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The Paratethys sea, Paratethys ocean, Paratethys realm or just Paratethys (meaning "beside Tethys"), was a large shallow inland sea that covered much of mainland Europe and parts of western Asia during the middle to late
Cenozoic The Cenozoic Era ( ; ) is Earth's current geological era, representing the last 66million years of Earth's history. It is characterized by the dominance of mammals, insects, birds and angiosperms (flowering plants). It is the latest of three g ...
, from the late
Paleogene The Paleogene Period ( ; also spelled Palaeogene or Palæogene) is a geologic period and system that spans 43 million years from the end of the Cretaceous Period Ma (million years ago) to the beginning of the Neogene Period Ma. It is the fir ...
to the late
Neogene The Neogene ( ,) is a geologic period and system that spans 20.45 million years from the end of the Paleogene Period million years ago ( Mya) to the beginning of the present Quaternary Period million years ago. It is the second period of th ...
, and is regarded as the largest inland sea in history. At its greatest extent, it stretched from the region north of the
Alps The Alps () are some of the highest and most extensive mountain ranges in Europe, stretching approximately across eight Alpine countries (from west to east): Monaco, France, Switzerland, Italy, Liechtenstein, Germany, Austria and Slovenia. ...
over
Central Europe Central Europe is a geographical region of Europe between Eastern Europe, Eastern, Southern Europe, Southern, Western Europe, Western and Northern Europe, Northern Europe. Central Europe is known for its cultural diversity; however, countries in ...
to the
Aral Sea The Aral Sea () was an endorheic lake lying between Kazakhstan to its north and Uzbekistan to its south, which began shrinking in the 1960s and had largely dried up into desert by the 2010s. It was in the Aktobe and Kyzylorda regions of Kazakhst ...
in
Central Asia Central Asia is a region of Asia consisting of Kazakhstan, Kyrgyzstan, Tajikistan, Turkmenistan, and Uzbekistan. The countries as a group are also colloquially referred to as the "-stans" as all have names ending with the Persian language, Pers ...
. Paratethys formed about 34 Mya (million years ago) at the beginning of the
Oligocene The Oligocene ( ) is a geologic epoch (geology), epoch of the Paleogene Geologic time scale, Period that extends from about 33.9 million to 23 million years before the present ( to ). As with other older geologic periods, the rock beds that defin ...
epoch, when the northern region of the
Tethys Ocean The Tethys Ocean ( ; ), also called the Tethys Sea or the Neo-Tethys, was a prehistoric ocean during much of the Mesozoic Era and early-mid Cenozoic Era. It was the predecessor to the modern Indian Ocean, the Mediterranean Sea, and the Eurasia ...
(Peri-Tethys) was separated from the Mediterranean region of the Tethys realm due to the formation of the Alps,
Carpathians The Carpathian Mountains or Carpathians () are a range of mountains forming an arc across Central Europe and Southeast Europe. Roughly long, it is the third-longest European mountain range after the Urals at and the Scandinavian Mountains ...
, Dinarides, Taurus and Elburz mountains. Paratethys was at times reconnected with the Tethys or its successors (the
Mediterranean Sea The Mediterranean Sea ( ) is a sea connected to the Atlantic Ocean, surrounded by the Mediterranean basin and almost completely enclosed by land: on the east by the Levant in West Asia, on the north by Anatolia in West Asia and Southern Eur ...
or the
Indian Ocean The Indian Ocean is the third-largest of the world's five oceanic divisions, covering or approximately 20% of the water area of Earth#Surface, Earth's surface. It is bounded by Asia to the north, Africa to the west and Australia (continent), ...
) during the
Oligocene The Oligocene ( ) is a geologic epoch (geology), epoch of the Paleogene Geologic time scale, Period that extends from about 33.9 million to 23 million years before the present ( to ). As with other older geologic periods, the rock beds that defin ...
and the early and middle
Miocene The Miocene ( ) is the first epoch (geology), geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and mea ...
times, but at the onset of the late Miocene epoch, the tectonically trapped sea turned into a megalake from the eastern Alps to what is now
Kazakhstan Kazakhstan, officially the Republic of Kazakhstan, is a landlocked country primarily in Central Asia, with a European Kazakhstan, small portion in Eastern Europe. It borders Russia to the Kazakhstan–Russia border, north and west, China to th ...
. From the
Pliocene The Pliocene ( ; also Pleiocene) is the epoch (geology), epoch in the geologic time scale that extends from 5.33 to 2.58Black Sea The Black Sea is a marginal sea, marginal Mediterranean sea (oceanography), mediterranean sea lying between Europe and Asia, east of the Balkans, south of the East European Plain, west of the Caucasus, and north of Anatolia. It is bound ...
,
Caspian Sea The Caspian Sea is the world's largest inland body of water, described as the List of lakes by area, world's largest lake and usually referred to as a full-fledged sea. An endorheic basin, it lies between Europe and Asia: east of the Caucasus, ...
,
Aral Sea The Aral Sea () was an endorheic lake lying between Kazakhstan to its north and Uzbekistan to its south, which began shrinking in the 1960s and had largely dried up into desert by the 2010s. It was in the Aktobe and Kyzylorda regions of Kazakhst ...
,
Lake Urmia Lake Urmia is an endorheic salt lake in Iran. The lake is located between the provinces of East Azerbaijan and West Azerbaijan in Iran, and west of the southern portion of the Caspian Sea. At its greatest extent, it was the largest lake in th ...
, Namak Lake and others are remnants of the Paratethys Sea.


Name and research

The name Paratethys was first used by Vladimir Laskarev in 1924. Laskarev's definition included only
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserve ...
s and sedimentary strata from the sea of the
Neogene The Neogene ( ,) is a geologic period and system that spans 20.45 million years from the end of the Paleogene Period million years ago ( Mya) to the beginning of the present Quaternary Period million years ago. It is the second period of th ...
system A system is a group of interacting or interrelated elements that act according to a set of rules to form a unified whole. A system, surrounded and influenced by its open system (systems theory), environment, is described by its boundaries, str ...
. This definition was later adjusted also to include the Oligocene
series Series may refer to: People with the name * Caroline Series (born 1951), English mathematician, daughter of George Series * George Series (1920–1995), English physicist Arts, entertainment, and media Music * Series, the ordered sets used i ...
. The existence of a separate water body in these periods was deduced from the fossil fauna, including mollusks, fish and
ostracod Ostracods, or ostracodes, are a Class (biology), class of the crustacean, Crustacea (class Ostracoda), sometimes known as seed shrimp. Some 33,000 species (only 13,000 of which are extant taxon, extant) have been identified,Brandão, S.N.; Antoni ...
s. In periods in which the Paratethys or parts of it were separated from each other or from other oceans, a separate fauna developed which is found in sedimentary deposits. In this way, the paleogeographical development of the Paratethys can be studied. Laskerev's description of the Paratethys was anticipated much earlier by Sir Roderick Murchison in chapter 13 of his 1845 book. One of the key characteristics of the Paratethys realm, that is differentiating it from the
Tethys Ocean The Tethys Ocean ( ; ), also called the Tethys Sea or the Neo-Tethys, was a prehistoric ocean during much of the Mesozoic Era and early-mid Cenozoic Era. It was the predecessor to the modern Indian Ocean, the Mediterranean Sea, and the Eurasia ...
, is the widespread development of
endemic Endemism is the state of a species being found only in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also foun ...
fauna, adapted to fresh and brackish waters like those that still exist in recent waters of the Caspian Sea. This distinctive fauna in which univalves of freshwater origin such as ''Limnex'' and ''Neritinex'' are associated with forms of Cardiacae and Mytili, common to partially saline or brackish waters, makes the geologic records from Paratethys particularly difficult to correlate with those from other oceans or seas because their faunas evolved separately at times. Stratigraphers of the Paratethys, therefore, have their own sets of stratigraphic stages which are still used as alternatives for the official
geologic timescale The geologic time scale or geological time scale (GTS) is a representation of time based on the rock record of Earth. It is a system of chronological dating that uses chronostratigraphy (the process of relating strata to time) and geochron ...
of the ICS.


Palaeogeographic evolution

Paratethys was peculiar due to its paleogeography: it consisted of a series of deep basins, formed during the Oxfordian stage of the
Late Jurassic The Late Jurassic is the third Epoch (geology), epoch of the Jurassic Period, and it spans the geologic time scale, geologic time from 161.5 ± 1.0 to 143.1 ± 0.8 million years ago (Ma), which is preserved in Upper Jurassic stratum, strata.Owen ...
as an extension of the rift that formed the Central
Atlantic Ocean The Atlantic Ocean is the second largest of the world's five borders of the oceans, oceanic divisions, with an area of about . It covers approximately 17% of Earth#Surface, Earth's surface and about 24% of its water surface area. During the ...
. During the
Jurassic The Jurassic ( ) is a Geological period, geologic period and System (stratigraphy), stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately 143.1 Mya. ...
and
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 143.1 to 66 mya (unit), million years ago (Mya). It is the third and final period of the Mesozoic Era (geology), Era, as well as the longest. At around 77.1 million years, it is the ...
periods, this part of Eurasia was covered by shallow seas that formed the northern margins of the Tethys Ocean. These basins were connected with each other and the global ocean by narrow and shallow seaways that often limited water exchange and caused widespread long-term anoxia. The Paratethys descends directly from the Peri-Tethys, the northwestern arm of the Tethys Ocean, which was separated from the rest of the ocean via the Alpide orogeny. The Paratethys spread over a large area in Central Europe and western Asia. In the west it included in some stages the Molasse basin north of the
Alps The Alps () are some of the highest and most extensive mountain ranges in Europe, stretching approximately across eight Alpine countries (from west to east): Monaco, France, Switzerland, Italy, Liechtenstein, Germany, Austria and Slovenia. ...
; the Vienna Basin, the Outer Carpathian Basin, the
Pannonian Basin The Pannonian Basin, with the term Carpathian Basin being sometimes preferred in Hungarian literature, is a large sedimentary basin situated in southeastern Central Europe. After the Treaty of Trianon following World War I, the geomorpholog ...
, and further east to the basin of the current
Black Sea The Black Sea is a marginal sea, marginal Mediterranean sea (oceanography), mediterranean sea lying between Europe and Asia, east of the Balkans, south of the East European Plain, west of the Caucasus, and north of Anatolia. It is bound ...
and the
Caspian Sea The Caspian Sea is the world's largest inland body of water, described as the List of lakes by area, world's largest lake and usually referred to as a full-fledged sea. An endorheic basin, it lies between Europe and Asia: east of the Caucasus, ...
until the current position of the
Aral Sea The Aral Sea () was an endorheic lake lying between Kazakhstan to its north and Uzbekistan to its south, which began shrinking in the 1960s and had largely dried up into desert by the 2010s. It was in the Aktobe and Kyzylorda regions of Kazakhst ...
.


Anoxic Giant

The boundary between the
Eocene The Eocene ( ) is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (Ma). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene'' comes ...
and
Oligocene The Oligocene ( ) is a geologic epoch (geology), epoch of the Paleogene Geologic time scale, Period that extends from about 33.9 million to 23 million years before the present ( to ). As with other older geologic periods, the rock beds that defin ...
epochs was characterized by a big drop of the global ( eustatic) sea level and sudden steep cooling of global climates. At the same time the Alpine orogeny, a
tectonic Tectonics ( via Latin ) are the processes that result in the structure and properties of the Earth's crust and its evolution through time. The field of ''planetary tectonics'' extends the concept to other planets and moons. These processes ...
phase by which the Alps,
Carpathians The Carpathian Mountains or Carpathians () are a range of mountains forming an arc across Central Europe and Southeast Europe. Roughly long, it is the third-longest European mountain range after the Urals at and the Scandinavian Mountains ...
, Dinarides, Taurus, Elburz and many other
mountain chain A mountain chain is a row of high mountain summits, a linear sequence of interconnected or related mountains,Whittow, John (1984). ''Dictionary of Physical Geography''. London: Penguin, p 87. . or a contiguous ridge of mountains within a larger mo ...
s along the southern rim of Eurasia were formed. The combination of a drop in sea level and tectonic uplift resulted in the partial disconnection of the Tethys and Paratethys domains. Due to poor connectivity with the global ocean, the Paratethys realm became stratified and turned into a giant anoxic sea. The western and central Paratethys basins experienced intense tectonic activity and anoxia during the Oligocene and early
Miocene The Miocene ( ) is the first epoch (geology), geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and mea ...
and became filled with
sediments Sediment is a solid material that is transported to a new location where it is deposited. It occurs naturally and, through the processes of weathering and erosion, is broken down and subsequently sediment transport, transported by the action of ...
. Local
gypsum Gypsum is a soft sulfate mineral composed of calcium sulfate Hydrate, dihydrate, with the chemical formula . It is widely mined and is used as a fertilizer and as the main constituent in many forms of plaster, drywall and blackboard or sidewalk ...
and salt evaporitic basins formed in the East Carpathian region during the early Miocene. The Eastern Paratethys basin, holding most of the water of Paratethys, remained
anoxic Anoxia means a total depletion in the level of oxygen, an extreme form of hypoxia or "low oxygen". The terms anoxia and hypoxia are used in various contexts: * Anoxic waters, sea water, fresh water or groundwater that are depleted of dissolved ox ...
for almost 20 million years (35–15 Mya), and during this time Paratethys acted as an enormous
carbon sink A carbon sink is a natural or artificial carbon sequestration process that "removes a  greenhouse gas, an aerosol or a precursor of a greenhouse gas from the atmosphere". These sinks form an important part of the natural carbon cycle. An overar ...
trapping organic matter in its sediments. The Paratethys anoxia was "shut down" during the middle Miocene, some 15 million years ago, when a widespread
marine transgression A marine transgression is a geologic event where sea level rises relative to the land and the shoreline moves toward higher ground, resulting in flooding. Transgressions can be caused by the land sinking or by the ocean basins filling with water ...
, known as the Badenian Flooding, improved connections with the global ocean and triggered the ventilation of the deep waters of Paratethys.


Short-lived open seas

After the Badenian Flooding, in the middle
Miocene The Miocene ( ) is the first epoch (geology), geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and mea ...
, Paratethys was characterized by open-marine environments. Brackish and lacustrine basins turned into ventilated seas. Rich marine fauna containing
shark Sharks are a group of elasmobranch cartilaginous fish characterized by a ribless endoskeleton, dermal denticles, five to seven gill slits on each side, and pectoral fins that are not fused to the head. Modern sharks are classified within the ...
s (e.g.,
megalodon ''Otodus megalodon'' ( ; meaning "big tooth"), Common name, commonly known as megalodon, is an extinction, extinct species of giant mackerel shark that lived approximately 23 to 3.6 million years ago (Mya), from the Early Miocene to the Earl ...
),
coral Corals are colonial marine invertebrates within the subphylum Anthozoa of the phylum Cnidaria. They typically form compact Colony (biology), colonies of many identical individual polyp (zoology), polyps. Coral species include the important Coral ...
s,
marine mammal Marine mammals are mammals that rely on marine ecosystems for their existence. They include animals such as cetaceans, pinnipeds, sirenians, sea otters and polar bears. They are an informal group, unified only by their reliance on marine enviro ...
s,
foraminifera Foraminifera ( ; Latin for "hole bearers"; informally called "forams") are unicellular organism, single-celled organisms, members of a phylum or class (biology), class of Rhizarian protists characterized by streaming granular Ectoplasm (cell bio ...
and nanoplankton spread throughout Paratethys from the neighbouring Mediterranean region, probably via the Trans-Tethyan Corridor, an ancient sea-strait located in modern
Slovenia Slovenia, officially the Republic of Slovenia, is a country in Central Europe. It borders Italy to the west, Austria to the north, Hungary to the northeast, Croatia to the south and southeast, and a short (46.6 km) coastline within the Adriati ...
.


Salt Giants

The open marine environments of Paratethys were short-lived, and halfway through the middle Miocene, progressive uplift of the central European mountain ranges and a eustatic drop isolated Paratethys from the global ocean triggering a salinity crisis in Central Paratethys. The " Badenian Salinity Crisis" spanned between 13.8 and 13.4 Mya. Thick evaporitic beds (salt and gypsum) formed in the Outer Carpathians, Transylvanian and Pannonian basins. Salt mines extract this middle-Miocene salt in Transylvania:
Turda Turda (; , ; ; ) is a Municipiu, city in Cluj County, Transylvania, Romania. It is located in the southeastern part of the county, from the county seat, Cluj-Napoca, to which it is connected by the European route E81, and from nearby Câmpia ...
, Ocna Mures, Ocna Sibiului and Praid; in the Eastern and Carpathians: Wieliczka, Bochnia, Cacica and Slanic Prahova; and Ocnele Mari in the Southern Carpathians, but evaporites are also present in areas west of the Carpathians:
Maramureș ( ; ; ; ) is a geographical, historical and cultural region in northern Romania and western Ukraine. It is situated in the northeastern Carpathians, along parts of the upper Tisza River drainage basin; it covers the Maramureș Depression and the ...
, eastern
Slovakia Slovakia, officially the Slovak Republic, is a landlocked country in Central Europe. It is bordered by Poland to the north, Ukraine to the east, Hungary to the south, Austria to the west, and the Czech Republic to the northwest. Slovakia's m ...
(Solivar mine near
Prešov Prešov () is a city in eastern Slovakia. It is the seat of administrative Prešov Region () and Šariš. With a population of approximately 85,000 for the city, and in total more than 100,000 with the urban area, it is the second-largest city i ...
) and, to a lesser extent, in the Pannonian depression in central
Hungary Hungary is a landlocked country in Central Europe. Spanning much of the Pannonian Basin, Carpathian Basin, it is bordered by Slovakia to the north, Ukraine to the northeast, Romania to the east and southeast, Serbia to the south, Croatia and ...
.


Megalake

Some 12 million years ago, slightly before the onset of the
late Miocene The Late Miocene (also known as Upper Miocene) is a sub-epoch of the Miocene epoch (geology), Epoch made up of two faunal stage, stages. The Tortonian and Messinian stages comprise the Late Miocene sub-epoch, which lasted from 11.63 Ma (million ye ...
, the ancient sea transformed into a megalake that covered more than 2.8 million square kilometers, from the eastern Alps to what is now
Kazakhstan Kazakhstan, officially the Republic of Kazakhstan, is a landlocked country primarily in Central Asia, with a European Kazakhstan, small portion in Eastern Europe. It borders Russia to the Kazakhstan–Russia border, north and west, China to th ...
, and characterized by salinities generally ranging between 12 and 14%. During its five-million-year lifetime, the megalake was home to many species found nowhere else, including molluscs and ostracods as well as miniature versions of whales, dolphins and seals. In 2023,
Guinness World Records ''Guinness World Records'', known from its inception in 1955 until 1999 as ''The Guinness Book of Records'' and in previous United States editions as ''The Guinness Book of World Records'', is a British reference book published annually, list ...
named this
lake A lake is often a naturally occurring, relatively large and fixed body of water on or near the Earth's surface. It is localized in a basin or interconnected basins surrounded by dry land. Lakes lie completely on land and are separate from ...
the largest in earth's history. Near the end of the Miocene, an event known as the Khersonian crisis, marked by rapidly fluctuating environmental factors and sea levels, wiped out much of the unique fish fauna of this megalake.


After Paratethys

When parts of the Mediterranean fell dry during the
Messinian salinity crisis In the Messinian salinity crisis (also referred to as the Messinian event, and in its latest stage as the Lago Mare event) the Mediterranean Sea went into a cycle of partial or nearly complete desiccation (drying-up) throughout the latter part of ...
(about 6 million years ago) there were phases when Paratethys water flowed into the deep Mediterranean basins. During the
Pliocene The Pliocene ( ; also Pleiocene) is the epoch (geology), epoch in the geologic time scale that extends from 5.33 to 2.58Pannonian Sea, a
brackish Brackish water, sometimes termed brack water, is water occurring in a natural environment that has more salinity than freshwater, but not as much as seawater. It may result from mixing seawater (salt water) and fresh water together, as in estuari ...
sea in the
Pannonian Basin The Pannonian Basin, with the term Carpathian Basin being sometimes preferred in Hungarian literature, is a large sedimentary basin situated in southeastern Central Europe. After the Treaty of Trianon following World War I, the geomorpholog ...
. Many of these would disappear before the start of the
Pleistocene The Pleistocene ( ; referred to colloquially as the ''ice age, Ice Age'') is the geological epoch (geology), epoch that lasted from to 11,700 years ago, spanning the Earth's most recent period of repeated glaciations. Before a change was fin ...
. At present, only the
Black Sea The Black Sea is a marginal sea, marginal Mediterranean sea (oceanography), mediterranean sea lying between Europe and Asia, east of the Balkans, south of the East European Plain, west of the Caucasus, and north of Anatolia. It is bound ...
,
Caspian Sea The Caspian Sea is the world's largest inland body of water, described as the List of lakes by area, world's largest lake and usually referred to as a full-fledged sea. An endorheic basin, it lies between Europe and Asia: east of the Caucasus, ...
and the
Aral Sea The Aral Sea () was an endorheic lake lying between Kazakhstan to its north and Uzbekistan to its south, which began shrinking in the 1960s and had largely dried up into desert by the 2010s. It was in the Aktobe and Kyzylorda regions of Kazakhst ...
remain of what was once a vast inland sea.


Paleoecology

During the Middle Miocene Climatic Optimum, the Paratethys supported a tropical to subtropical marine ecosystem with very high biodiversity and
endemism Endemism is the state of a species being found only in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also foun ...
, including the establishment of
coral reef A coral reef is an underwater ecosystem characterized by reef-building corals. Reefs are formed of colonies of coral polyps held together by calcium carbonate. Most coral reefs are built from stony corals, whose polyps cluster in group ...
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 The Oligocene ( ) is a geologic epoch (geology), epoch of the Paleogene Geologic time scale, Period that extends from about 33.9 million to 23 million years before the present ( to ). As with other older geologic periods, the rock beds that defin ...
-aged Menilite Formation, a
flysch Flysch () is a sequence of sedimentary rock layers that progress from deep-water and turbidity flow deposits to shallow-water shales and sandstones. It is deposited when a deep basin forms rapidly on the continental side of a mountain building ep ...
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 The Middle Miocene Climatic Transition (MMCT) was a relatively steady period of climatic cooling that occurred around the middle of the Miocene, roughly 14 million years ago (Ma), during the Langhian stage, and resulted in the growth of ice sheet ...
, 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 Zooplankton are the heterotrophic component of the planktonic community (the " zoo-" prefix comes from ), having to consume other organisms to thrive. Plankton are aquatic organisms that are unable to swim effectively against currents. Consequent ...
populations. Despite its diversity, the brief "Paratethyan biodiversity hotspot" was short-lived, lasting for only 3 million years.


Fishes

The isolation of the Paratethys from other ocean basins saw the adaptive radiation and diversification of numerous
endemic Endemism is the state of a species being found only in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also foun ...
fish lineages. These endemic fish were closely related to modern groups, but belonged to their own distinct, now-lost evolutionary radiations. For example,
gobies Gobiidae or gobies is a family of bony fish in the order Gobiiformes, one of the largest fish families comprising over 2,000 species in more than 200 genera. Most of gobiid fish are relatively small, typically less than in length, and the fam ...
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 The Alosidae, or the shads, are a family (biology), family of clupeiform fishes. The family currently comprises four genera worldwide, and about 32 species. The shads are Pelagic fish, pelagic (open water) schooling fish, of which many are anadr ...
and allies) also saw extensive diversification in the Paratethys, with many fossil genera known. Connections with the Mediterranean Sea allowed for many
cartilaginous fish Chondrichthyes (; ) is a class of jawed fish that contains the cartilaginous fish or chondrichthyans, which all have skeletons primarily composed of cartilage. They can be contrasted with the Osteichthyes or ''bony fish'', which have skeleto ...
(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 ''Otodus megalodon'' ( ; meaning "big tooth"), Common name, commonly known as megalodon, is an extinction, extinct species of giant mackerel shark that lived approximately 23 to 3.6 million years ago (Mya), from the Early Miocene to the Earl ...
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 Cetacea (; , ) is an infraorder of aquatic mammals belonging to the order Artiodactyla that includes whales, dolphins and porpoises. Key characteristics are their fully aquatic lifestyle, streamlined body shape, often large size and exclusively c ...
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

* * * *


References


Further reading

*


External links

* * {{Authority control Historical oceans Oligocene Europe Miocene Europe Pliocene Europe History of the Mediterranean Seas of the Mediterranean Sea