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Triassic) Conodont Genus Metapolygnathus Hayashi

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Triassic) Conodont Genus Metapolygnathus Hayashi Journal of Earth Science, Vol. 26, No. 2, p. 219–223, April 2015 ISSN 1674-487X Printed in China DOI: 10.1007/s12583-015-0534-y The Case of the Carnian (Triassic) Conodont Genus Metapolygnathus Hayashi Ali Murat Kiliç*1, Pablo Plasencia2, Keisuke Ishida3, Francis Hirsch4 1. Department of Geological Engineering, Balıkesir University 10145, Balıkesir, Turkey 2. Department of Geology and ICBIBE, University of Valencia, Dr. Moliner 50. 46100, Burjassot, Spain 3. Laboratory of Geology, Faculty of Sciences, Tokushima University, Tokushima 770-8502, Japan 4. Laboratory of Geology, Faculty of Sciences, Naruto University, Naruto 772-8502, Japan ABSTRACT: Successive evolutionary trends control the genera Paragondolella Mosher and Meta- polygnathus Hayashi over the about 20 Ma long Aegean–Tuvalian timespan. In consideration of their evolutionary criteria, these genera have been retained together with Carnepigondolella Kozur and the two new genera proposed herein, Mazzaella (type species M. carnica) and Hayashiella (type species H. nodosa). Mazzaella n. gen. represents a Late Julian side branch of Metapolygnathus, harbouring a plat- form ornamentation similar to the several million years younger Tuvalian lineage of Carnepigondolella, issued from Hayashiella n. gen. that is intermediary between Metapolygnathus and Carnepigondolella, based on appearance and platform ornamentation. KEY WORDS: Carnian, conodont, phylogeny, evolutionary trend. 0 INTRODUCTION polygnathiformis (Budurov and Stefanov), M. acuminatus Or- The beginning of the Carnian (Upper Triassic) is marked chard, M. intermedius Orchard, M. zonneveldi Orchard, and M. by the extinction of many conodont forms that characterized lobatus Orchard. For the Tuvalian, Orchard (2007b) listed M. the Middle Triassic; one of the survivors, the genus Paragon- carpathicus (Mock), M. nodosus Hayashi, M. noah Hayashi, M. dolella, diversified during the Carnian in one of the last sig- aff. communisti, M. echinatus (Hayashi), M. parvus Kozur, M. nificant events of radiation that conodonts experimented. The stephanae Orchard, M. polygnathiformis (Budurov and Ste- nature and evolutionary relationships of the many lineages fanov), M. communisti Hayashi, M. reversus (Mosher), M. descendant from Paragondolella have been a subject that in- pseudoechinatus Kozur, M. samueli Orchard, M. pseudodiebeli cited vivid debate during the last decades. (Kozur), M. zoae Orchard, M. lindae Orchard, and M. primitius It was Hayashi (1968) who established the genus Meta- (Mosher). Orchard (2007b) redefined the variations within the polygnathus, encompassing the species M. communisti (type), genus in terms of lineages, based on following morphological M. linguiformis and M. noah. Hayashi (1987) put the species criteria: (1) platform margin profile; (2) pit position; (3) blade that he attributed to the genus Gladigondolella, on the base of length; (4) posterior carina development; (5) node distribution, their centrally located basal pit, to Metapolygnathus echinatus and (6) platform shape. However, Orchard (2013) restricted the (Hayashi), M. nodosus Hayashi, M. abneptis permicus (Hayashi) genus Metapolygnathus to only two groups: communisti and and M. spatulatus (Hayashi). parvus, introducing new genera to accommodate each of his Notwithstanding that Mosher (1973) admitted the priority other six lineages, judging Paragondolella and of Metapolygnathus Hayashi 1968 over Paragondolella Metapolygnathus as inappropriate. Mosher 1968, both genera have been used indistinctly for dec- Significant was the numerical cladistic analysis that led ades. Later, Budurov (1977), Budurov and Sudar (1990) and Mazza et al. (2012a) to recognise three different lineages: Buryi (1996) limited the definition of Metapolygnathus to bifid Paragondolella-Norigondolella, Paragondolella-Metapolygnathus morphs derived from evolved Paragondolella, using the taxon and Paragondolella-Carnepigondolella-Epigondolella, in which Ancyrogondolella Budurov (1972: type A. triangularis) for the genus Metapolygnathus consists of M. praecommunisti Mazza, extreme bifid forms. Rigo and Nicora, M. mersinensis Kozur and Moix, M. cf. primitius Orchard (2007a, 1991a, b, 1983) included into Meta- (Mosher), M. linguiformis Hayashi, M. communisti Hayashi, M. polygnathus the Julian species M. tadpole (Hayashi), M. echinatus (Hayashi) and M. parvus Kozur. In considering the resulting phylogeny from Aegean *Corresponding author: [email protected] through Tuvalian forms, based on the development of a free © China University of Geosciences and Springer-Verlag Berlin blade and bifurcation of the basal cavity, it appears justified to Heidelberg 2015 maintain Paragondolella as the ancestor of Metapolygnathus. While the former genus conforms with smooth (naked) “all Manuscript received October 13, 2014. platform” units without significant free blade, the latter is Manuscript accepted November 24, 2014. characterized by oscillating between “fair platform” units with Kiliç, A. M., Plasencia, P., Ishida, K., et al., 2015. The Case of the Carnian (Triassic) Conodont Genus Metapolygnathus Hayashi. Journal of Earth Science, 26(2): 219–223. doi:10.1007/s12583-015-0534-y 220 Ali Murat Kiliç, Pablo Plasencia, Keisuke Ishida and Francis Hirsch short free blades and “residual” platforms with an “almost all” The genus Carnepigondolella Kozur (Type species: free blade. Moreover, forms classified as Metapolygnathus that Metapolygnathus zoae Orchard), according to the original di- developed platform ornamentation in the form of increasing agnosis, adopted by Mazza et al. (2010), a long to moderately numbers of marginal nodes and denticles are classified into the long platform, bears nodes or short node-like denticles on the genus Carnepigondolella. anterior part of the platform and often on its entire length. The In the present paper we analyse some of the difficulties free blade, when present, is short to moderately long. The basal generated by the wide morphological discrepancies between the cavity is sub-terminal with respect to the keel end and to the ornamented Carnian taxa, as already sensed by our predeces- platform, or it may be slightly forward shifted, but it always sors. lies distinctly behind the middle of the platform. The keel ter- mination is broadly rounded, blunt or little bifurcated, but never 1 CASE pointed. We summarise the respective original diagnoses. Orchard (2007b) observed that platform shapes and orna- The genus Paragondolella Mosher (Type species: P. ex- ment vary in all Carnepigondolella Kozur species, but that the celsa Mosher) is defined in Mazza et al. (2010) by a character- progressive platform morphogenesis, blade ratio and anterior istic lower side with a subterminal to backward shifted pit, with migration of the pit are distinctive. respect both to keel and platform (Kozur, 1989; Mosher, 1968). Summing up, the array of species referred to the genus has The platform is rounded or blunt at the posterior end, while it varied over the years and according to the authors. For Orchard may extend completely to the anterior end or leave a short free (2007b), Carnepigondolella Kozur includes C. zoae (=Orchard, blade. Platform margins never bear any nodes. The keel is nar- 1991a, sp. F), C. samueli (=Orchard, 1991a, sp. G), C. “pseu- row or broad and its termination may be pointed, squared or doechinata” Kozur as well as Epigondolella orchardi Kozur, rounded, but never bifurcated. This long living taxon evolved Metapolygnathus nodosus Hayashi, Gondolella carpathica during the Early Anisian, persisted until the Carnian, when its Mock, Metapolygnathus lindae (=Orchard, 1991a, sp. E), M. advanced form-characteristics eventually evolved into pseudodiebeli (Kozur) and Epigondolella carnica Krystyn. At Metapolygnathus. Pizzo Mondello, Mazza et al. (2012b) listed Carnepigondolella The genus Metapolygnathus Hayashi (Type species: M. baloghi (Kovacs), C. nodosa (Hayashi), C. orchardi (Kozur), C. communisti) is characterized for Mazza et al. (2010) by a lack carpathica (Mock), C. tuvalica Mazza and Rigo, C. pseu- of denticles on the platform margins and the forward-shifted dodiebeli, C. zoae (Orchard), C. angulata Mazza, Cau and Rigo, position of the pit in respect to keel end. The lower side has a C. samueli (Orchard), C. pseudoechinata (Kozur), and C. gul- very narrow basal cavity, centrally located in juvenile speci- loae Mazza and Rigo. Orchard (2013) added C. pseudodiebeli mens to strongly forward-shifted in advanced forms. The posi- (Kozur), C. samueli (Orchard), C. zoae (Orchard), C. eozoae tion of the pit is advanced with respect to both platform and (Orchard), and C. medioconstricta (Orchard). keel end (Kozur, 2003). The robust platform generally lacks Consequently, as platform ornamentation became a key ornamentation. In some species a few nodes may be present, criterion in classification of species within the genus Meta- but they are always confined to the anterior platform or in cor- polygnathus, discrepancies could not be preserved under “one respondence to the geniculation point. A distinct free blade is roof”. This has resulted in the redefinition of existing genera, present, with a high carina with highly fused denticles. The keel such as Paragondolella, but also in the introduction of addi- end is often, but not necessarily, bifurcated. tional genera, such as Carnepigondolella. Orchard (2013) has In the case of M. samueli Orchard, which corresponds to introduced five new genera for mostly endemic Orchard’s (1991a) sp. G, Mazza et al. (2010) wrote that
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