Elsevier

Cretaceous Research

Volume 112, August 2020, 104448
Cretaceous Research

Scombroclupea javieri sp. nov., an enigmatic Cenomanian clupeomorph fish (Teleostei, Clupeomorpha) from the marine deposits of the Cintalapa Formation, Ocozocoautla, Chiapas, southeastern Mexico

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Abstract

Scombroclupea javieri sp. nov. is erected based on the description of well-preserved specimens collected in the lower Cenomanian marine outcrop at the El Chango quarry, near Ocozocoautla de Espinosa, Chiapas State, southeastern Mexico. This new species shows the distinctive feature of the superorder Clupeomorpha, the presence of the abdominal scute series, as well as the diagnostic characters of Scombroclupea, as the lack of the dorsal scutes and the presence of two supramaxillae, the comb-like dentition in the maxilla, the horizontal laminae in the anterior caudal vertebrae, the flat epineurals and epipleural in caudal region of the trunk, and the finlets present along the anal fin, and differs from the other three nominal species of this genus in meristic data of the fins and vertebrae. Scombroclupea was previously thought as an incertae sedis members of the order Clupeiformes; however, S. javieri displays two primitive features never recorded before in this clupeomorph order and commonly observed in the order Ellimmichthyiformes, this new species has a large beryciform foramen piercing the anterior ceratohyal and its parietal bones are sutured to each other. The review of specimens of S. occidentalis and S. macrophthalma confirms that these species share the presence of a large beryciform foramen in the anterior ceratohyal. These characters make it difficult to include Scombroclupea as a member of any of the orders Clupeiformes and Ellimmichthyiformes. This record of Scombroclupea, the oldest in America and the second in México, widens the geographical Cenomanian distribution of the genus along the Tethys Sea, from Middle East to southern North America.

Introduction

The nearly 400 living species of herrings, anchovies, and sardines, and their about 160 extinct allies species, constitute a diverse and widespread teleostean fish group formally named as the superorder Clupeomorpha (Greenwood et al., 1966, Nelson et al., 2016). Members of this group inhabit practically all aquatic environments around the world and its fossil record dates back to the Early Cretaceous (Grande, 1985, Helfman et al., 2009). For a long time this group of fishes were long treated as a “wastebasket” group (e.g. Berg, 1940, Jordan, 1923, Lecointre and Nelson, 1996; among others), however, Greenwood et al. (1966: 358) and later Grande (1985) recognized their naturalness. Until today, the general aspects of clupeomorphs classification as set by last author, it being grouped into the orders Ellimmichthyiformes and Clupeiformes. Despite the recent achieves on the knowledge of the interrelationships and the arrangement of a natural classification of the clupeomorphs; today, the early diversification of this group still problematic (e.g. Grande, 1985, Zaragüeta-Bagils, 2004, Alvarado-Ortega, 2014, Vernygora et al., 2016). Therefore, any contribution to the knowledge of the fossils of this group is relevant to solve these and other problems of its natural history.

Undoubtedly, Mexico is the North American territory with the richest, more diverse and better preserved clupeomorphs fossil record and counts with numerous sites with ages ranging between the Albian to Oligocene (Maldonado-Koerdell, 1949, Ovalles-Damián, 2004, Alvarado-Ortega et al., 2006, Alvarado-Ortega et al., 2008, González-Barba and Alvarado Ortega, 2009, Pérez-Sánchez and González-Rodríguez, 2010; among others). This was demonstrated by Than-Marchese and Alvarado-Ortega (2013), who launched a project to study the numerous clupeomorph fossils recently discovered in different locations of Mexico; hence, the aim of the present paper is to describe a peculiar fishes recovered from the Lower Cenomanian marine deposits of the El Chango quarry, previously reported by Than-Marchese et al., 2011, Than-Marchese et al., 2015, Than-Marchese et al., 2016. Although many of these Mexican fossils still require to be studied accurately; recently, significant strides have been made in this respect. Specimens of Diplomystus Cope, 1877, from an uncertain locality in northern Mexico were identified by Aguilera (1896); however, this report is doubtful because, after more than a hundred years, this genus was never found again this country. Applegate (1996) and Chang and Grande (1997) reported the presence of Diplomystus-like and Ellimmichthys Jordan, 1919, in the Albian limestones of the Tlayúa quarry, Puebla; however, Melgarejo-Damián, 2012, Melgarejo-Damián and Alvarado-Ortega, 2010 identified four potential new ellimmichthyifom species among these, of which they named Paraclupea seilacheri Alvarado-Ortega and Melgarejo-Damián (2017). Additionally, the most abundant clupeomorph fish in Tlayúa was named as Ranulfoichthys dorsonudum Alvarado-Ortega (2014). Triplomystus applegatei Alvarado-Ortega and Ovalles-Damián (2008), later renamed as Scutatoclupea applegatei by Bannikov (2015) and a specimen similar to Paraclupea Chang and Chou, 1977, were the first clupeomorphs studied from the Cenomanian deposits of the El Chango quarry, Chiapas (Ovalles-Damián, 2004, Alvarado-Ortega et al., 2008). Scombroclupea occidentalis Giersch, Frey, Stinnesbeck, Ifrim, and Padilla Gutierrez (2011), was described on with specimens recovered in the Cenomanian deposits of the Loma La Mula site, near Múzquiz, Coahuila.

The El Chango quarry together with its coeval and sister paleontological site, the El Espinal quarry, are real lagerstätten discovered in the first decade of the present century (Ovalles-Damián and Alvarado Ortega, 2002), near Ocozocoautla de Espinosa Municipality, Chiapas, southeastern Mexico (Fig. 1). In these small paleontological sites, a rich and well-preserved fossil assemblage has been recovered from the Cenomanian laminar and parallel limestones strata of the Cintalapa Formation, deposited under marine shallow and tropical marine conditions. The fossil assemblage as far recovered in the El Chango include plants, ammonites, bivalves, arthropods, and fishes (Alvarado-Ortega et al., 2009, Alvarado-Ortega and Than-Marchese, 2012b, Alvarado-Ortega and Than-Marchese, 2013, Díaz-Cruz et al., 2016, Guerrero-Márquez et al., 2012, Moreno-Bedmar et al., 2014, Vega et al., 2006; among others).

The genus Scombroclupea was erected by Kner (1863), to include his species S. pinnulata from Comen, today Slovenia, as well as S. macrophthalma (Heckel, 1849), previously described as Clupea macrophthalma from the Cenomanian deposits of Hakel and Hujula, Lebanon. Subsequently, the last of these species was removed the genus Clupea Linnaeus, 1758, and the first species was recognized as its junior synonymous (Woodward, 1901, Patterson, 1967, Patterson, 1970). Before this paper, the fossil record of Scombroclupea included three Cenomanian species with a geographical distribution at both sides of the Tethys Sea, in the Middle East-Europe and North America. The other nominal species of this genus are Scombroclupea diminuta Forey, Yi, Patterson, and Davis (2003), recently discovered in deposits of Namoura, Lebanon, and S. occidentalis Giersch, Frey, Stinnesbeck, Ifrim, and Padilla-Gutiérrez (2011), from the marine deposits of the “Loma La Mula” site, in Coahuila, northern Mexico [other authors (e.g. Alvarado-Ortega and Porras-Múzquiz, 2012) named this site as the La Mula quarry and suggest its age as Turonian]. Unfortunately, since Grande (1985) suggested that Scombroclupea is an incertae sedis clupeomorph, its phylogenetic relationships have never been properly evaluated.

Here we report a clupeomorph that presents characters that place it in the genus Scombroclupea, recognizing as fourth new and valid species, as well as the second for America. In addition, the gaol of this work is to present the evidence of complex morphological characters that support as incertae sedis genus due to its shared distribution with the main groups of Clupeomorpha.

Section snippets

Preparation methods

Part of the specimens described in this paper were prepared following the acid method and transfer technique (Toombs and Rixon, 1959). They were bathed into an aqueous solution of 5–15% of acetic acid and hardened with Plexygum. Needles and pin vises were used under the stereoscopic microscope to remove the remaining patches of sediments from the skeletons. Prepared specimens were coated with magnesium dust to get black and white photographs of high contrast under white light.

Anatomical abbreviations and measurements

The anatomical

Systematic paleontology

  • Superorder Clupeomorpha Greenwood et al., 1966

  • Order incertae sedis

  • Family incertae sedis

  • Genus Scombroclupea Kner, 1863

  • Scombroclupea javieri sp. nov

  • Holotype. IHNFG-5297, a complete specimen of 90 mm standard length exposing its left side (Fig. 2).

  • Paratypes. IHNFG-2990, a complete specimen preserved in part and contrapart. IHNFG-2992, complete specimen with the scales out of place and scattered on and around the trunk. IHNFG-5298 articulated specimen preserved in part and counterpart, with the

Discussion

Actually, the genus has three valid species: S. macrophthalma [Líbano (Hakel y Hajula), Slovenia (Komen) y Croatia], S. diminuta [Líbano (Namoura)] and S. occidentalis [Mexico (Coahuila)]. The Scombroclupea diagnosis was amended by Woodward, 1901, Forey et al., 2003 and Giersch et al. (2011). The latter defines the genus as follows: 1) two supramaxillae; 2) tiny teeth in the caudal half of the jaw that have a comb-like dentition [Initially observed by Woodward (1901) and removed by Forey et al.

Conclusions

Scombroclupea is a clupeomorph genus at to have ventral scutes, who is a synapomorphy characters of the clade. Scombroclupea javieri sp. nov. is a member of the genus by having a lack of the dorsal scutes and the presence of two supramaxilla, the comb-like dentition in the maxilla, the horizontal laminae in the anterior caudal vertebrae, the flat epineurals and epipleural in caudal region of the trunk and the finlets present along the anal fin, last are interpreted as diagnostic characters of

CRediT authorship contribution statement

Bruno Andrés Than-Marchese: Conceptualization, Data curation, Formal analysis, Methodology, Investigation, Project administration, Resources, Validation, Visualization, Writing - original draft, Writing - review & editing. Jesús Alvarado-Ortega: Conceptualization, Formal analysis, Funding acquisition, Methodology, Investigation, Project administration, Resources, Supervision, Validation, Visualization, Writing - original draft, Writing - review & editing. Wilfredo A. Matamoros:

Acknowledgements

Authors thanks Marco A. Coutiño José, Luis E. Gómez Pérez, Gerardo F. Carbot Chanona, who are on charge of the project Prospección y Resguardo del Patrimonio Paleontológico de Chiapas that allow the paleontological research in the El Chango quarry. Oksana Vernygora, Emilia Sfeco, Yuri Modesto and Rene Zaragüeta Bagils kindly provided us with important articles used in the paper. Likewise, Hector Porras lent us part of the Scombroclupea specimens of the studied in this investigation and gave us

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