Messinian Lago-Mare deposits near the Strait of Gibraltar (Malaga Basin,S Spain)

A new stratigraphic unit (named LM unit) located at the base of the Pliocene marine succession in the eastern sector of the Malaga Basin has been distinguished. This unit is composed of two fining-upward sequences bounded by a discontinuity related to synsedimentary tectonic activity. Both sequences are made up of basal alluvial deposits overlain by subaquatic deposits. The lower sequence contains rich mollusc faunas characterized by bivalves Lymnocardiinae and Dreissenidae of Paratethyan origin, and non-marine gastropods, typical of oligo-low mesohaline shallow waters, and low diversified ostracod faunas of Paratethyan origin (Loxocorniculina djafarovi, Tyrrhenocythere pontica, Euxinocythere (Maeotocythere) praebaquana and Amnicythere propinqua) typical of low mesohaline waters. Alternances of monospecific ostracod assemblages made of Cyprideis agrigentina or oligotypic assemblages made of C. agrigentina and Loxoconcha spp. and richer assemblages made of L. djafarovi accompanied by several Amnicythere spp. and Camptocypria sp.1 characterize the upper sequence, pointing to more unstable environment affected by salinity and depth changes. Some samples collected from this upper sequence yielded rare, planktonic and benthic foraminifera, which could suggest the sporadic establishment of quasi-normal marine conditions in the westernmost Mediterranean during the deposition of this sequence. The palaeontological data and the palaeomagnetic results obtained in both the LM unit and the overlying Pliocene marine sediments restrict the age of the former to the post-evaporitic late Messinian. The age and the environmental conditions correlate these deposits with the Lago-Mare event occurring throughout the Mediterranean before the quick and widespread marine flooding of the Mediterranean at the beginning of the Pliocene.     

Late Messinian rodents from Verduno (Piedmont,NW Italy): Biochronological,paleoecological and paleobiogeographic implications

The stratigraphic and paleoenvironmental context of the Verduno fossil vertebrate locality is discussed herein based on its rodent record. The Verduno section crops out in the southern part of the Tertiary Piedmont Basin (TPB), and can be included in the Messinian post-evaporitic Cassano Spinola Fm., chronologically corresponding to the so-called Lago-Mare event. Rodents are represented by a relatively rich assemblage. Murids are by far the most diverse and abundant, with at least four taxa, including the common Centralomys benericettii and Paraethomys meini, and the rare Apodemus gudrunae and Occitanomys sp. Cricetids are represented by a single species, Apocricetus cf. A. barrierei. Muscardinus aff. M. vireti appears to be the only glirid present at Verduno. The Verduno rodent assemblage shares some taxa with other Messinian post-evaporitic localities from Italy bearing continental vertebrate remains, such as Brisighella (central Italy) and Moncucco Torinese (NW Italy) (e.g., C. benericettii, P. meini) and, possibly, with Ciabòt Cagna (NW Italy). However, the general structure of these four Messinian assemblages displays substantial differences, which may reflect different palaeoenvironmental conditions.     

Messinian Lago-Mare mollusc fauna from the Gorgona Island slope, Tyrrhenian Sea

Late Miocene Lago-Mare macrofossiliferous sediments were recovered in the northeastern Tyrrhenian Sea by dredging the continental slope off Gorgona Island, Tuscan Archipelago, at 300-470 m depth. The fossil assemblage consists of a rich lymnocardiid bivalve fauna dominated by Pontalmyra ex gr. P. incerta (Deshayes), associated with Dreissena ex gr. D. rostriformis (Deshayes), Pontalmyra cf. partschi (Mayer), “Limnocardium” sp., the gastropods Melanopsis narzolina (D’Archiac), Melanopsis sp. and cf. Saccoia sp. All bivalve taxa recognized at species level are of Paratethyan (Pontian) affinity and widespread in the Late Miocene of the Mediterranean Basin while M. narzolina has so far only recorded from the Mediterranean Basin. This finding represents the most diverse Lago-Mare macrofauna reported thus far from any submerged location in the Mediterranean Basin and documents that the post-evaporitic Cusercoli Formation contributes to the syn-rift neoautochthonous units of this sector of the Northern Tyrrhenian Sea.     

«Pachylebias crassicaudus (Agassiz) (Poisson téléostéen,cyprinodontiforme), un constituant majeur de l'ichthyofaune du Messinien continental du bassin méditerranéen

«Pachylebias crassicaudus (Agassiz) is a fossilfish widely distributed throughout the continental Messinian of Italy and Crete. Its detailed anatomical study permits the assessment of its systematic status within the subfamily Cyprinodontinae. A new locality with P. crassicaudus is pointed out in Italy, in the vicinity of Bologna. The fact that the same species occurs also in the gypsum deposits of Western Crete (Greece) leads one to suggest that these gypsums are Messinian in age. Finally, the palaeoecological study of P. crassicaudus makes possible the definition of the sedimentation conditions of the sediments belonging to the «gypsumsulphur formation.     

The genus Cyprideis Jones, 1857 (Crustacea,Ostracoda) in the Neogene of Italy: A geometric morphometric approach

As a result of the use of geometric morphometrics, multivariate statistics and classical morphological observations the authors propose a taxonomic revision of the Neogene Italian Cyprideis species. On this basis, the Cyprideis valves collected in Tortonian, Messinian and Pliocene brackish deposits of Italy are referred to 12 species, six of which are new: Cyprideis alexandri nov. sp., Cyprideis arvedoi nov. sp., Cyprideis lepianensis nov. sp., Cyprideis rooki nov. sp., Cyprideis strollae nov. sp., and Cyprideis toscana nov. sp. Some species are rather widespread in the Italian Peninsula (Cyprideis ruggierii Decima and Cyprideis crotonensis Decima) or even in the whole Palaeomediterranean (Cyprideis agrigentina Decima and Cyprideis anlavauxensis Carbonnel). For C. ruggierii, C. agrigentina and C. crotonensis, a phyletic relation is suggested. The new species are confined both geographically and stratigraphically in several Tuscan athalassic brackish basins, suggesting endemic speciation similar to those occurring in ancient lakes.     

La microflore de diatomées au passage mio-pliocène en Corse

The “Aleria formation”, described in northeastern Corsica, consists of siliciclastic deposits referred to a deltaic environment and comprises diatomitic lenses exposed at Casabianda. The Aleria formation fills a depression of the Messinian erosional surface and is overlain by Early Pliocene sediments in several places. This suggests a late Messinian-earliest Pliocene age. Diatom microflora is well-preserved and contains 59 species. Pennates indicate a very large diversity with 54 species, while Centrics are represented only by five species. Diatom assemblages show a relative constant composition along the studied interval. The species Aulacoseiragranulata is the dominant taxon with about 90% in all samples. Taxa are of different salinity classes and different modes of life (planktic, tychoplanktic, benthic, epiphytic…), suggesting multiple environmental factor interactions. The environmental model suggested by sedimentological and biological data led as to assume a pond-like coastal receptacle common in deltaic system. This receptacle was slightly influenced by the close marine waters and was supplied by permanent freshwaters from continental run-off and alluvial sheet. This model based on diatom assemblages provides new data for palaeoenvironmental and chronological interpretations of the so-called Messinian Lago-Mare deposits.     

Paracamelus (Mammalia,Camelidae) remains from the late Messinian of Italy: insights into the last camels of western Europe

In this paper we describe fossil remains of an indeterminate species of the genus Paracamelus (Artiodactyla, Camelidae) from the Messinian post-evaporitic deposits (5.55–5.40 Ma) of Verduno (Piedmont, NW Italy). Camelins dispersed into Eurasia from North America in the late Miocene and almost instantaneously spread in western Europe and Africa. The size and morphology of the fossils found at Verduno are consistent in with those of Paracamelus, the earliest Old World camelin. Up to now, the only fossil camels recovered in the Neogene of Western Europe have been found at Venta del Moro and Librilla in Spain at 6.2 Ma. The remains from Verduno represent the first evidence of a camelin in the Neogene of Italy and they considerably expand the paleobiogeographic range of the Old World early camelins. The presence of a camelid at Verduno reinforces and confirms the importance of the fossiliferous deposits of NW Italy in defining the complex paleobiogeographic patterns of Europe during the Messinian, at the end of the Messinian salinity crisis.     

Latest Messinian “Lago-Mare” Lymnocardiinae from Italy: Close relations with the Pontian fauna from the Dacic Basin

Rich hypo- to mesohaline molluscan assemblages characterising the latest Messinian “Lago-Mare” biofacies, composed of prosobranch gastropods (Neritidae, Thiaridae, Melanopsidae, Hydrobiidae) and bivalves of the families Cardiidae (subfamily Lymnocardiinae) and Dreissenidae, are widespread in shallow water basins characterized by low salinities within the Mediterranean realm, during the post-evaporitic phase in the time-span 5.5-5.3 Ma. Several genera and species are recorded in the Italian uppermost Messinian sediments. While the gastropods show endemic character being linked to continental water-systems, Lymnocardiinae and Dreissenidae have strong Paratethyan affinity. New records of significant species of Lymnocardiinae from the uppermost Messinian sediments of Tuscany, Marches and Sicily and the systematic review of the old literature data point out close relations of the Italian fauna with that from the Pontian sediments of the Dacic Basin. The palaeobiogeographical data referred to Messinian and Pontian Lymnocardiinae suggest that the Aegean Basin could be an intermediate basin from whence the Pontian Paratethyan-type fauna migrated into the Mediterranean area during the latest Messinian. Since the ecology of Lymnocardiinae is mainly tied to oligo- and mesohaline water, spreading of suitable habitats in depositional systems of marginal settings characterized by increasing freshwater influx after the “salinity crisis” favoured their dispersal into the Mediterranean area from the Paratethys realm.     

Paratethyan Ostracod immigrants in Italy during the Late Miocene

In this paper the ostracod assemblages recovered from several brackish Late Miocene Italian deposits have been analysed from a palaeobiogeographical perspective. During late Tortonian-early Messinian it is possible to recognize in Italy rich ostracod assemblages characterized by a wide contingent of taxa with central European or Mediterranean affinity, while only few brackish and freshwater ostracods show Paratethyan affinity. The recognized composition of the ostracod assemblages matches the palaeogeographic setting of the palaeo-Mediterranean/Paratethys at that moment. In fact during late Tortonian-early Messinian the palaeo-Mediterranean and Paratethysian domains were divided and, even if the connection via the present Marmara Sea-Strimon Basin was still open, the different salinity between them represented an ecological barrier, preventing faunal exchanges. Since normal aquatic migration was impossible, it must be assumed that the Paratethyan-like taxa entered the palaeo-Mediterranean area via passive dispersal by aquatic birds. On the contrary, the ostracod assemblages from the Italian Lago-Mare deposits show the absolute predominance of Paratethyan taxa, which, according to the known palaeogeographic setting during the late Messinian Lago-Mare event, could actively migrate from the Paratethys domain, colonizing the palaeo-Mediterranean, whose endemic fauna was severely impoverished by the Messinian salinity crisis and the following water dilution.     

First record ofBorelis melo andDendritina sp. in the Messinian of SE Spain (Cabo de Gata, Province Almeria)

Borelis melo andDendritina sp. occur in Messinian carbonates of the Cabo de Gata near Almeria. This is the first record of these larger benthic foraminifera from Messinian deposits in SE Spain. Two foraminiferal assemblages are differentiated: firstly, aBorelis- Dendritina assemblage which occurs on reef flanks, and secondly a lagoonal monospecificDendritina fauna.     

Coastal evolution and Holocene ostracods in Melides lagoon (SW Portugal)

A multidisciplinary study covering sedimentology (texture, carbonate and organic matter content, clay mineralogy), geochemistry (major, minor and trace elements, isotopes), palaeoecolgy (ostracods) and isotopic dating (¹⁴C) has been carried out on two long cores (MB-40.0 m and MIGM-34.0 m) retrieved from the barrier and the alluvial plain margining the Melides lagoon (south-western coast of Portugal). The interpretation of the whole data set allowed the establishment of an evolutionary model for this lowland since the Late Glacial. This model includes four stages. It starts with a fluvial terrestrial environment, replaced by an estuarine to fully marine one (circa 9500 cal BP), due to the rapid rise in sea level recorded during the first phase of the Holocene transgression. The deceleration of sea level rise favoured the emplacement of a sandy barrier (circa 5900 cal BP) that promoted the differentiation of a coastal lagoon which has been progressively filled with fluvial sediments.The ostracod fauna in both locations includes 38 species of littoral to sublittoral/phytal marine forms (mainly Urocythereis britannica, Carinocythereis whitei, Loxoconcha rhomboidea, Pontocythere elongata, Semicytherura sella, Semicytherura acuta, Basslerites cf. berchoni, Hiltermannicythere emaciata) and brackish water forms (Cyprideis torosa, Loxoconcha elliptica), which clearly characterize the signature of both the marine and lagoonal episodes, respectively.     

Sierra del Colmenar 1A,a new late Messinian (Late Miocene) locality in the Bajo Segura Basin (SE Spain): biostratigraphic and palaeoenvironmental implications

Abstract

In this paper, we have studied a new micromammal site from the Sierra del Colmenar section (Elche, SE Spain), named Sierra del Colmenar 1A (SCO-1A), representing the uppermost levels of Messinian age of the Bajo Segura Basin. The sedimentary context of this locality corresponds to a costal lagoon with marine influence. The fossil site has yielded remains of Apodemus aff. gorafensis, Paraethomys meini, Apocricetus alberti, Occitanomys alcalai, Ruscinomys sp., Eliomys cf. truci, Muscardinus sp., Parasorex ibericus, Prolagus michauxi and Soricidae indet. Based on this assemblage, we propose a Late Miocene age (MN13) for the locality SCO-1A, matching the previously inferred age for the stratigraphic unit in which the fossil site is situated. According to its faunal composition, we infer warm and humid climate conditions in the moment of formation of this locality, agreeing with the environmental conditions of the Messinian age in a parallic sedimentary context.     

Late miocene magnetobiostratigraphy of Crete

Six Upper Miocene marine clay sections on Crete (Greece) have been subjected to a detailed magnetobiostratigraphic analysis. Six correlatable polarity zones are recognized and these demonstrate the regional synchrony of planktonic foraminiferal biohorizons.By way of correlation to the magnetic polarity time-scale of Lowrie and Alvarez (1981), the Cretan sequence is assigned to polarity chronozones 5 (anomaly 3A) and 6. The new chronology provides an age of 5.6 Ma for the first occurrence datum (FOD) of theGloborotalia conomiozea group in the Mediterranean, an age of 6.0 Ma for the FOD ofG. menardii form 5 and an age of 6.6 Ma for the last occurrence datum (LOD) ofG. menardii form 4.Correlating the polarity record of the New Zealand Blind River section (Kennett and Watkins, 1974), with the magnetic polarity time-scale provides an age of 6.0 Ma for the evolutionary appearance ofGloborotalia conomiozea, which is in complete agreement with the age of 6.1 ± 0.1Ma given by Loutit and Kennett (1979). The demonstrated diachrony of 0.4 Ma between the New Zealand FOD ofG. conomiozea and its Mediterranean counterpart is explicable in view of the different nature of the two events, the one in New Zealand being evolutionary and the one in the Mediterranean migrational.The FOD of theG. conomiozea group in the Tortonian/Messinian boundary stratotype section coincides with the level proposed by Colalongo et al. (1979) to mark the base of the Messinian. Since the FOD of theG. conomiozea group in Crete and in Sicily are most probably time-equivalent, the age of the Tortonian/Messinian boundary is fixed at 5.6 Ma.The youngest sediments incorporated in this study extend into the Gilbert chronozone and antedate the main evaporitic phase. Consequently, the Messinian evaporitic body is younger than the base of the Gilbert chronozone, the age of which is fixed at 5.3 Ma. Adopting an age of 5.0 Ma for the Miocene/Pliocene boundary would imply that evaporites and post-evaporitic Lago Mare sediments were deposited in some 300,000 years and suggests that in the central parts of the Mediterranean basins evaporites must have accumulated at rates of some 3 m per 1000 years.     

Lower Pleistocene stratigraphy of the Burdur Basin of SW Anatolia

The Burdur Basin is one of the NE-trending extensional orogen-top basins of SW Anatolia hosting alluvial-fan, fluvial and lacustrine deposits from the Late Miocene onward. The remains of Equus (Allohippus) sp., Paracamelus cf. gigas, and medium-sized deer (Cervidae gen. indet.) from the uppermost reach of the basin-fill succession show an interregional palaeobiogeographical linkage. The composition of fossil associations is typical for the Villafranchian of eastern Europe and central Asia. The architecture of the basin-fill stratigraphy and associated fossil taxa refines the Early Pleistocene regional palaeogeography and biodiversity of Anatolia drawn over the Eurasian migration pattern.     

Tarbellastraea (Scleractinia): A new stable isotope archive for Late Miocene paleoenvironments in the Mediterranean

Geochemical proxy records of sea surface temperature (SST) or sea surface salinity (SSS) variability on intra- and interannual time-scales in corals from geological periods older than Pleistocene are extremely rare due to pervasive diagenetic alteration of coralline aragonite. Very recently, however, stable isotope data (δ¹⁸O, δ¹³C) from specimens of Porites of Late Miocene age (10 Ma) have been shown to preserve original environmental signatures. In this paper we describe new finds of the zooxanthellate corals Porites and Tarbellastraea in exceptional aragonite preservation from the island of Crete in sediments of Tortonian (～ 9 Ma) and Early Messinian (～ 7 Ma) age. Systematic, comparative stable isotope analysis of massive Tarbellastraea and Porites sampled from the same beds and localities reveal identical stable isotope fractionation patterns in both genera. Therefore, extinct Tarbellastraea represents an additional environmental archive fully compatible and mutually exchangeable with Porites. Provided that seasonal variations in δ¹⁸O reflect SST changes only, seasonal SST contrasts of 7.3 °C for the Tortonian and 4.8 °C for the Early Messinian are inferred, implying warmer summer and cooler winter SSTs during the Tortonian than during the Messinian. However, reduced δ¹⁸O seasonality (1.1‰ in the Tortonian and 0.7‰ in the Messinian) and slightly less negative mean δ¹⁸O in Messinian corals (? 2.4‰) compared to Tortonian specimens (? 2.7‰) may not necessarily indicate a long-term fall in SSTs, but changes in surface water δ¹⁸O, i.e. global ice build-up or enhanced evaporation during summer or increased precipitation/river discharge during winter and changes in insolation. On the other hand, coral communities of Tortonian and Messinian age in central Crete are identical, and compatible annual extension rates indicate similar average SSTs during the two investigated time periods. In addition, lithological and paleobotanical data from Central Crete document a change from humid to dry climatic conditions during the Late Miocene. Therefore, a likely explanation for the observed shift in coral mean δ¹⁸O and reduced δ¹⁸O seasonality from the Tortonian to the Early Messinian is a change in ambient seawater δ¹⁸O caused by a change in the hydrological balance towards high evaporation/high salinity during summer.     

Messinian bottom-grown selenitic gypsum: An archive of microbial life

Primary gypsum deposits, which accumulated in the Mediterranean Basin during the so-called Messinian salinity crisis (5.97–5.33 Ma), represent an excellent archive of microbial life. We investigated the molecular fossil inventory and the corresponding compound-specific δ¹³C values of bottom-grown gypsum formed during the first stage of the crisis in four marginal basins across the Mediterranean (Nijar, Spain; Vena del Gesso, Italy; Heraklion, Crete; and Psematismenos, Cyprus). All studied gypsum samples contain intricate networks of filamentous microfossils, whose phylogenetic affiliation has been debated for a long time. Petrographic analysis, molecular fossil inventories (hydrocarbons, alcohols, and carboxylic acids), and carbon stable isotope patterns suggest that the mazes of filamentous fossils represent benthic microbial assemblages dominated by chemotrophic sulfide-oxidizing bacteria; in some of the samples, the body fossils are accompanied by lipids produced by sulfate-reducing bacteria. Abundant isoprenoid alcohols including diphytanyl glycerol diethers (DGDs) and glycerol dibiphytanyl glycerol tetraethers (GDGTs), typified by highly variable carbon stable isotope composition with δ¹³C values spanning from ?40 to ?14‰, reveal the presence of planktic and benthic archaeal communities dwelling in Messinian paleoenvironments. The compound inventory of archaeal lipids indicates the existence of a stratified water column, with a normal marine to diluted upper water column and more saline deeper waters. This study documents the lipid biomarker inventory of microbial life preserved in ancient gypsum deposits, helping to reconstruct the widely debated conditions under which Messinian gypsum formed.     

Late Miocene brackish Loxoconchidae (Crustacea, Ostracoda) from Italy

The Late Miocene Italian brackish Loxoconchidae are herein discussed and illustrated. Three genera and two subgenera have been recognized in the brackish Italian basins: Loxoconcha, Loxocorniculina, Loxoconchissa (Loxoconchissa) and Loxoconchissa (Loxocaspia). Taking into account the diagnostic characters of several Loxoconchidae genera, in this paper Loxocorniculina is raised at a generic rank, Loxocaspia is confirmed as a subgenus within genus Loxoconchissa and several new species are established: Loxoconchissa (Loxoconchissa) kinoi nov. sp., Loxoconchissa (Loxocaspia) cosentinoi nov. sp., Loxoconchissa (Loxocaspia) nuda nov. sp., Loxoconchissa (Loxocaspia) punctata nov. sp., Loxoconchissa (Loxocaspia) reticulata nov. sp., Loxoconchissa (Loxocaspia) tuberosa nov. sp. and Loxoconchissa (Loxocaspia) velonae nov. sp. Loxocorniculina is a typical Paratethyan genus which widespread into the Palaeo-Mediterranean during the late Messinian Lago-Mare event. Loxoconchissa was known to be widespread only in the Paratethyan realm and in this paper it is signalled for the first time in the Late Tortonian-early Messinian of Italy. The palaeobiogeography of these genera is discussed and the observed continuous distribution of Loxocorniculina against the disjunct distribution of Loxoconchissa leads to suggest that this latter genus underwent a passive dispersal via aquatic birds.     

Late Miocene ostracod assemblages from eastern Mediterranean coral reef complexes (central Crete,Greece)

Ostracods from ten Late Miocene coral reef complexes built by Siderastrea, Tarbellastrea and Porites, cropping out in the Messara Plain (southern Iraklion basin, central Crete), have been investigated and five assemblages have been recognised, which point to different marine environments: (1) assemblage from the basal sandy silts, dominated by very shallow inner-infralittoral species, such as Cyamocytheridea meniscus, Cyamocytheridea obstipa, Cyamocytheridea dertonensis, Cytheretta semiornata and Nonurocythereis seminulum; (2) assemblage from the coral reef complexes within which Grinioneis haidingeri, Aurila cicatricosa, Cimbaurila diecii, Tenedocythere cruciata, Pokornyella italica and Callistocythere quadrangula are dominant and point to a stable inner-infralittoral environment characterised by warm, quiet and well-oxygenated waters; (3) assemblage from the silts intercalated among the coral reef complexes, mainly characterised by Neomonoceratina laskarevi, Cytheridea acuminata, Phlyctenophora farkasi and Aurila albicans together with Callistocythere spp., Xestoleberis communis and Xestoleberis dispar, which points to a very shallow marine environment rich in aquatic vegetation; (4) assemblage from the upper silts, which records the absolute dominance of Xestoleberis species, reflecting a very shallow and highly-vegetated environment and (5) assemblage from the uppermost silty clays, dominated by Hemicytherura defiorei, Xestoleberis spp. and Palmoconcha dertobrevis, accompanied by Acanthocythereis hystrix, Cytherella scutulum, Bairdoppilata conformis, Semicytherura spp., Krithe sp., Cytheropteron alatum, Bythocypris sp. and Pseudocythere caudata, which suggest deeper marine environments probably located in the outer infralittoral/inner-circalittoral zones. The studied section has been dated by means of calcareous nannoplankton to be not younger than Zone MNN9 (Early Tortonian), which is the biostratigraphical datum recorded in the fine-grained deposits that overlie the coral reef complexes. An age not older than Tortonian can be inferred by the stratigraphical distribution of the recognized ostracods. Thus, the coral reef complexes have been tentatively referred to the Early Tortonian.     

Fischreste (Selachii,Actinopterygii) aus der Wealdentonscholle von Lobber Ort (Mönchgut/Rügen/DDR)

A fossil fish fauna from an erratic Wealden clay deposit in the tillitic cliff of Lobber Ort (Isle of Rügen/GDR) is described (Hybodus sp.,Polyacrodus sp.,Lissodus rugianus n. sp.,Lepidotes cf.mantelli Agassiz,Lepidotes sp., Pycnodontiformes fam., gen. et sp. indet). The fauna indicates brackish-fresh water conditions and resembles those from British Purbeck and Wealden deposits and from the Wealden deposits of Hannover. The exceptional environmental features within the depositional area of the NW-European Wealden facies have obviously accelerated speciation processes within the genusLissodus Brough 1935 during the Earliest Cretaceous.