During the
Middle Miocene Climatic Optimum, the Paratethys supported a tropical to subtropical marine ecosystem with very high biodiversity and
endemism, including the establishment of
coral reef ecosystems. Some portions of the Paratethys, over modern Poland, were deep enough to support a
bathypelagic ecosystem with significant endemism. An exceptionally-preserved record of this ecosystem is known from the
Oligocene-aged
Menilite Formation, a
flysch containing fossils of pelagic and deep-sea fish taxa, as well as
microbial mats. This biodiversity was badly hit by environmental changes later in the Miocene, with the coral reefs being wiped out following cooling during the
Middle Miocene disruption, while changing circulation patterns and the resulting
anoxia wiped out the deepwater habitats. Surface-dwelling species saw a significant decline from a collapse in the
zooplankton populations. Despite its diversity, the brief "Paratethyan biodiversity hotspot" was short-lived, lasting for only 3 million years. These endemic fish were closely related to modern groups, but belonged to their own distinct, now-lost evolutionary radiations. For example,
gobies were a particularly successful group in the Paratethys, with numerous endemic genera and species known from both fossil skeletons and
otoliths. The modern diversity of Ponto-Caspian gobies (
Benthophilinae) likely originates from survivors of this radiation. Many different families of
clupeoids (
herrings,
shad and allies) also saw extensive diversification in the Paratethys, with many fossil genera known. Connections with the Mediterranean Sea allowed for many
cartilaginous fish (sharks and rays) to colonize the Paratethys Sea by the Early Miocene. A Miocene-aged deepwater shark fauna from Slovakia is depauperate & mostly dominated by
squaliforms, and appears to suggest a highly stressed paleoenvironment. Later fossil assemblages suggest that the increased isolation of the Paratethys by the Middle Miocene caused significant extirpation among most small-to-medium sized deepwater and pelagic sharks. However, larger sharks, such as the
megalodon and
Cosmopolitodus, continued to persist in the Paratethys and did not see such extirpations, likely due to the widespread occurrence of marine mammals to feed on.
Marine mammals The Paratethys also supported numerous marine mammal lineages, including
cetaceans and
pinnipeds. It included several genera of dwarf baleen whales within the family
Cetotheriidae, including
Cetotherium rathkii and
Ciuciulea davidi. These are among the smallest baleen whales ever known to have existed, and it has been suggested that the group may have originated in the Paratethys. The
eurhinodelphinids, an unusual family of
toothed whales, appear to have invaded the Paratethys via the Mediterranean during the middle Miocene, with remains of the widespread genus
Xiphiacetus recovered from Austria. The more primitive toothed whale
Romaleodelphis also appears to have been a Paratethyan endemic. Over time, in response to the increased salinity in the Paratethys from its isolation, marine mammals independently evolved
pachyosteosclerosis, leading to dense, bulky bones. This condition appears to have independently evolved in pinnipeds, toothed whales & baleen whales, and first started appearing the Central Paratethys after the
Badenian Salinity Crisis before spreading eastwards. ==See also==