Scombroclupea javieri sp. nov., an enigmatic Cenomanian clupeomorph fish (Teleostei, Clupeomorpha) from the marine deposits of the Cintalapa Formation, Ocozocoautla, Chiapas, southeastern Mexico
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
References (56)
Ancient herring from the Tlayúa Quarry (Cretaceous, Albian) near Tepexi de Rodíguez, Puebla State, central Mexico, closing the gap in the early diversification of Clupeomorpha
Cretaceous Research
(2014)- et al.
The Cenomanian short snout enchodontid fishes (Aulopifomes, Enchodontidae) from Sierra Madre Formation, Chiapas, southeastern Mexico
Cretaceous Research
(2016) - et al.
Clupeomorpha, sister-group of Ostariophysi
- et al.
Ammonite age-calibration of the EL Chango Fossil-Lagerstätte, Chiapas state (SE Mexico)
Journal of South American Earth Sciences
(2014) Sinopsis de Geología Mexicana
- et al.
Mesozoic Osteichthyans of Mexico
- et al.
Triplomystus applegatei sp. nov. (Teleostei:Ellimmichthyiformes), a rare “Triple armored herring” from El Espinsal Quarry (Early Cretaceous), Chiapas, Southeastern Mexico
Journal of Vertebrate Paleontology
(2008) - et al.
A review of the interrelationships of the order Ellimmichthyiformes (Teleostei: Clupeomorpha)
- et al.
The fossil fishes from the Sierra Madre Formation, Ocozocoautla, Chiapas, Southern Mexico
Palaeontologia Electronica
(2009) - et al.
The first American record of Aspidopleurus (Teleostei, Aulopiformes) from La Mula quarry (Turonian), Coahuila State, Mexico
Revista Brasileira de Paleontologia
(2012)
A Cenomanian aipichthyoid fish (Teleostei, Acanthomorpha) from America, Zoqueichthys carolinae gen. and sp. nov. from El Chango quarry (Cintalapa Member, Sierra Madre Formation), Chiapas, Mexico
Revista Mexicana de Ciencias Geológicas
Paraclupea seilacheri sp. nov., a double armored herring (Clupeomorpha, Ellimmichthyiformes) from the Albian limestones of Tlayúa quarry, Puebla, Mexico
Revista Mexicana de Ciencias Geológicas
The first record of a North American Cenomanian Trachichthyidae fish (Acanthomorpha, Acanthopterygii), Pepemkay maya, gen. et sp. nov., from El Chango Quarry (Sierra Madre Formation), Chiapas, Mexico
Journal of Vertebrate Paleontology
An overview of the quarries near Tepexi de Rodríguez, Puebla, México
Basal teleosts and teleostean phylogeny
Palaeo-Ichthyologica
A new genus for the Cenomanian ellimmichthyiform fishes from Lebanon and Mexico
Bollettino Paleontologica, Societa Academy
Ueber die Morphogenie der Wirbelsäule der Amnioten
Biologisches Centralblatt
Classification of fishes, both Recent and fossil
Trudy Zoologicheskogo Instituta, Akademiya Nauk SSSR, Leningrad
On Late Mesozoic fossil fishes from Zhejiang Province, China
Memoirs of the Institute of Vertebrate Paleontology and Paleoantropology, Academia Sinica
Redescription of Paraclupea chetungensis, an Early Cretaceous Clupeomorph from the Lower Cretaceous of southeastern China. Fieldiana, 1489
Geology, New Series
Redescription of Ellimma branneri and Diplomystus shengliensis, and relationships of some basal clupeomorphs
American Museum Novitates
A contribution to the knowledge of the ichthyological fauna of the Green River Shales
Bulletin of the United States Geological and Geographical Survey
Homology between the recessus lateralis and cephalic sensory canals, with the proposition of additional synapomorphies for the Clupeiformes and the Clupeoidei
Zoological Journal of the Linnean Society
A three-dimensional skull of a primitive clupeomorph from the Cenomanian English Chalk, and implications for the evolution of the clupeomorph acusticolateralis system
Fossil fishes from the Cenomanian (Upper Cretaceous) of Namoura, Lebanon
Journal of Systematic Palaeontology
Scombroclupea occidentalis sp. nov. (Clupeiformes, Teleostei) from the Cenomanian (Cretaceous) Plattenkalk deposits of NE Mexico
Swiss Journal of Geosciences
Los peces Clupeiformes del Oligoceno Tardío (Chattiano) en las Formaciones EL Cien y San Gregorio de Baja California Sur, México. XI Congreso Nacional de Paleontología, Centro de Geociencias, Campus UNAM-Juriquilla, Queretaro, 25 -27 de febrero del 2009
A revision of the fossil genus Diplomystus, with comments on the interrelationships of clupeomorph fishes
American Museum Novitates
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Armigatus felixi sp. nov. An Albian double armored herring (Clupeomorpha, Ellimmichthyiformes) from the Tlayúa lagerstätte, Mexico
2022, Journal of South American Earth SciencesCitation Excerpt :Mexico has been revealed as a region with numerous Cretaceous paleontological sites bearing diverse and abundant fish assemblages that help to minimize such bias in the fossil record (Alvarado-Ortega, 2014; Alvarado-Ortega et al., 2006). The potential new basal clupeomorph species from Mexico that wait for accurate descriptive studies include at least two species from Albian marine deposits of Tlayúa (Puebla), one from the Albian-Cenomanian marine strata of the Muhi Quarry (Hidalgo), and two marine species from the Cenomanian sites of the El Chango and the El Espinal quarries (Chiapas), and one from the Turonian deposits of Huehuetla Quarry (Puebla) (Pérez-Sánchez and González-Rodríguez, 2010; Than-Marchese et al., 2020; Alvarado-Ortega, pers. observ.).
Apuliadercetis gonzalezae sp. nov., a North American Campanian dercetid fish (Teleostei, Aulopiformes) from Tzimol, Chiapas, Mexico
2022, Cretaceous ResearchCitation Excerpt :The eastern-western marine biotic interchange of the Tethys Sea was established since the Jurassic and continued up to the Cretaceous (e.g. Smith, 1983; Hallam, 1983; Damborenea, 2000; Gasparini and Fernández, 2005; Cavin, 2008). This biotic process involved marine reptiles (ichthyosaurs, metriorhynchids, and pliosaurs) and fishes (lepisosteiforms, macrosemiids, ophiophsids, ichthyodectiforms, ellimmichthyiforms, and enchodontids, among others) recently described from different Mexican Jurassic and Early Cretaceous paleontological sites (e.g. Blanco-Piñón, 2003; González-Rodríguez et al., 2004, 2016; Fielitz and González-Rodríguez, 2010; Giersch, 2014; Barrientos-Lara et al., 2015, 2016, 2018; Brito et al., 2017; Cantalice et al., 2019; Baños-Rodríguez et al., 2020; Díaz-Cruz et al., 2019, 2020; Than-Marchese et al., 2020). An important pending task, beyond the scope of the present work, is to carry out a comprehensive biogeographic study to include vertebrate and invertebrate marine taxa with allied species and genera distributed in the western and eastern regions of the Tethys Sea.