Andrea Engelbrecht

Ultimate Eocene (Priabonian) chondrichthyans (Holocephali, Elasmobranchii) of Antarctica

ABSTRACT The Eocene La Meseta Formation on Seymour Island, Antarctic Peninsula, is known for its remarkable wealth of fossil remains of chondrichthyans and teleosts. Chondrichthyans seemingly were dominant elements in the Antarctic Paleogene fish fauna, but decreased in abundance from middle to late Eocene, during which time remains of bony fishes increase. This decline of chondrichthyans at the end of the Eocene generally is related to sudden cooling of seawater, reduction in shelf area, and increasing shelf depth due to the onset of the Antarctic thermal isolation. The last chondrichthyan records known so far include a chimeroid tooth plate from TELM 6 (Lutetian) and a single pristiophorid rostral spine from TELM 7 (Priabonian). Here, we present new chondrichthyan records of Squalus, Squatina, Pristiophorus, Striatolamia, Palaeohypotodus, Carcharocles, and Ischyodus from the upper parts of TELM 7 (Priabonian), including the first record of Carcharocles sokolovi from Antarctica. This assemblage suggests that chondrichthyans persisted much longer in Antarctic waters despite rather cool sea surface temperatures of approximately 5°C. The final disappearance of chondrichthyans at the Eocene—Oligocene boundary concurs with abrupt ice sheet formation in Antarctica. Diversity patterns of chondrichthyans throughout the La Meseta Formation appear to be related to climatic conditions rather than plate tectonics.

Before the freeze: otoliths from the Eocene of Seymour Island, Antarctica, reveal dominance of gadiform fishes (Teleostei)

The first record of fossil teleostean otoliths from Antarctica is reported. The fossils were obtained from late Early Eocene shell beds of the La Meseta Formation, Seymour Island that represent the last temperate marine climate phase in Antarctica prior to the onset of cooling and subsequent glaciation during the late Eocene. A total of 17 otolith-based teleost taxa are recognized, with 10 being identifiable to species level containing nine new species and one new genus: Argentina antarctica sp. nov., Diaphus? marambionis sp. nov., Macruronus eastmani sp. nov., Coelorinchus balushkini sp. nov., Coelorinchus nordenskjoeldi sp. nov., Palimphemus seymourensis sp. nov., Hoplobrotula? antipoda sp. nov., Notoberyx cionei gen. et sp. nov. and Cepola anderssoni sp. nov. Macruronus eastmani sp. nov. is also known from the late Eocene of Southern Australia, and Tripterophycis immutatus Schwarzhans, widespread in the southern oceans during the Eocene, has been recorded from New Zealand, southern Australia, and now Antarctica. The otolith assemblage shows a typical composition of temperate fishes dominated by gadiforms, very similar at genus and family levels to associations known from middle Eocene strata of New Zealand and the late Eocene of southern Australia, but also to the temperate Northern Hemisphere associations from the Paleocene of Denmark. The Seymour Island fauna bridges a gap in the record of global temperate marine teleost faunas during the early Eocene climate maximum. The dominant gadiforms are interpreted as the main temperate faunal component, as in the Paleocene of Denmark. Here they are represented by the families Moridae, Merlucciidae (Macruroninae), Macrouridae and Gadidae. Nowadays Gadidae are a chiefly Northern Hemisphere temperate family. Moridae, Macruroninae and Macrouridae live today on the lower shelf to deep-water or mesopelagically with Macruroninae being restricted to the Southern Ocean. The extant endemic Antarctic gadiform family Muraenolepididae is missing, as are the dominant modern Antarctic fishes of the perciform suborder Notothenioidei. Recently, there has been much debate on isolated jaw bones of teleost fishes found in the La Meseta Formation and whether they would represent gadiforms (Merlucciidae in this case) or some early, primitive notothenioid. Otoliths are known to often complement rather than duplicate skeletal finds. With this in mind, we conclude that our otolith data support the presence of gadiforms in the early Eocene of Antarctica while it does not rule out the presence of notothenioids at the same time. http://zoobank.org/urn:lsid:zoobank.org:pub:A30E5364-0003-4467-B902-43A41AD456CC

Eocene sand tiger sharks (Lamniformes, Odontaspididae) from the Bolca Konservat-Lagerstätte, Italy: palaeobiology, palaeobiogeography and evolutionary significance

Abstract Here we report the first record of one of the most common and widespread Palaeogene selachians, the sand tiger shark Brachycarcharias, from the Ypresian Bolca Konservat-Lagerstätte. The combination of dental character of the 15 isolated teeth collected from the Pesciara and Monte Postale sites (e.g. anterior teeth up to 25 mm with fairly low triangular cusp decreasing regularly in width; one to two pairs of well-developed lateral cusplets; root with broadly separated lobes; upper teeth with a cusp bent distally) supports their assignment to the odontaspidid Brachycarcharias lerichei (Casier, 1946), a species widely spread across the North Hemisphere during the early Palaeogene. The unambiguous first report of this lamniform shark in the Eocene Bolca Konservat-Lagerstätte improves our knowledge concerning the diversity and palaeobiology of the cartilaginous fishes of this palaeontological site, and provides new insights about the biotic turnovers that involved the high trophic levels of the marine settings after the end-Cretaceous extinction.

A synoptic review of the Eocene (Ypresian) cartilaginous fishes (Chondrichthyes: Holocephali, Elasmobranchii) of the Bolca Konservat-Lagerstätte, Italy

Here, we review and discuss the records and taxonomy of the Ypresian (Eocene) chondrichthyans from the famous Bolca Konservat-Lagerstätte in northeastern Italy. Despite the outstanding diversity and the numerous studies focusing on the actinopterygian faunas from Pesciara and Monte Postale, the current knowledge about the systematics, taxonomy and phylogenetic relationships of the cartilaginous fishes from these Eocene sites remains elusive and largely inadequate. The celebrated Eocene Bolca Lagerstätte has yielded several exquisitely preserved articulated remains of chondrichthyan fishes in which delicate structures and soft tissues are preserved, as well as isolated teeth. The cartilaginous fish assemblage of Bolca comprises at least 17 species-level taxa belonging to 10 families in 6 orders, including selachians (Carcharhiniformes, Lamniformes), batoids (Torpediniformes, Myliobatiformes, Rajiformes) and holocephalans (Chimaeriformes). The occurrence of holocephalans represented by an isolated fin-spine of the chimeroid Ischyodus in the Bolca assemblage is reported here for the first time and represents the first record of chimeroids in the Eocene of Italy and also southern Europe. The Bolca chondrichthyan assemblage is remarkably different from those of other contemporaneous Boreal or Tethyan deposits, suggesting that its taxonomic composition is largely influenced by the palaeoenvironmental context. However, this synoptic review also highlights the importance of detailed revisions of all chondrichthyan remains from the Bolca Konservat-Lagerstätten.KurzfassungObwohl Knochenfische in der berühmten eozänen (Ypresium) Bolca-Konservatlagerstätte in NE-Italien äußerst divers sind und in den vergangenen Jahren Gegenstand zahlreicher Studien waren, ist der derzeitige Kenntnisstand über die Systematik, Taxonomie und Phylogenie der Knorpelfische aus dieser Lagerstätte überraschend gering und ungenau. In dieser Studie geben wir einen Überblick über die Knorpelfische der Bolca-Konservatlagerstätte und diskutieren ihre Verbreitung und ihre Taxonomie. Aus den beiden Fundstellen Pesciara und Monte Postale stammen einige sehr gut erhaltene Exemplare, bei denen auch sehr feine Strukturen und Weichgewebe fossilisiert sind, sowie isolierte Zähne. Die Knorpelfischfauna von Bolca umfasst mindestens 17 Arten, die zu zehn Familien und sechs Ordnungen gehören wie Haie (Carcharhiniformes, Lamniformes), Rochen (Torpediniformes, Myliobatiformes, Rajiformes) und Chimären. Chimären sind erstmals für Bolca durch einen isolierten Flossenstachel von Ischyodus belegt, was gleichzeitig der erste Nachweis für Chimären im Eozän von Italien und Süd-Europa ist. Die Knorpelfischassoziation von Bolca unterscheidet sich in ihrer taxonomischen Zusammensetzung auffällig von anderen tethyalen und auch borealen Faunen. Dies lässt vermuten, dass die taxonomische Zusammensetzung im Wesentlichen durch Umweltbedingungen bestimmt ist. Der hier präsentierte synoptische Überblick der Bolca-Knorpelfische zeigt aber auch deutlich, dass detaillierte Revisionen aller Knorpelfischreste der Bolca-Konservatlagerstätte für ein besseres Verständnis dieser wichtigen eozänen Fundstelle dringend nötig sind.

A new sawshark, Pristiophorus laevis, from the Eocene of Antarctica with comments on Pristiophorus lanceolatus

Abstract The highly fossiliferous Eocene deposits of the Antarctic Peninsula are among the most productive sites for fossil remains in the Southern Hemisphere and offer rare insights into high-latitude faunas during the Palaeogene. Chondrichthyans, which are represented by abundant isolated remains, seemingly dominate the marine assemblages. Eocene Antarctic sawsharks have only been known from few isolated rostral spines up to now, that were assigned to Pristiophorus lanceolatus. Here, we present the first oral teeth of a sawshark from the Eocene of Seymour Island and a re-evaluation of previously described Pristiophorus remains from Gondwana consisting exclusively of rostral spines. The holotype of Pristiophorus lanceolatus represents a single, abraded and insufficiently illustrated spine from the Oligocene of New Zealand. All other Cenozoic rostral spines assigned to this species are morphologically very indistinct and closely resemble those of living taxa. Consequently, we regard this species as dubious and introduce a new species, Pristiophorus laevis, based on oral teeth. The combination of dental characteristics of the new species makes it unique compared to all other described species based on oral teeth. Rostral spines from the Eocene of Seymour Island are assigned to this new species whereas those from other Cenozoic Gondwana localities remain ambiguous. LSID urn:lsid:zoobank.org:pub:7177A373-527B-4315-85F6-25180DB5E087

Revision of Eocene Antarctic carpet sharks (Elasmobranchii, Orectolobiformes) from Seymour Island, Antarctic Peninsula.

Seymour Island, Antarctic Peninsula, was once called the ‘Rosetta Stone’ of Southern Hemisphere palaeobiology, because this small island provides the most complete and richly fossiliferous Palaeogene sequence in Antarctica. Among fossil marine vertebrate remains, chondrichthyans seemingly were dominant elements in the Eocene Antarctic fish fauna. The fossiliferous sediments on Seymour Island are from the La Meseta Formation, which was originally divided into seven stratigraphical levels, TELMs 1–7 (acronym for Tertiary Eocene La Meseta) ranging from the upper Ypresian (early Eocene) to the late Priabonian (late Eocene). Bulk sampling of unconsolidated sediments from TELMs 5 and 6, which are Ypresian (early Eocene) and Lutetian (middle Eocene) in age, respectively, yielded very rich and diverse chondrichthyan assemblages including over 40 teeth of carpet sharks representing two new taxa, Notoramphoscyllium woodwardi gen. et sp. nov. and Ceolometlaouia pannucae gen. et sp. nov. Two additional teeth from TELM 5 represent two different taxa that cannot be assigned to any specific taxon and thus are left in open nomenclature. The new material not only increases the diversity of Eocene Antarctic selachian faunas but also allows two previous orectolobiform records to be re-evaluated. Accordingly, Stegostoma cf. faciatum is synonymized with Notoramphoscyllium woodwardi gen. et sp. nov., whereas Pseudoginglymostoma cf. brevicaudatum represents a nomen dubium. The two new taxa, and probably the additional two unidentified taxa, are interpreted as permanent residents, which most likely were endemic to Antarctic waters during the Eocene and adapted to shallow and estuarine environments. http://zoobank.org/urn:lsid:zoobank.org:pub:DD064382-3802-41A9-9B89-3095A6533B8A

THE SOUTHERNMOST OCCURRENCE OF BRACHYCARCHARIAS (LAMNIFORMES, ODONTASPIDIDAE) FROM THE EOCENE OF ANTARCTICA PROVIDES NEW INFORMATION ABOUT THE PALEOBIOGEOGRAPHY AND PALEOBIOLOGY OF PALEOGENE SAND TIGER SHARKS

The first record of one of the most common and widespread Paleogene selachians, the sand tiger shark Brachycarcharias , in the Ypresian strata of the La Meseta Formation, Seymour Island, Antarctica, is provided herein. Selachians from the early Eocene horizons of this deposit represent the southernmost Paleogene occurrences in the fossil record, and are represented by isolated teeth belonging to orectolobiforms, lamniforms, carcharhiniforms, squatiniforms and pristiophoriforms. The combination of dental characters of the 49 isolated teeth collected from the horizons TELMs 2, 4 and 5 supports their assignment to the odontaspidid Brachycarcharias lerichei (Casier, 1946), a lamniform species widely spread across the Northern Hemisphere during the early Paleogene. The unambiguous first report of this lamniform shark in the Southern Hemisphere in the Eocene of the La Meseta Formation improves our knowledge concerning the diversity and paleobiology of the cartilaginous fishes of this deposit, and provides new insights about the biotic turnovers that involved the high trophic levels of the marine settings after the end-Cretaceous extinction and before the establishment of the modern marine ecosystems.

Skates and rays (Elasmobranchii, Batomorphii) from the Eocene La Meseta and Submeseta formations, Seymour Island, Antarctica

Abstract Eocene deposits of the famous La Meseta Formation of Seymour Island, Antarctic Peninsula, yielded the most diverse Paleogene fossil elasmobranch association of the Southern Hemisphere. In this assemblage, sharks clearly dominate the fauna, whereas batoids are very rare components. Herein, we describe two new taxa of cold water tolerant skates, Marambioraja leiostemma gen. et sp. nov., and Mesetaraja maleficapelli gen. et sp. nov., two new species of the genus Raja, Raja amphitrita sp. nov. and Raja manitaria sp. nov., as well as remains of warm water adapted myliobatiforms. It is, however, not possible to unambiguously assign these remains either to Myliobatidae or Rhinopteridae, or to any specific genus. Previously reported remains of Raja/Bathyraja sp. are assigned to the new described species Raja manitaria sp. nov. The biogeographic distribution of extant and extinct rays and skates clearly shows that both groups are more widely distributed today than in the past, and additionally seem to have been more diverse in the Northern than the Southern Hemisphere. The occurrence, albeit rare of isolated teeth of skates (Rajidae) and rays (Myliobatidae) in the La Meseta Formation representes a minimum age constraint for their first appearance in the Southern Ocean. http://www.zoobank.org/urn:lsid:zoobank.org:pub:E507D86C-FFEC-4047-A337-AE77606BB1A6

New carcharhiniform sharks (Chondrichthyes, Elasmobranchii) from the early to middle Eocene of Seymour Island, Antarctic Peninsula

ABSTRACT Seymour Island, Antarctic Peninsula, is known for its wealth of fossil remains. This island provides one of the richest fossiliferous Paleogene sequences in the world. Chondrichthyans seemingly dominate this Eocene marine fauna and offer a rare insight into high-latitude faunas during the Palaeogene. So far, only a few isolated teeth of carcharhinid sharks have been reported from Seymour Island. Bulk sampling in the well-exposed La Meseta and Submeseta formations yielded new and abundant chondrichthyan material, including numerous teeth of carcharhinid and triakid sharks. Here, we present a reevaluation of the previously described carcharhinid remains and a description of new taxa: Meridiogaleus cristatus, gen. et sp. nov., Kallodentis rythistemma, gen. et sp. nov., Abdounia richteri, sp. nov., and Abdounia mesetae, sp. nov. The carcharhiniforms Mustelus sp. and Galeorhinus sp. are reported based on rare material, whereas teeth previously assigned to Scoliodon represent a nomen dubium.

Eocene squalomorph sharks (Chondrichthyes, Elasmobranchii) from Antarctica

Rare remains of predominantly deep-water sharks of the families Hexanchidae, Squalidae, Dalatiidae, Centrophoridae, and Squatinidae are described from the Eocene La Meseta Formation, Seymour Island, Antarctic Peninsula, which has yielded the most abundant chondrichthyan assemblage from the Southern Hemisphere to date. Previously described representatives of Hexanchus sp., Squalus weltoni, Squalus woodburnei, Centrophorus sp., and Squatina sp. are confirmed and dental variations are documented. Although the teeth of Squatina sp. differ from other Palaeogene squatinid species, we refrain from introducing a new species. A new dalatiid taxon, Eodalatias austrinalis gen. et sp. nov. is described. This new material not only increases the diversity of Eocene Antarctic elasmobranchs but also allows assuming that favourable deep-water habitats were available in the Eocene Antarctic Ocean off Antarctica in the Eocene. The occurrences of deep-water inhabitants in shallow, near-coastal waters of the Antarctic Peninsula agrees well with extant distribution patterns.