The coquina on the banks of the Oued Beth in the Rharb Basin in northwestern Morocco has long been known to be exceptionally rich in fossils. The stratigraphic position ranging from the Late Miocene ...to the Pliocene has been controversial, however. In the course of my master’s degree field work in 1975/76, I mapped the right bank of the Oued Beth from Dar bel Hamri to El Kansera. Following multiple recent studies in the general region, I here review my results and present an updated comprehensive stratigraphic and geologic frame for the first time. The coquina near Dar bel Hamri is interpreted to be of Early Pliocene age, possibly containing some reworking of Late Miocene fossils. The coquina and other locations along the Oued Beth have yielded a rich otolith assemblage, which is described in this article. It represents the first fossil otolith-based fish fauna described from Northwest Africa and contains 96 species, 16 of which are new. The new species in the order of their description are
Diaphus maghrebensis
n. sp.,
Ophidion tuseti
n. sp.,
Centroberyx vonderhochti
n. sp.,
Myripristis ouarredi
n. sp.,
Deltentosteus planus
n. sp.,
Caranx rharbensis
n. sp.,
Trachurus insectus
n. sp.,
Parapristipoma bethensis
n. sp.,
Pomadasys zemmourensis
n. sp.,
Cepola lombartei
n. sp.,
Trachinus maroccanus
n. sp.,
Trachinus wernlii
n. sp.,
Uranoscopus hoedemakersi
n. sp.,
Uranoscopus vanhinsberghi
n. sp.,
Spondyliosoma tingitana
n. sp., and
Opsodentex mordax
n. sp. In addition, a new species is described from the Tortonian and Zanclean of Italy:
Rhynchoconger carnevalei
n. sp. Some additional otoliths are described from another Northwest Moroccan location of Early Pliocene age near Asilah, 50 km south of the Strait of Gibraltar. The Early Pliocene fish fauna from Dar bel Hamri in the Rharb Basin is also of interest, because it constitutes the nearest Atlantic fauna of the time of the reconnection of the Mediterranean with the Atlantic and may have acted as a hosting area for the remigration of fishes into the Mediterranean. Indeed, the correlation is high between the Northwest Moroccan and the well-known time-equivalent Mediterranean fish fauna, but the Moroccan fauna also contains a good proportion of putative endemic taxa and taxa with tropical West African affinities that apparently did not migrate into the Mediterranean. Thus, the Early Pliocene fish fauna from the Rharb Basin represents a unique assemblage for which I propose the biogeographic term “Maghrebian bioprovince.”
Almost fifty years after the first definition of the Messinian salinity crisis (MSC), the events that occurred in the Mediterranean during the terminal portion of the Miocene still attract the ...attention of a large and diverse scientific community. Although fossils are relatively common in the deposits that accumulated during the MSC, their significance for the interpretation of the latest Miocene paleoenvironmental evolution of the Mediterranean has been underevalued. In this paper, we summarize the marine paleoichthyological record of the three stages of the MSC based on both articulated and isolated skeletal remains and otoliths, the latter almost exclusively known from the Lago-Mare phase. We focus on the composition of the marine ichthyofauna of the Mediterranean during the three main stages of the MSC, showing the persistent continuity of marine stenohaline taxa throughout most of the interval between 5.97 and 5.33 Ma. While the record of articulated fish skeletons is unquestionably autochthonous, thereby providing unambiguous evidence of the occurrence of open marine environments in the MSC preceding the Lago-Mare phase, the autochthonous nature of the otolith record has often been questioned. For this reason, the otolith record of marine fishes has been examined in detail from a taxonomic and paleoecological point of view. Three species, Bellottia verecunda n. sp., Benthosema taurinense n. sp., and Bostrychus marsilii n. sp., are described as new and a thorough discussion about the possible origin of the otoliths is provided. Alternative explanations for the occurrence of otoliths of marine fish during the Lago-Mare phase, such as reworking, contamination from overlying Pliocene sediments or import from outside the Mediterranean through aquatic birds are considered unlikely. In our assessment, the occurrence of marine fish otoliths in the Lago-Mare phase can be explained with the presence of normal marine environments in the Mediterranean, at least temporarily. Therefore, we suggest that the paleoichthyological data provided herein should be integrated in the future evolutionary paleoenvironmental reconstructions of the MSC.
Otolith assemblages have rarely been studied across the K-Pg boundary. The late Maastrichtian Kemp Clay of northeastern Texas and the Fox Hills Formation of North Dakota, and the early Danian Clayton ...Formation of Arkansas therefore offer new insights into how teleost fishes managed across the K-Pg boundary as reconstructed from their otoliths. The Kemp Clay contains 25 species, with 6 new species and 2 in open nomenclature and the Fox Hills Formation contains 4 species including 1 new species. The two otolith associations constitute the Western Interior Seaway (WIS) community. It contains the earliest unambiguous representatives of the Gadiformes (cods and hakes) and the Heterenchelyidae (mud eels). The WIS community differs significantly from other Maastrichtian otolith assemblages previously studied from Mississippi and Maryland, which constitute the Appalachian community, with only 4 shared species (similarity percentage of 7.3%) between both communities. The difference is interpreted to be related to cold-water influence in the WIS community, which may have still been connected to the Arctic Basin, and to the depostional environment (muddy bottom) in the Kemp Clay. The Kemp Clay is unusually rich in taxa that survived the end-Cretaceous extinction event and are still present in the Danian of the Clayton Formation, or, as the case may be, in the Danian and Selandian of the boreal northern European community known from Denmark. Approximately 54% of all otolith-based teleost species identified from the Maastrichtian WIS community survived the K-Pg boundary event (versus 11-12% in other communities) and 73% of the genera (versus 40-50% in other communities). The early Danian Clayton Formation contains an impoverished inherited association with 14 species, of which 11 are survivors from late Maastrichtian times, 1 species is new, and 2 remain in open nomenclature. This compares to a significantly higher degree of newly evolved species in only slightly younger faunas from the middle to late Danian and Selandian of Europe indicating an initially slow pace of recovery. The observed differences in survival and the composition of survived and extinct taxa are discussed in the light of the ongoing discussions concerning the consequences and effects that led to the end-Cretaceous extinction event commonly thought to have been caused by a large meteorite impact. In our assessment, an ‘impact winter’ could have had a major influence on the live cycle of tropical to subtropical fishes while perturbations in the pelagic food web or ocean surface acidification might have had a minor and more selective effect. Overall, teleost fishes were significantly affected by the end-Cretaceous mass extinction, but to a much lesser extent than in many other biota. This study provides more evidence of the importance of Late Cretaceous otolith assemblages in the USA for interpreting teleostean evolution. The newly described taxa are: Elopothrissus carsonsloani n. sp., Pythonichthys arkansasensis n. sp., Congrophichthus transterminus n. gen., n. sp., Rhynchoconger brettwoodwardi n. sp., Palaeogadus weltoni n. sp., Dakotaichthys hogansoni n. gen., n. sp., and Ampheristus americanus n. sp.
A small collection of fish otoliths, mostly myctophids, from a bathyal environment are described from the Bartonian (late middle Eocene) Marne di Monte Piano Formation of San Genesio, Turin Hill, ...Piedmont. Myctophids have been rarely recorded from the Eocene in general, and few bathyal environments have been sampled for otoliths of that time interval globally. In this context, the assemblage described herein fills an important stratigraphic gap in the record and evolution of the Myctophidae. The findings are discussed in a regional and stratigraphic context. Seven species are recorded, of which one is new, Oligophus bartonensis n. sp., belonging to the family Myctophidae.
Otoliths are common and diverse in the Neogene of tropical America. Following previous studies of Neogene tropical American otoliths of the lanternfishes (Myctophidae), marine catfishes (Ariidae), ...croakers (Sciaenidae), and cusk-eels (Ophidiiformes), we describe here the otoliths of the gobies (Gobiidae). The Gobiidae represent the richest marine fish family, with more than 2000 species worldwide and about 250 in America. In the fossil record too they are the species richest family in the Neogene of tropical America. We have investigated otoliths sampled from Ecuador, Pacific and Atlantic Panama, Atlantic Costa Rica, Dominican Republic, Venezuela, and Trinidad, ranging in age from late Early Miocene (late Burdigalian) to late Early Pleistocene (Calabrian). Most of the studied material originates from the collection expeditions of the Panama Paleontology Project (PPP). Our study represents the first comprehensive record of fossil gobies from America, and we recognize 107 species, of which 51 are new to science, 35 are in open nomenclature, and 19 represent species that also live in the region today. Previously, only two fossil otolith-based goby species have been described from the Neogene of tropical America. The dominant gobies in the fossil record of the region are from the Gobiosomatini, particularly of genera living over soft bottoms or in deeper water such as
Bollmannia
,
Microgobius
,
Antilligobius
, and
Palatogobius
. Another purpose of our study is to provide a first comprehensive account of otoliths of the extant Gobiidae of America, which we consider necessary for an adequate identification and interpretation of the Neogene otoliths. We studied otoliths of 130 extant American gobiid species and figured 106 of them for comparison. We also present a morphological analysis and characterization of the extant otoliths as a basis for the identification of fossil otoliths. Problems that commonly arise with the identification of fossil otoliths and specifically of fossil goby otoliths are addressed and discussed. A comparison of the history of the Gobiidae in tropical America reveals a high percentage of shared species between the Pacific and the Atlantic basins during the Late Miocene (Tortonian and Messinian) from at least 11 to 6 Ma. A recording gap on the Pacific side across the Pliocene allows a comparison again only in the late Early Pleistocene (Calabrian, 1.8 to 0.78 Ma), which shows a complete lack of shared species. These observations support the effective closure of the former Central American Seaway and emersion of the Isthmus of Panama in the intervening time. Groups that today only exist in the East Pacific were also identified in the Miocene and Pliocene of the West Atlantic, and there is also at least one instance of a genus now restricted to the West Atlantic having occurred in the East Pacific as late as the Pleistocene. The evolution of gobies in tropical America and the implications thereof are extensively discussed. Furthermore, observations of fossil gobies in the region are discussed in respect to paleoenvironmental indications and paleobiogeographic aspects.
The first fossil otolith association from the middle Paleocene (Selandian) of Scania, southern Sweden is described. Forty-seven otoliths were retrieved from shallow wells representing 14 teleost ...taxa. Many specimens are small and/or eroded and, therefore, not identifiable to species level. Nevertheless, our findings indicate the potential for further fossil otolith discoveries in the region. The Scanian otolith-based fauna greatly resembles the better-known coeval association from Copenhagen, Denmark, but is relatively rich and diverse in perciform otoliths. The fauna records the first occurrence of Serranus? caribbaeus from the European Paleocene, and of Archaemacruroides ornatus from the Selandian of the North Sea Basin.
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The Lago Mare phase at the end of the Messinian Salinity Crisis (MSC) in the Mediterranean has long been known for the Paratethyan affinities of its biota. The taxonomic level of these affinities and ...their origin, however, is subject to divergent interpretations. Here, we have studied otoliths of the Gobioidei from late Miocene and early Pliocene deposits from Italy and Greece and revised earlier studies including time-equivalent data from Romania and NW Turkey. Nowadays, gobies constitute the most speciose marine fish family in Europe and the most diverse endemic family in the Ponto-Caspian region. Furthermore, they are highly adapted to nearshore shallow marine, brackish, and freshwater environments, which makes them excellent candidates to explore short-term connections of waterways of different nature. The normal marine late Tortonian and pre-evaporitic Messinian deposits of the studied Italian and Greek locations contained a rich and diverse goby assamblage, but species with Paratethyan affinities were very rare and possibly originated from connections during the early Serravallian (late Badenian) and early Messinian (early Meotian). A rare fauna from a paralic environment from Cessaniti, Calabria, similar to the mangrove environments found in the present-day Guinea coastal area, revealed two species of the family Eleotridae and no gobies with Paratethyan affinities. Cessaniti provides a unique opportunity to recognize the presence of this type of environmental context with a unique gobioid assemblage in the Neogene of Europe. The shallow-water brackish-influenced sediments of Strada degli Archi just below the MSC show an increasing influence of Paratethyan gobiid taxa (Proterorhinus yigitbasi, Zosterisessor exsul n. sp.), which are not present in time-equivalent normal marine environments, but have also been identified in the Dacic Basin of Romania. Sediments of the stage 3 ot the MSC, especially those that accumulated during the Lago Mare event, were dominated by a diverse goby assemblage with Paratethyan affinities, while indigenous Mediterranean goby taxa disappeared probably because of the lack of suitable conditions for neritic demersal stenohaline fishes, except for the nektonic Aphia minuta. The sediments of the stage 3 of the MSC also included in one particular location and level the unusual Enigmacottus socialis n. gen., n. sp., a putative member of the family Psychrolutidae of uncertain origin, which constitutes about 70% of all otoliths obtained from that interval in Italy. Beginning with the Zanclean, the goby assemblage of the Mediterranean shows an almost exclusively Atlantic-Mediterranean composition. A total of 15 new species are described, 12 in the family Gobiidae, two of the Eleotridae, and one putative Psychrolutidae. The new taxa are: Eleotris omuamuaensis n. sp., Eleotris tyrrhenicus n. sp., Lesueurigobius stazzanensis n. sp., Gobius peloponnesus n. sp., Proterorhinus cretensis n. sp., Zosterisessor exsul n. sp., Benthophilus aprutinus n. sp., Benthophilus labronicus n. sp., Caspiosoma lini n. sp., Caspiosoma paulisulcata n. sp., Thorogobius petilus n. sp., Buenia pisiformis n. sp., Hesperichthys gironeae n. sp., Knipowitschia etrusca n. sp., and Enigmacottus socialis n. gen. et n. sp.
Reconstructing fossil bony fish faunas using otoliths is a well-established method that allows a diverse and dense record in time and space to be assembled. Here we report about a rich otolith-based ...fish fauna from the middle Sarmatian s.l. (middle Bessarabian) from Jurkine, Kerch Peninsula, Crimea. The study is based on more than 5,000 specimens constituting 36 different species, 24 of which are new and 2 remain in open nomenclature. This assemblage represents the first major otolith association described from the Bessarabian. It also represents a fish fauna from the last continuous restricted marine environment that evolved in the Eastern Paratethys, was recruited from the Badenian/Tarkhanian fauna, and was not affected by the subsequent Khersonian crisis. The association of otoliths is characterized by a high content of endemic fishes that derived from the relatively well-known early Sarmatian s.l. (Volhynian) fish fauna, and it contains certain faunal elements that were trapped in the then-secluded Eastern Paratethys and did not range into younger strata. This forced endemic evolution explains the unusually high percentage of new taxa.
The fish fauna is dominated by stenohaline marine shelf fishes apparently recruited from the Konkian and earlier Sarmatian s.l. (Volhynian) fauna after the Karaganian crisis. The families Gobiidae and Gadidae benefited most in this restricted marine environment, while deep-water fishes disappeared with the Karaganian crisis. In this study, we discuss the further evolution of Eastern Paratethyan fishes as far as can be reconstructed from the relatively limited data from post-Bessarabian strata, and we also outline targets for future research in the field.
The stratigraphic sequence of the Jurkine section is being revised based on a detailed suite of benthic foraminifera. Implications for the stratigraphy of the middle and upper Sarmatian s.l., their boundary, and the paleoenvironments of this part of the Kerch Peninsula are discussed.
Myctophid otoliths are the most common fossil otoliths to be found in Neogene deep water sediments below approximately 200 m depositional depth. The southeastern part of Japan is particularly rich in ...such locations. Here we describe myctophid otoliths from late Pliocene to early Pleistocene locations on Okinawa, Shikoku and central Honshu and review previous publications. These faunas represent the tropical to subtropical faunal zone, which was established as the “Kakegawa Fauna” based on molluscs. A total of 37 species are recognized by means of otoliths in the family Myctophidae and one in the family Neoscopelidae. Two species are recorded in open nomenclature, 23 as persistent extant species and 12 extinct species, thereof 7 new. The new species are: Diaphus caurus, Diaphus endoi, Diaphus kakegawaensis, Diaphus nafpaktitisi, Diaphus noboriensis, Symbolophorus moriguchii and Notoscopelus praejaponicus. Two very large myctophid otoliths (>8 mm length) from the putative Piacenzian of Fiji are also described, one being a new species - Diaphus grebneffi - representing the largest by far otolith associated with the So-group of species in the genus so far known. The myctophid otolith assemblage is characterized by an abundance of large specimens (>5 mm length) which certainly derived from fully adult fishes and represents the biggest association of its kind described so far from the fossil record of the northern Pacific. Its composition is interpreted for stratigraphic and environmental purposes. The occurrence of a few, rare species with links to coeval finds in the Caribbean exemplify the potential of myctophid otoliths for supraregional biostratigraphic purposes. The comparison of the fossil assemblage with the distribution in the Recent supports the concept of a warm “Kakegawa Fauna” as established by molluscs and at the same time documents that certain tropical species reached further north during the late Pliocene than they do today.
Large and diverse Eocene otolith assemblages have been described from the US Gulf and Atlantic coastal plains, various basins in Europe, and New Zealand. Here we described a highly diverse otolith ...association from the middle Eocene (Lutetian and Bartonian) strata of southern California, San Diego County, which was retrieved from the heritage of John E. Fitch (1918-1982) in the archives of the Ichthyology Department of the Natural History Museum of Los Angeles County, Los Angeles, Califronia. The collection represents the first Eocene otolith assemblage described from the Northeastern Pacific and stems from two localities: the Ardath Shale at Black’s Beach in the Torrey Pines Park just south of Black’s Canyon Road and north of the Salk Fault on the Pacific shore face, and the MissionValley Formation from the west-facing graded hill just east of the Miramar Water Treatment Plant near the southern shore of Miramar Reservoir. The Ardath Shale was deposited on the deep shelf and continental slope while the Mission Valley Formation represent an inland, nearshore environment during times of the maximum transgression. A total of more than15,000 otoliths were found representing 96 species of which 53 are described as new and 12 remain in open nomenclature. The fauna of the Ardath Shale is distinctly richer containing 88 species while the one from the Miramar Reservoir site yielded 38 species. The low diversity at the Miramar Reservoir site is thought to be related to the shallow-water environment in which it was deposited whereas the high diversity at Black’s Beach reflects a mixture of shelf and bathyal fishes. In addition, 24 new otolith-based genera are established, four of which are in the category “incertae sedis”. Furthermore, five new species are described from a comparative otolith collection from the US Gulf Coast Eocene and one new species from a comparative collection from the Lutetian of Balegem in Belgium. The Eocene otolith-based fish fauna from California is assessed for its paleoecological, paleobiogeographic and evolutionary significance. Particular emphasis is directed towards the elucidation of the evolution of the early Ophidiiformes, which during the early Paleogene were predominantly adapted to warm, clastic shallow-water environments and there were competing in abundance and diversity with other percomorphs (e.g., perciforms s.l.). A warm Eocene paleobioprovince is proposed for the Californian fish fauna which is not related to today’s endemic Northwest Pacific fish fauna along the regime of the cold-water California Current.
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