Journal of Paleontology, 91(4), 2017, p. 715–734 Copyright © 2017, The Paleontological Society 0022-3360/17/0088-0906 doi: 10.1017/jpa.2016.135
Filling the Gondwanan gap: paleobiogeographic implications of new crinoids from the Castillejo and Fombuena formations (Middle and Upper Ordovician, Iberian Chains, Spain)
Selina R. Cole,1 William I. Ausich,1 Jorge Colmenar,2 and Samuel Zamora3,4
1School of Earth Sciences, The Ohio State University, Columbus, OH 43210, USA ⟨
cole.678@osu.edu⟩; ⟨
ausich.1@osu.edu⟩ 2Natural History Museum of Denmark, University of Copenhagen, Øster Voldgade 5–7, 1350, Copenhagen, Denmark
⟨
jorgecolmenarlallena@gmail.com⟩ 3Instituto Geológico y Minero de España, C/Manuel Lasala, 44, 9ºB, 50006, Zaragoza, Spain ⟨
s.zamora@
igme.es⟩ 4Department of Paleobiology, National Museum of Natural History, Smithsonian Institution, Washington, DC 20013–7012, USA
Abstract.—A diverse crinoid fauna is described from the Upper Ordovician (Katian) Fombuena Formation from the eastern Iberian Chains of Spain. New crinoids include the diplobathrid camerates Fombuenacrinus nodulus n. gen. n. sp., Goyacrinus gutierrezi n. gen. n. sp., Dalicrinus hammanni n. gen. n. sp., and Ambonacrinus decorus n. gen. n. sp.; the monobathrid camerate Eopatelliocrinus hispaniensis n. sp.; and the cladid Picassocrinus villasi n. gen. n. sp. A new occurrence of Heviacrinus melendezi Gil Cid, Domínguez Alonso, and Silván Pobes, 1996 is also documented from the Castillejo Formation (Darriwilian, Middle Ordovician) from the eastern Iberian Chains of Spain. The Fombuena Formation comprises a Gondwanan crinoid assemblage from a high paleolatitude and has the highest crinoid diversity of any currently known Katian Gondwanan fauna. This assemblage is compared to other Katian age faunas around the globe, and its paleobiogeographic implications are discussed.
Introduction
The Ordovician was a pivotal time in the evolutionary history of the Crinoidea. The oldest known crinoids are from the earliest Ordovician (early Tremadocian), by which time most major clades of Paleozoic crinoids were already present. By the Floian, crinoids had appeared on most paleocontinents. Current under- standing of Ordovician pelmatozoan faunas indicates they were highly endemic during the Early and Middle Ordovician but became more cosmopolitan during the Late Ordovician (Paul, 1976; Lefebvre and Fatka, 2003; Lefebvre et al., 2013). However, interpretations of the paleobiogeographic patterns of crinoids and other pelmatozoans during the Ordovician are inhibited by uneven global sampling. In particular, most Ordovician paleocontinents are undersampled relative to Laurentia, which has likely introduced a Laurentian bias into hypotheses of Ordovician macroevolutionary and paleobiogeo- graphic patterns (Ausich and Kammer, 2011). For this reason, the recovery and description of faunas from non-Laurentian paleocontinents is critical to an improved understanding of the dynamics of both the initial Early Ordovician radiation of crinoids and the subsequent Middle Ordovician radiation (Sprinkle and Guensburg, 2004; Peters and Ausich, 2008) that occurred during the Great Ordovician Biodiversification Event (GOBE; Webby et al., 2004). Herein, we describe five new crinoid genera and species,
one new species of Eopatelliocrinus Brower, 1973, and two taxa left in open nomenclature from the Upper Ordovician (Katian)
Fombuena Formation of northeastern Spain. We also document a new occurrence of Heviacrinus melendezi Gil Cid, Domínguez Alonso, and Silván Pobes, 1996 from the Middle Ordovician (Darriwilian) Castillejo Formation of northeastern Spain. Notably, the Fombuena fauna has the highest taxonomic richness of any Katian crinoid assemblage known from Gondwana and represents a diverse, camerate-dominated fauna from a high-latitude siliciclastic environment. Description of this new fauna serves to improve sampling of underrepresented Middle Ordovician paleocontinents, and comparison to other Katian-age crinoid assemblages enhances interpretations of paleobiogeographic patterns.
Materials and methods
Collection localities.—The studied crinoid material was collected from three localities in the eastern Iberian Chains of northeastern Spain (Fig. 1). Material collected from the Fombuena Formation came from two localities at the same stratigraphic horizon (Fig. 2). This horizon is located in the Huerva Member of the Fombuena Formation and is middle to early late Berounian (lower to middle Katian, Ka1-2 stage slices) in age (Villas, 1995). Locality 1 is located about 1 km west of the village of Fombuena (41º8'53''N, 1º12'16''W) and corresponds to the F4 locality (Peña del Tormo section) of Villas (1985). Locality 2 is located about 2 km south of the village of Fombuena (41º7'45''N, 1º12'4W) and corresponds to the F8 locality (Santa Catalina section), RC of Villas (1992). Material
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