoogle Scholar
Cisne, JL, Chandlee, GO, Rabe, BD and Cohen, JA (1982) Clinal variation, episodic evolution, and possible parapatric speciation: the trilobite Flexicalymene senaria along an Ordovician depth gradient: Lethaia 15, 325–41.CrossRefGoogle Scholar
Clausen, S, Álvaro, JJ and Zamora, S (2014) Replacement of benthic communities in two Neoproterozoic–Cambrian subtropical-to-temperate rift basins, High Atlas and Anti-Atlas, Morocco. Journal of African Earth Sciences 98, 72–93.CrossRefGoogle Scholar
Destombes, J, Hollard, H and Willefert, S (1985) Lower Palaeozoic rocks of Morocco. In Lower Palaeozoic Rocks of the World, vol. 4: Lower Palaeozoic of North-Western and West Central Africa (ed. Holland, CH), pp. 57–184. Chichester: John Wiley and Sons.Google Scholar
Elicki, O and Geyer, G (2013) The Cambrian trilobites of Jordan – taxonomy, systematic and stratigraphic significance. Acta Geologica Polonica 63, 1–56.CrossRefGoogle Scholar
Esteve, J, Zhao, YL and Peng, J (2017) Morphological assessment of the Cambrian trilobites Oryctocephalus indicus (Reed, 1910) from China and Oryctocephalus ‘reticulatus’ (Lermontova, 1940) from Siberia. Lethaia 50, 175–93.CrossRefGoogle Scholar
Esteve, J, Zhao, YL, Maté-González, MA, Gómez Heras, M and Peng, J (2018) A new high-resolution 3-D quantitative method for analysing small morphological features: an example using a Cambrian trilobite. Scientific Reports 8, 2868. doi: 10.1038/s41598-018-21088-4.CrossRefGoogle ScholarPubMed
Forey, PL, Fortey, RA, Kenrick, P and Smith, A (2004) Taxonomy and fossils: a critical appraisal. Philosophical Transactions of the Royal Society, London B 359, 639–53.CrossRefGoogle ScholarPubMed
Fortey, RA (2009) A new giant Asaphid trilobite from the Lower Ordovician of Morocco. Memoirs of the Association of Australasian Paleontologists 37, 9–16.Google Scholar
Gasquet, D, Levresse, G, Cheilletz, A, Azizi-Samir, MR and Mouttaqi, A (2005) Contribution to a geodynamic reconstruction of the Anti-Atlas (Morocco) during Pan-African times with the emphasis on inversion tectonics and metallogenic activity in the Precambrian-Cambrian transition. Precambrian Research 140, 157–82.CrossRefGoogle Scholar
Geyer, G (1989) Late Precambrian to early Middle Cambrian lithostratigraphy of southern Morocco. Beringeria 1, 115–43.Google Scholar
Geyer, G (1993) The giant Cambrian trilobites of Morocco. Beringeria 8, 71–107.Google Scholar
Geyer, G (1998) Intercontinental, trilobite-based correlation of the Moroccan early Middle Cambrian. Canadian Journal of Earth Sciences 35, 374–401.CrossRefGoogle Scholar
Geyer, G and Landing, E (eds) (1995) Morocco’95. The Lower–Middle Cambrian Standard of Gondwana. Beringeria, Special Issue 2, 171 pp.Google Scholar
Geyer, G and Landing, E (2004) A unified Lower–Middle Cambrian chronostratigraphy for West Gondwana. Acta Geologica Polonica 54, 179–218.Google Scholar
Geyer, G and Vincent, T (2015) The Paradoxides puzzle resolved: the appearance of the oldest paradoxidine and its bearing on the Cambrian Series 3 lower boundary. Paläontologische Zeitschrift 89, 335–98.CrossRefGoogle Scholar
Geyer, G, Nowicki, J, Żylińska, A and Landing, E (2019) Comment on: Álvaro, J.J., Esteve, J. & Zamora, S. 2019. Morphological assessment of the earliest paradoxidid trilobites (Cambrian Series 3) from Morocco and Spain [Geological Magazine]. Geological Magazine 156, 1691–707.CrossRefGoogle Scholar
Gozalo, R, Dies Álvarez, ME, Gámez Vintaned, JA, Zhuravlev, AY, Bauluz, B, Subías, I, Chirivella Martorell, JB, Mayoral, E, Gursky, HJ, Andrés, JA and Liñán, E (2013) Proposal of a reference section and point for the Cambrian Series 2–3 boundary in the Mediterranean subprovince in Murero (NE Spain) and its intercontinental correlation. Geological Journal 48, 142–55.CrossRefGoogle Scholar
Gutiérrez-Marco, JC and García-Bellido, D (in press) The international fossil trade from the Paleozoic of the Anti-Atlas, Morocco. In The Great Ordovician Biodiversification Event: Insights from the Tafilalt Biota, Morocco (eds AW Hunter, JJ Álvaro, B Lefebvre, P Van Roy and S Zamora). Geological Society of London, Special Publication no. 485. doi: 10.1144/SP485.1.CrossRefGoogle Scholar
Hopkins, MJ and Webster, M (2009) Ontogeny and geographic variation of a new species of the corynexochine trilobite Zacanthopsis (Dyeran, Cambrian). Journal of Paleontology 83, 524–47.CrossRefGoogle Scholar
Hughes, NC (1994) Ontogeny, intraspecific variation, and systematics of the Late Cambrian trilobite Dikelocephalus . Smithsonian Contributions to Paleobiology 79, 1–89.CrossRefGoogle Scholar
Hughes, NC and Chapman, RE (1995) Growth and variation in the Silurian proetide trilobite Aulacopleura konincki and its implications for trilobite palaeobiology. Lethaia 28, 333–53.CrossRefGoogle Scholar
Hughes, NC and Jell, PA (1992) A statistical/computer-graphic technique for assessing variation in tectonically deformed fossils and its application to Cambrian trilobites from Kashmir. Lethaia 25, 317–30.CrossRefGoogle Scholar
Hunda, BR and Hughes, NC (2007) Evaluating paedomorphic heterochrony in trilobites: the case of the diminutive trilobite Flexicalymene retrorsa minuens from the Cincinnatian Series (Upper Ordovician), Cincinnati region. Evolution & Development 9, 483–98.CrossRefGoogle ScholarPubMed
Jackson, ISC and Budd, GE (2017) Intraspecific morphological variation of Agnostus pisiformis, a Cambrian Series 3 trilobite-like arthropod. Lethaia 50, 467–85.CrossRefGoogle Scholar
Kim, K, Sheets, HD, Haney, RA and Mitchell, CE (2002) Morphometric analysis of ontogeny and allometry of the Middle Ordovician trilobite Triarthrus beckii . Paleobiology 28, 364–77.2.0.CO;2>CrossRefGoogle Scholar
Kim, K, Sheets, HD and Mitchell, CE (2009) Geographic and stratigraphic change in the morphology of Triarthrus beckii (Green) (Trilobita): a test of the Plus ça change model of evolution. Lethaia 42, 108–25.CrossRefGoogle Scholar
Klingenberg, CP (2004) Integration, modules, and development: molecules to morphology to evolution. In Phenotypic Integration: Studying the Ecology and Evolution of Complex Phenotypes (eds Pigliucci, M and Preston, KA), pp. 213–30. Oxford: Oxford University Press.Google Scholar
Klingenberg, CP, Barluenga, M and Meyer, A (2002) Shape analysis of symmetric structures: quantifying variation among individuals and asymmetry. Evolution 56, 1909–20.CrossRefGoogle ScholarPubMed
Labandeira, C and Hughes, NC (1995) Biometry of the Late Cambrian trilobite genus Dikelocephalus and its implications for trilobite systematics. Journal of Paleontology 68, 492–517.CrossRefGoogle Scholar
Landing, E, Geyer, G and Heldmaier, W (2006) Distinguishing eustatic and epeirogenic controls on Lower-Middle Cambrian boundary successions in West Gondwana (Morocco and Iberia). Sedimentology 53, 899–911.CrossRefGoogle Scholar
Lloyd, GT (2012) A refined modelling approach to assess the influence of sampling on palaeobiodiversity curves: new support for declining Cretaceous dinosaur richness. Biology Letters 8, 123–6.CrossRefGoogle ScholarPubMed
Michard, A, Hoepffner, C, Soulaimani, A and Baidder, L (2008) The Variscan Belt. In Continental Evolution: The Geology of Morocco (eds Michard, A, Saddiqi, O, Chalouan, A and Frizon de Lamotte, D), pp. 65–131. Berlin: Springer.CrossRefGoogle Scholar
Nowicki, J and Żylińska, A (2019) The first occurrence of the earliest species of Acadoparadoxides outside West Gondwana (Cambrian; Holy Cross Mountains, Poland). Geological Magazine 156, 1027–51.CrossRefGoogle Scholar
Posamentier, HW and Allen, GP (1999) Siliciclastic sequence stratigraphy: concepts and applications. SEPM Concepts in Sedimentology and Paleontology 7, 1–210.Google Scholar
Pouclet, A, Aarab, A, Fekkak, A and Benharref, M (2007) Geodynamic evolution of the northwestern Paleo-Gondwana margin in the Moroccan Atlas at the Precambrian–Cambrian boundary. In The Evolution of the Rheic Ocean: From Avalonian–Cadomian Active Margin to Allegheniane Variscan Collision (eds U Linneman, RD Nance, P Kraft and G Zulauf), pp. 27–60. Geological Society of America Special Paper no. 423.Google Scholar
Pouclet, A, Ouazzani, H and Fekkak, A (2008) The Cambrian volcanosedimentary formations of the westernmost High Atlas (Morocco): their place in the geodynamic evolution of the West African Paleo-Gondwana northern margin. In The Boundaries of the West African Craton (eds N Ennih and JP Liégeois, pp. 303–27. Geological Society of London Special Publication no. 297.Google Scholar
Pouclet, A, El Hadi, H, Álvaro, JJ, Bardintzeff, JM, Benharref, M and Fekkak, A (2018) Review of the Cambrian volcanic activity in Morocco: geochemical fingerprints and geotectonic implications for the rifting of West Gondwana. International Journal of Earth Sciences 107, 2101–23.CrossRefGoogle Scholar
Rushton, AWA and Hughes, NC (1996) Biometry, systematics and biogeography of the late Cambrian trilobite Maladioidella . Transactions of the Royal Society of Edinburgh, Earth Sciences 86, 247–56.CrossRefGoogle Scholar
Scott, H (1985) Homotaxy and biostratigraphical theory. Palaeontology 28, 777–82.Google Scholar
Sdzuy, K (1958) Neue trilobiten aux dem Mittelkambrium von Spanien. Senckenbergiana lethaea 39, 235–53.Google Scholar
Sheets, HD, Kim, K and Mitchell, CE (2004) A combined landmark and outline-based approach to ontogenetic shape change in the Ordovician trilobite Triarthrus becki . In Morphometrics: Applications in Biology and Paleontology (ed. Elewa, AMT), pp. 67–82. New York: Springer.CrossRefGoogle Scholar
Smith, AB (2000) Stratigraphy in phylogeny reconstructions. Journal of Paleontology 74, 763–6.CrossRefGoogle Scholar
Smith, AB (2001) Large-scale heterogeneity of the fossil record: implications for Phanerozoic biodiversity studies. Philosophical Transactions of the Royal Society of London B 356, 351–67.CrossRefGoogle ScholarPubMed
Srivastava, DC and Shah, J (2006) Digital method for strain estimation and retrodeformation of bilaterally symmetric fossils. Geology 34, 593–6.CrossRefGoogle Scholar
Wagner, PJ (1995) Stratigraphic tests of cladistics hypotheses. Paleobiology 21, 153–78.CrossRefGoogle Scholar
Webber, AJ and Hunda, BR (2007) Quantitatively comparing morphological trends to environment in the fossil record (Cincinnatian Series; Upper Ordovician). Evolution 61, 1455–65.CrossRefGoogle Scholar
Webster, M (2007) Ontogeny and evolution of the Early Cambrian trilobite genus Nephrolenellus (Olenelloidea). Journal of Paleontology 81, 1168–93.CrossRefGoogle Scholar
Webster, M (2009) Ontogeny, systematics, and evolution of the effaced early Cambrian trilobites Peachella Walcott, 1910 and Eopeachella new genus (Olenelloidea). Journal of Paleontology 83, 197–218.CrossRefGoogle Scholar
Westrop, SR and Landing, E (2000) Lower Cambrian (Branchian) trilobites and biostratigraphy of the Hanford Brook Formation, southern New Brunswick. Journal of Paleontology 74, 858–78.CrossRefGoogle Scholar
Wheeler, QD and Platnick, NI (2000) The phylogenetic species concept (sensu Wheeler and Platnick). In Species Concepts and Phylogenetic Theory: A Debate (eds Wheeler, QD and Meier, R), pp. 55–69. New York: Columbia University Press.Google Scholar
Whittington, HB, Chatterton, BDE, Speyer, SE, Fortey, RA, Owens, RM, Chang, WT, Dean, T, Jell, PA, Laurie, JR, Palmer, AR, Repina, LN, Rushton, AWA, Shergold, JH, Clarkson, ENK Wilmot, NV and Kelly, SRA (1997) Trilobita. Introduction, Order Agnostida, Order Redlichiida. Treatise on Invertebrate Paleontology, Part O, Revised, vol. 1. Lawrence, KS: University of Kansas; New York: Geological Society of America, xxiv+530 pp.Google Scholar