A 104-Ma record of deep-sea Atelostomata (Holasterioda, Spatangoida, irregular echinoids) - a story of persistence, food availability and a big bang.

Deep-sea macrobenthic body fossils are scarce due to the lack of deep-sea sedimentary archives in onshore settings. Therefore, hypothesized migrations of shallow shelf taxa into the deep-sea after phases of mass extinction (onshore-offshore pattern in the literature) due to anoxic events is not constrained by the fossil record. To resolve this conundrum, we investigated 1,475 deep-sea sediment samples from the Atlantic, Pacific and Southern oceans (water depth ranging from 200 to 4,700 m), providing 41,460 spine fragments of the crown group Atelostomata (Holasteroida, Spatangoida). We show that the scarce fossil record of deep-sea echinoids is in fact a methodological artefact because it is limited by the almost exclusive use of onshore fossil archives. Our data advocate for a continuous record of deep-sea Atelostomata back to at least 104 Ma (late early Cretaceous), and literature records suggest even an older age (115 Ma). A gradual increase of different spine tip morphologies from the Albian to the Maastrichtian is observed. A subsequent, abrupt reduction in spine size and the loss of morphological inventory in the lowermost Paleogene is interpreted to be an expression of the "Lilliput Effect", related to nourishment depletion on the sea floor in the course of the Cretaceous-Paleogene (K-Pg) Boundary Event. The recovery from this event lasted at least 5 Ma, and post-K-Pg Boundary Event assemblages progress-without any further morphological breaks-towards the assemblages observed in modern deep-sea environments. Because atelostomate spine morphology is often species-specific, the variations in spine tip morphology trough time would indicate species changes taking place in the deep-sea. This observation is, therefore, interpreted to result from in-situ evolution in the deep-sea and not from onshore-offshore migrations. The calculation of the "atelostomate spine accumulation rate" (ASAR) reveals low values in pre-Campanian times, possibly related to high remineralization rates of organic matter in the water column in the course of the mid-Cretaceous Thermal Maximum and its aftermath. A Maastrichtian cooling pulse marks the irreversible onset of fluctuating but generally higher atelostomate biomass that continues throughout the Cenozoic.

Njoerdichthys dyckerhoffi gen. et sp. nov. (Pycnodontiformes, lower Turonian) northward migration caused by the Cretaceous Thermal Maximum.

A new pycnodont taxon, Njoerdichthys dyckerhoffi gen. et sp. nov., from the Turonian of the Lower Saxony Basin of Germany is described and its systematic positions is established based on phylogenetic analyses of three specimens using slightly altered data matrices. All analyses display some differences to previous analyses but show very similar results to each other apart from the interpretation of the position of several taxa such as, e.g., Palaeobalistum. The new pycnodontiform specimens from northwestern Germany are unambiguously identified as a derived member of Pycnodontidae with close relationships to Abdobalistum and Nursallia? goedeli because of the unique combination of characters. One of the three specimens represents a juvenile form. Its morphological characters are limited, but it shares some characters with Njoerdichthys dyckerhoffi gen. et sp. nov. and is consequently allocated to the new taxon. The systematic placement of the new taxon, Njoerdichthys gen. nov., within Pycnodontidae is surprising since it does not display the one autapomorphic character (postparietal brush-like extension for muscle attachment) previously proposed to define this clade, but rather displays a combination of derived and homoplastic characters indicating that the definition of supra-generic taxa needs to be re-evaluated in the future by including more and new taxa. The distribution of pycnodontiform fishes in the Cretaceous appears to concur with changes in global climatic conditions, where high upper-ocean temperatures and high sea levels allow these fishes to migrate into higher latitudes as evidenced by the occurrence of the new taxon and Anomoeodus subclavatus in the Campanian of Sweden.