Clupeomorph Fish Research Articles

Double-armoured herrings (Clupeomorpha: Ellimmichthyiformes) from the Lower Cretaceous of Pietraroja (Southern Italy)

The Lower Cretaceous (Albian) clupeomorph fishes of the Pietraroja Plattenkalk were among the first fishes described for this celebrated Italian Konservat-Lagerstätte. However, a detailed anatomical and phylogenetic study has never been carried out on these fishes in order to clarify their taxonomic status, systematic position, and relationships within the Clupeomorpha. The specimens examined herein exhibit diagnostic features that clearly support their alignment with the extinct double-armoured herring order Ellimmichthyiformes. Based on the presence of an incomplete series of heart-shaped predorsal scutes, a new combination, Armigatus elatus (Costa, 1850), and the new species Armigatus plinii sp. nov. are recognized as the oldest representatives of the family Armigatidae, whereas a unique combination of features allows us to recognize most of the specimens as a new representative of the family Paraclupeidae, Paraclupea pietrarojae sp. nov. The presence of lower Albian species of Armigatus and Paraclupea in the Western Tethys extends the temporal and biogeographical range of these widely distributed genera, providing additional information for our understanding of the evolutionary and biogeographical history of the double-armoured herrings. http://zoobank.org/urn:lsid:zoobank.org:pub:98F0ECD2-19C3-4399-89D2-7567058AE812

Revision of †Ellimmichthys longicostatus (Clupeomorpha: †Ellimmichthyiformes) from the Lower Cretaceous of Brazil with comments on the taxonomy of related species

†Ellimmichthys longicostatus is a deep-bodied and double-armored clupeomorph fish only known from non-marine Berriasian-Valanginian beds of the Recôncavo Basin, northeastern Brazil. It is the oldest herring-like fossil fish described from South America. This species was recently separated in two distinct forms that have not yet been formally classified: the first has the pelvic fin anterior to the origin of the dorsal fin, enlarged, thorn-shaped postpelvic scutes, posteriormost ribs on parapophyses, and long dorsal pterygiophores; the second one has the pelvic fin opposite the base of the dorsal fin and all the ribs inserted in deep excavations on centra. The latter corresponds to the fish used as a reference by Edward Drinker Cope, in 1886, in the original description of the species and herein referred as †Ellimmichthys longicostatus s. str. Consequently, the current diagnosis of †Ellimmichthys is not sufficient to distinguish species and closely related taxa. Based on new and old articulated specimens, we provide an amended diagnosis for †Ellimmichthys longicostatus s. str. and erect †Ellimmichthys spinosus nov. sp. for the rare form that was mistakenly identified to be the same species.

Morphological variation among the species of Armigatus (Teleostei, Clupeomorpha, Ellimmichthyiformes) and new material of Armigatus alticorpus from the Upper Cretaceous (Cenomanian) of Hakel, Lebanon

The clupeomorph genus Armigatus is an extinct group of fish which comprises five species of small marine fishes described from the Late Cretaceous Tethys Sea. Of the five species, A. alticorpus is the least well known and least studied species. The most recent cladistic analyses of the group exclude A. alticorpus due to a lack of sufficient information for the taxon. Here we report new material of Armigatus from the Cenomanian of Hakel, Lebanon. The mass mortality slab preserves multiple clupeomorph fishes including specimens of two species of Armigatus, A. namourensis and A. alticorpus, which have previously been known only from Namoura, Lebanon. The good preservation of skull elements and caudal skeletons of the A. alticorpus specimens provides necessary information for inclusion of this species in a phylogenetic analysis. Results of the phylogenetic reassessment of A. alticorpus indicate that the species had close affinities to the two younger species of the genus, A. oligodentatus and A. dalmaticus. The three species together (A. alticorpus (A. oligodentatus + A. dalmaticus)) belong to a separate lineage from their sister-clade, (A. brevissimus + A. namourensis). Comparative examination of the multiple specimens of each species enabled us to establish a set of morphological traits which allow a more reliable delimitation of the species of Armigatus; this includes number of abdominal scutes, vertebrae, predorsal bones, as well as position of the dorsal fin.

The relationships of Gasteroclupea branisai Signeux, 1964, a freshwater double-armored herring (Clupeomorpha, Ellimmichthyiformes) from the Late Cretaceous-Paleocene of South America

The clupeomorph fish Gasteroclupea branisai Signeux, 1964 from the Late Cretaceous-Paleocene of El Molino Formation, Cayara locality, Bolivia is redescribed in detail. This South American double-armored herring is the sole member of the family Gasteroclupeidae and exhibits a unique combination of features (skull roof ornamented with fine tubercles, beryciform foramen, endopterygoid teeth, about 16–18 supraneurals, a series of 35–38 predorsal and 40 ventral scutes, median fins small and containing 12–14 rays) that clearly supports its assignment to the extinct clupeomorph order Ellimmichthyiformes. G. branisai shares a number of synapomorphies with Sorbinichthys elusivo and S. africanus (five hypurals, second hypural autogenous, distal end of the second uroneural not reaching the distal end of the first uroneural) with which it forms a monophyletic assemblage recognized herein for the first time as the new suborder Sorbinichthyoidei. The peculiar morphology of Gasteroclupea that is characterized by having an extremely deep abdominal profile, hypertrophied coracoid, and elongated pectoral fins, resembles in many aspects the body plan of extant freshwater hatchetfishes of the characiform family Gasteropelecidae, suggesting that the morphological adaptations of G. branisai can be related, at least in part, to a similar lifestyle.

Relationships of †Codoichthys carnavalii Santos, 1994 (Teleostei, Clupeomorpha, †Ellimmichthyiformes) from the Late Aptian of São Luís-Grajaú Basin, NE Brazil.

†Codoichthys carnavalii is a clupeomorph fish only found in calcareous concretions of Codó Formation, State of Maranhão. It is known based on three specimens housed in the paleontological collection of the Museu de Ciências da Terra of Departamento Nacional da Produção Mineral, Rio de Janeiro. It was omitted in most of recent cladistic analyses about clupeomorphs. We revisited its anatomy furnishing new data and additional restorations. Furthermore we explored the relationships of †Codoichthys with the computer program TNT based on a matrix with 30 taxa and 60 unordered and unweight characters. Elops was used to root the tree. The strict consensus was obtained from three shortest trees (L=181; CI=0.387; RI=0.632). The monophyly of †Ellimmichthyiformes is supported by a sigmoid cleithrum and an uniquely derived predorsal scute series. Most of subgroups showed low support indices. †Sorbinichthyidae and †Horseshoeichthys appear in the most basal position, and not closely related to †Armigatus or †Diplomystus. A †Diplomystus clade is more advanced than †Armigatus and sister-group of remaining †ellimmichthyiforms. Within †Paraclupeidae, †Codoichthys is sister-group of remaining †paraclupeids (including †thorectichthyines and †paraclupeines). Within †Paraclupeinae, †Triplomystini includes a †Triplomystus clade, a sister group of †Rhombichthys plus †Tycheroichthys, and †Paraclupeini with †S. itapagipensis and all other †ellimmichthyiform taxa.

A new species of Sorbinichthys (Teleostei: Clupeomorpha: Ellimmichthyiformes) from the Late Cretaceous of Morocco

A new species of ellimmichthyiform fish, represented by three specimens, has been recovered from deposits of the Akrabou Formation of Morocco. The new species is described in the existing genus Sorbinichthys , family Sorbinichthyidae, as Sorbinichthys africanus , sp. nov., closely related to the type species Sorbinichthys elusivo . The Ellimmichthyiformes is an extinct order of clupeomorph fishes that includes both freshwater and marine species ranging from the Early Cretaceous through the Eocene. Sorbinichthys elusivo is known from Cenomanian deposits of the eastern Mediterranean Tethys (Lebanon), whereas the new species is possibly early Turonian but more probably late Cenomanian in age, from the western Tethys (eastern Morocco). At lower taxonomic levels, the assemblage from the Akrabou Formation has a primarily Tethyan composition, exemplified by the genus Sorbinichthys, whereas supra-generic taxa found there include groups with trans-Atlantic (e.g., Sorbinichthyidae, Macrosemiidae) or near-cosmopolitan (e.g., Paraclupeidae) distributions.

First record of a clupeomorph fish in the Neuquén Group (Portezuelo Formation), Upper Cretaceous of Patagonia, Argentina

A new genus and species of clupeomorph fish, Leufuichthys minimus, is described from the fluvial deposits of the Portezuelo Formation, Upper Cretaceous (Turonian–Coniacian) of the Neuquén Group, Patagonia, Argentina. It is a small-sized fish with an estimated body length up to 46mm. Among other characters, the new species shows the following: abdominal scutes; abdomen moderately convex; anal fin elongate-based; three uroneurals; two epurals; caudal fin bearing very elongate rays; and cycloid scales. Leufuichthys minimus gen. et sp. nov. shows a greater similarity with Kwangoclupea dartevellei, a clupeomorph described from a marine Cenomanian deposit of the Democratic Republic of Congo (Africa), mainly due to the presence of an elongate-based anal fin, bearing more than 20 fin-rays, differing from it by the presence of a not hypertrophied abdomen. As far as known, L. minimus gen. et sp. nov. is the first clupeomorph described in the Upper Cretaceous of Patagonia and represents one fortuitous preservation of an articulated fish in fluvial deposits.

A new clupeiform fish from the Lower Cretaceous (Barremian) of Sergipe-Alagoas Basin, northeastern Brazil

ABSTRACT Fossil remains of clupeomorph fishes are commonly found in Cretaceous and Tertiary deposits from the northeastern Brazil. The taxonomic diversity is relatively high and include †ellimmichthyiforms, early clupeiforms, and advanced clupeoids. However, well-preserved specimens are very scarce so that very little is known about affinities of most taxa. Recently, new localities containing clupeomorphs have been discovered and prospected in Alagoas State, in Northeastern Brazil. From them, Atol Quarry, in which Barremian shales from the Coqueiro Seco Formation crop out, has demonstrated to be meaningful by having a diverse fish assemblage. It contains holosteans, coelacanths, gonorynchiforms, and enchodontids besides clupeomorphs. Most of fishes correspond to new taxa. In this paper, an early clupeiform fish, †Pseudoellimma gallae gen. et sp. nov., is described from those shales. The fish shows many informative characters and revealed to be advanced in comparison with other early and relatively well-known clupeomorphs such as †Spratticeps and †Santanaclupea. Although lacking pleurostyle, it shares with clupeoids derived features such as accessory anterior foramen of temporal canal, hidden post-orbital branch of supraorbital canal on frontal, longitudinal and parallel ridges sculpturing the skull roof, and reduced first ural centrum. If so, many derived features found in advanced extant clupeoids appeared early in the history of clupeiform fishes. Furthermore the present study indicates divergence between Clupeoidei and Denticipitoidei lineages older than Barremian age.

A new marine clupeoid fish from the Lower Cretaceous of the Sergipe-Alagoas Basin, northeastern Brazil

Clupeomorph fishes are largely diversified and widespread in Cretaceous strata of northeastern Brazil. They are represented by basal clupeiforms, †ellimmichthyiforms, and advanced clupeoids. In this paper, a new clupeoid fish, †Nolfia riachuelensis sp. nov., is described based on a specimen found in marine shale yielded in the Taquari Member (Albian) of Riachuelo Formation, Sergipe-Alagoas Basin. Although partially preserved (most of skull is lacking), the fish shows many informative characters, particularly in the axial skeleton. The fish is morphologically very similar to †Nolfia kwangoensis, a clupeoid fish originally described from marine deposit from the Upper Cretaceous (Cenomanian) of Kipala, Democratic Republic of Congo, with which it shares: 18 caudal vertebrae; long and cylindrical pleural ribs; posteriormost pleural ribs supported by long parapophyses and gradually decreasing in size posteriorly; very reduced ural centra; presence of long and distally congruent neural spines of third, second, and first preural centra; three uroneural bones (the first probably forming the pleurostyle). Otherwise, †Nolfia riachuelensis sp. nov. differs from other clupeomorphs by a unique combination of features. The number of supraneural bones, aligned rod-like postcleithra, less than 30 rays in the anal fin, and derived absence of dorsal scutes indicate its placement within Clupeidae. As far as is known, †Nolfia riachuelensis sp. nov. is the most ancient clupeoid fish known from the fossil record.

Triplomystus applegatei, sp. nov. (Teleostei: Ellimmichthyiformes), a rare “triple armored herring” from El Espinal Quarry (Early Cretaceous), Chiapas, southeastern Mexico

ABSTRACT Triplomystus applegatei, sp. nov., is described as a new clupeomorph fish belonging to the order Ellimmichthyiformes, based on a single specimen collected from the Early Cretaceous (Albian) limestones of El Espinal quarry, near Ocozocuautla de Espinosa, Chiapas, southeastern Mexico. T. applegatei is differentiated from the other two species of the genus on the basis of body proportions and meristic data, as well as a peculiar development of a complex shape and articulation between the last scutes in the posterodorsal series. This species extends the geographical and geological domain of this genus from the Cenomanian Eastern Tethys Sea to Albian western Tethys Sea.

A new clupeid fish (Clupeomorpha) from the oligocene of Fayum, Egypt, with notes on some other fossil clupeomorphs

ABSTRACT A new genus and species of Clupeidae, Chasmoclupea aegyptica, is described from early Oligocene fluviatile deposits of the Jebel Qatrani Formation in the Fayum, Egypt. The new species is preserved in three dimensions, allowing details of the skull roof, as well as lateral skull bones, to be seen. The new species is considered to be most closely related to the Alosinae and Clupeinae. This is the first record of a clupeomorph fish from Oligocene freshwater deposits of Africa. Most clupeomorph fishes are preserved in marine or lacustrine sediments, and it is only the rare preservation of this riverine fossil, as an articulated specimen, that has allowed it to be identified as a clupeomorph.

A new clupeomorph fish from the Cretaceous Maceió Formation, Alagoas Basin, NE Brazil

A new clupeomorph fish from the Cretaceous Maceió Formation, Alagoas Basin, NE Brazil

An Elasmosaur with Stomach Contents and Gastroliths from the Pierre Shale (Late Cretaceous) of Kansas

A nearly complete skeleton of an elasmosaurid plesiosaur (NJSM 15435) from the Sharon Springs Member (Middle Campanian) of the Pierre Shale, Logan County, Kansas, is associated intimately with fragmentary fish remains and numerous gastroliths. The fish bones and gastroliths were located just behind the pectoral girdle in the abdominal region. Identifiable prey includes Enchodus and other small clupeomorph fishes. An isolated tooth of the anacoracid shark Squalicorax cf. S. pristodontus also was recovered in this area. Ninety-five gastroliths (6.8 kg) were present, with the largest stone measuring 15.1 × 8.5 × 5.7 cm (5.0 × 3.3 × 2.2 in.) and weighing 1.06 kg (2.3 lb.). Many of the gastroliths are composed of pink or gray Sioux Quartzite, which suggests that the source of these stones was about 600 km (475 mi) to the northeast of where the elasmosaur remains were discovered. The association of fragmentary fish remains and gastroliths within the abdomen of NJSM 15435 supports the contention that the stones aided in the breakdown of food in plesiosaurs.

A new early Cretaceous clupeomorph fish from the Arratia Valley, Basque Country, Spain

Fish-bearing beds of early Cretaceous age crop out in the Arratia Valley, province of Biscay, along the nucleus of the Bilbao Anticlinorium in the Basque-Cantabrian Basin, northern Spain. Teleost fossils have been found in several stratigraphic levels, dated as Valanginian–Barremian, in the Villaro Member of the Villaro Formation. The Villaro Member is a 1100-m-thick succession of black shales, sandstones and scattered limestones that represent deposition in a shallow lacustrine system periodically fed by delta prograding bars and channels. Deposition took place in a restricted freshwater–brackish environment that was occasionally affected by marine incursions. The first new taxon from the Arratia Valley, described here, isEzkutuberezi carmeni gen. nov., sp. nov., a deep-bodied clupeomorph whose autapomorphies include: a posterior supramaxilla with a long anterior process forming an angle of 90° with the main body of the bone; a hypertrophied, claw-shaped last dorsal scute; only two epurals; and hypural 3 with a submedian longitudinal crest. The morphology of the dorsal scutes supports its assignment to the paraclupeid subfamily Paraclupeinae within the order Ellimmichthyiformes. The new taxon is the oldest European clupeomorph, and the first articulated vertebrate fossil to be described from the Basque-Cantabrian Basin.

A remarkable clupeomorph fish (Pisces, Teleostei) from a new Upper Cretaceous marine locality in Lebanon

A new clupeomorph fish,Sorbinichthys elusivo n. g., n. sp. is described from a new Upper Cretaceous (Middle ? Cenomanian) marine locality at Nammoura (Lebanon). The characteristic features of the new genus, such as dorsally expanded posttemporal, subrhomboid spiny dorsal scutes, a very wide and deep gap between second and third hypurals, second ray of dorsal and pectoral fins greatly elongated and filamentous, and anterior procurrent caudal fin rays deeply extended between vertebral spines are unusual for clupeomorphs and warrant the establishment of the new family Sorbinichthyidae. The new fish displays several clupeomorph and clupeiform characters, but its placement within the clupeiforms is problematic. The new family is referred to the Clupeomorpha incertae ordinis.

Laterophysic Connection: A Unique Link between the Swimbladder and the Lateral Line System in Chaetodon (Perciformes: Chaetodontidae)

The auditory and lateral line systems of teleost fishes are complementary in their ability to detect mechanical stimuli in water (Kalmijn, 1988, 1989; Coombs, 1994). The lateral line system is normally sensitive only to local incompressible flow fields arising from predators, prey, neighbors in a school, and environmental obstacles (reviewed by Montgomery et al., 1995). In contrast, the otolithic organs of the inner ear respond to whole-body accelerations generated by the same local flow fields to which the lateral line system responds. Enhanced sensitivity of the inner ear to propagated pressure waves (sound) is a specialized function of the inner ear of teleost fishes (Kalmijn, 1989), which is the result of the presence of a volume of compressible fluid (i.e., the swimbladder) in the vicinity of the inner ear. Morphological specializations in which the swimbladder is brought into close or direct contact with the inner ear are generally known as otophysic connections and occur in some or all species in all four lineages of teleost fishes (Osteoglossomorpha, Elopomorpha, Clupeomorpha, Euteleostei, reviewed by Schellart and Popper, 1992). It seems reasonable that the lateral line system could also be made pressure sensitive by placing an air cavity in the vicinity of, or in direct contact with, a lateral line canal. There are only two known examples of this type of sensory specialization: the recessus lateralis of clupeomorph fishes; and the unique connection between the inner ear and the lateral line

A fossil clupeomorph fish from the Albian of the Northwest Territories of Canada, with notes on cladistic relationships of clupeomorphs

A primitive clupeomorph, Erichalcis arctagen. et sp. nov. is described from the Albian of Northwest Territories, Canada. Erichalcis is uniquely specialized among clupeomorphs in the presence of enlarged lateral line scales and in the structure of the caudal skeleton. Erichalcis is classified as Clupeiformes incertae sedis. The two recognized suborders, Clupeoidei and Denticipitoidei are examined with respect to primitive and derived character states. It is concluded that the clupeoids and denticipitoids are divergently specialized in details of the union between the supraorbital and infraorbital sensory canals and the recessus lateralis. It is further suggested that the reduction in size and the posterior shift of the dermosphenotic in clupeoids is correlated with an upper division of the levator arcus palatini muscle and an antero‐dorsal process on the hyomandibular. Cladistic relationships of some Cretaceous clupeomorphs are proposed and it is suggested that Diplomystus brevissimus Cope not be included in the Clupeoidei.

A Primitive Clupeomorph Fish from the Middle Cenomanian of Hakel, Lebanon

A primitive clupeomorph fish Ornategulum gen. nov. sardinioides (Pictet) is described from the middle Cenomanian of Hakel in the Lebanon. The caudal skeleton shows a free first hypural, a fusion between the second hypural and the reduced first ural centrum, and a complete neural spine associated with the second preural centrum. The cranial anatomy shows this genus to be the most primitive clupeomorph yet known, lacking a temporal foramen, a well-defined pre-epiotic fossa, frontal fontanelles, a pterotic bulla, and a recessus lateralis. The fish is too primitive to be placed in either suborder of the Clupeomorpha and there is no firm evidence to suggest that it belongs to a separate clupeomorph lineage; the genus is placed as Clupeomorpha incertae sedis.

A clupeomorph fish from the Gault (Lower Cretaceous)

A primitive clupeomorph fish, Spratticepsgaultinus gen. et sp. nov., is described on the basis of incomplete braincases from the Gault (Albian) of Folkestone, Kent. This is the earliest clupeomorph in which any details of cranial structure are known. Spratticeps is more primitive than living clupeomorphs in having no recessus lateralis, auditory fenestra, pterotic bulla or sinus temporalis, and in having a heavily ornamented, entire skull roof, with the sensory canals bone enclosed and the post-temporal fossa roofed. The fish is too primitive to be placed in either suborder of Clupeomorpha, and is left incertae sedis. The primary characters of the Clupeomorpha are discussed.