Эоцен: Вторая эпоха палеогена

Second epoch of the Paleogene Period

The Eocene (/ipa/'/iː//ə//s//iː//n//, /'/iː//oʊ// / EEəseen, EEoh ) is a geological epoch that lasted from about 56 to 33.9 million years ago (Ma). It is the second epoch of the Paleogene Period in the modern Cenozoic Era. The name Eocene comes from the Ancient Greek ἠώς (ēṓs, "dawn") and καινός (kainós, "new") and refers to the "dawn" of modern ('new') fauna that appeared during the epoch. The Eocene spans the time from the end of the Paleocene Epoch to the beginning of the Oligocene Epoch. The start of the Eocene is marked by a brief period in which the concentration of the carbon isotope 13C in the atmosphere was exceptionally low in comparison with the more common isotope 12C. The average temperature of Earth in the beginning of the Eocene was about 27 degrees Celsius. The end is set at a major extinction event called the Grande Coupure (the "Great Break" in continuity) or the Eocene–Oligocene extinction event, which may be related to the impact of one or more large bolides in Siberia and in what is now Chesapeake Bay. As with other geologic periods, the strata that define the start and end of the epoch are well identified, though their exact dates are slightly uncertain.

The term "Eocene" is derived from Ancient Greek ἠώς eos meaning "dawn", and καινός kainos meaning "new" or "recent", as the epoch saw the dawn of recent, or modern, life. Scottish geologist Charles Lyell (ignoring the Quaternary) divided the Tertiary Epoch into the Eocene, Miocene, Pliocene, and New Pliocene (Holocene) Periods in 1833. British geologist John Phillips proposed the Cenozoic in 1840 in place of the Tertiary, and Austrian paleontologist Moritz Hörnes introduced the Paleogene for the Eocene and Neogene for the Miocene and Pliocene in 1853. After decades of inconsistent usage, the newly formed International Commission on Stratigraphy (ICS), in 1969, standardized stratigraphy based on the prevailing opinions in Europe: the Cenozoic Era subdivided into the Tertiary and Quaternary sub eras, and the Tertiary subdivided into the Paleogene and Neogene periods. In 1978, the Paleogene was officially defined as the Paleocene, Eocene, and Oligocene epochs; and the Neogene as the Miocene and Pliocene epochs. In 1989, Tertiary and Quaternary were removed from the time scale due to the arbitrary nature of their boundary, but Quaternary was reinstated in 2009.

The Eocene is a dynamic epoch that represents global climatic transitions between two climatic extremes, transitioning from the hot house to the cold house. The beginning of the Eocene is marked by the Paleocene–Eocene Thermal Maximum, a short period of intense warming and ocean acidification brought about by the release of carbon en masse into the atmosphere and ocean systems, This event happened around 55.8 Ma, and was one of the most significant periods of global change during the Cenozoic. The middle Eocene was characterized by the shift towards a cooler climate at the end of the EECO, around 47.8 Ma, which was briefly interrupted by another warming event called the middle Eocene climatic optimum (MECO). Lasting for about 400,000 years, the MECO was responsible for a globally uniform 4° to 6°C warming of both the surface and deep oceans, as inferred from foraminiferal stable oxygen isotope records. The resumption of a long term gradual cooling trend resulted in a glacial maximum at the late Eocene/early Oligocene boundary. The end of the Eocene was also marked by the Eocene–Oligocene extinction event, also known as the Grande Coupure.

The Eocene is conventionally divided into early (56–47.8 Ma), middle (47.8–38 Ma), and late (38–33.9 Ma) subdivisions. The corresponding rocks are referred to as lower, middle, and upper Eocene. The Ypresian Stage constitutes the lower, the Priabonian Stage the upper; and the Lutetian and Bartonian stages are united as the middle Eocene. The Western North American floras of the Eocene were divided into four floral "stages" by Jack Wolfe (1968) based on work with the Puget Group fossils of King County, Washington. The four stages, Franklinian, Fultonian, Ravenian, and Kummerian covered the Early Eocene through early Oligocene, and three of the four were given informal early/late substages. Wolfe tentatively deemed the Franklinian as Early Eocene, the Fultonian as Middle Eocene, the Ravenian as Late, and the Kummerian as Early Oligocene. The beginning of the Kummerian was refined by Gregory Retallack et al (2004) as 40 mya, with a refined end at the Eocene Oligocene boundary where the younger Angoonian floral stage starts.

During the Eocene, the continents continued to drift toward their present positions. At the beginning of the period, Australia and Antarctica remained connected, and warm equatorial currents may have mixed with colder Antarctic waters, distributing the heat around the planet and keeping global temperatures high. When Australia split from the southern continent around 45 Ma, the warm equatorial currents were routed away from Antarctica. An isolated cold water channel developed between the two continents. However, modeling results call into question the thermal isolation model for late Eocene cooling, and decreasing carbon dioxide levels in the atmosphere may have been more important. Once the Antarctic region began to cool down, the ocean surrounding Antarctica began to freeze, sending cold water and icefloes north and reinforcing the cooling. The northern supercontinent of Laurasia began to fragment, as Europe, Greenland and North America drifted apart. In western North America, the Laramide Orogeny came to an end in the Eocene, and compression was replaced with crustal extension that ultimately gave rise to the Basin and Range Province. The Kishenehn Basin, around 1.5 km in elevation during the Lutetian, was uplifted to an altitude of 2.5 km by the Priabonian. Huge lakes formed in the high flat basins among uplifts, resulting in the deposition of the Green River Formation lagerstätte. At about 35 Ma, an asteroid impact on the eastern coast of North America formed the Chesapeake Bay impact crater. The Tethys Ocean finally closed with the collision of Africa and Eurasia, while the uplift of the Alps isolated its final remnant, the Mediterranean, and created another shallow sea with island archipelagos to the north. Planktonic foraminifera in the northwestern Peri Tethys are very similar to those of the Tethys in the middle Lutetian but become completely disparate in the Bartonian, indicating biogeographic separation. Though the North Atlantic was opening, a land connection appears to have remained between North America and Europe since the faunas of the two regions are very similar. Eurasia was separated in three different landmasses 50 Ma; Western Europe, Balkanatolia and Asia. About 40 Ma, Balkanatolia and Asia were connected, while Europe was connected 34 Ma. The Fushun Basin contained large, suboxic lakes known as the paleo Jijuntun Lakes. India collided with Asia, folding to initiate formation of the Himalayas. The incipient subcontinent collided with the Kohistan–Ladakh Arc around 50.2 Ma and with Karakoram around 40.4 Ma, with the final collision between Asia and India occurring ~40 Ma.

The Eocene Epoch contained a wide variety of different climate conditions that includes the warmest climate in the Cenozoic Era, and arguably the warmest time interval since the Permian Triassic mass extinction and Early Triassic, and ends in an icehouse climate. The evolution of the Eocene climate began with warming after the end of the Paleocene–Eocene Thermal Maximum (PETM) at 56 Ma to a maximum during the Eocene Optimum at around 49 Ma. During this period of time, little to no ice was present on Earth with a smaller difference in temperature from the equator to the poles. Because of this the maximum sea level was 150 meters higher than current levels. Following the maximum was a descent into an icehouse climate from the Eocene Optimum to the Eocene–Oligocene transition at 34 Ma. During this decrease, ice began to reappear at the poles, and the Eocene–Oligocene transition is the period of time when the Antarctic ice sheet began to rapidly expand.

Greenhouse gases, in particular carbon dioxide and methane, played a significant role during the Eocene in controlling the surface temperature. The end of the PETM was met with very large sequestration of carbon dioxide into the forms of methane clathrate, coal, and crude oil at the bottom of the Arctic Ocean, that reduced the atmospheric carbon dioxide. Other proxies such as pedogenic (soil building) carbonate and marine boron isotopes indicate large changes of carbon dioxide of over 2,000 ppm over periods of time of less than 1 million years. Most of the methane released to the atmosphere during this period of time would have been from wetlands, swamps, and forests. During the warming in the early Eocene between 55 and 52 Ma, there were a series of short term changes of carbon isotope composition in the ocean. These isotope changes occurred due to the release of carbon from the ocean into the atmosphere that led to a temperature increase of at the surface of the ocean. Recent analysis of and research into these hyperthermals in the early Eocene has led to hypotheses that the hyperthermals are based on orbital parameters, in particular eccentricity and obliquity. The hyperthermals in the early Eocene, notably the Palaeocene–Eocene Thermal Maximum (PETM), the Eocene Thermal Maximum 2 (ETM2), and the Eocene Thermal Maximum 3 (ETM3), were analyzed and found that orbital control may have had a role in triggering the ETM2 and ETM3. An enhancement of the biological pump proved effective at sequestering excess carbon during the recovery phases of these hyperthermals. These hyperthermals led to increased perturbations in planktonic and benthic foraminifera, with a higher rate of fluvial sedimentation as a consequence of the warmer temperatures. Unlike the PETM, the lesser hyperthermals of the Early Eocene had negligible consequences for terrestrial mammals. These Early Eocene hyperthermals produced a sustained period of extremely hot climate known as the Early Eocene Climatic Optimum (EECO). During the early and middle EECO, the superabundance of the euryhaline dinocyst Homotryblium in New Zealand indicates elevated ocean salinity in the region.

One of the unique features of the Eocene's climate as mentioned before was the equable and homogeneous climate that existed in the early parts of the Eocene. A multitude of proxies support the presence of a warmer equable climate being present during this period of time. A few of these proxies include the presence of fossils native to warm climates, such as crocodiles, located in the higher latitudes, although clumped isotope analyses point to a maximum low latitude sea surface temperature of ± during the EECO. Relative to present day values, bottom water temperatures are higher according to isotope proxies. led to stagnant waters and as the azolla sank to the sea floor, they became part of the sediments on the seabed and effectively sequestered the carbon by locking it out of the atmosphere for good. The ability for the azolla to sequester carbon is exceptional, and the enhanced burial of azolla could have had a significant effect on the world atmospheric carbon content and may have been the event to begin the transition into an ice house climate. continued due to continual decrease in atmospheric carbon dioxide from organic productivity and weathering from mountain building. Global cooling continued until there was a major reversal from cooling to warming in the Bartonian. This warming event, signifying a sudden and temporary reversal of the cooling conditions, is known as the Middle Eocene Climatic Optimum (MECO). At around 41.5 Ma, stable isotopic analysis of samples from Southern Ocean drilling sites indicated a warming event for 600,000 years. Oxygen isotope analysis showed a large negative change in the proportion of heavier oxygen isotopes to lighter oxygen isotopes, which indicates an increase in global temperatures. The warming is considered to be primarily due to carbon dioxide increases, because carbon isotope signatures rule out major methane release during this short term warming. The increase in atmospheric carbon dioxide has also been hypothesised to have been driven by increased seafloor spreading rates and metamorphic decarbonation reactions between Australia and Antarctica and increased amounts of volcanism in the region. One possible cause of atmospheric carbon dioxide increase could have been a sudden increase due to metamorphic release due to continental drift and collision of India with Asia and the resulting formation of the Himalayas; however, data on the exact timing of metamorphic release of atmospheric carbon dioxide is not well resolved in the data. Yet another explanation hypothesises that MECO warming was caused by the simultaneous occurrence of minima in both the 400 kyr and 2.4 Myr eccentricity cycles. During the MECO, sea surface temperatures in the Tethys Ocean jumped to 32–36 °C, and Tethyan seawater became more dysoxic. A decline in carbonate accumulation at ocean depths of greater than three kilometres took place synchronously with the peak of the MECO, signifying ocean acidification took place in the deep ocean. On top of that, MECO warming caused an increase in the respiration rates of pelagic heterotrophs, leading to a decreased proportion of primary productivity making its way down to the seafloor and causing a corresponding decline in populations of benthic foraminifera. An abrupt decrease in lakewater salinity in western North America occurred during this warming interval. This warming is short lived, as benthic oxygen isotope records indicate a return to cooling at ~40 Ma. The post MECO cooling brought with it a major aridification trend in Asia, enhanced by retreating seas. The cooling during the initial stages of the opening of the Drake Passage ~38.5 Ma was not global, as evidenced by an absence of cooling in the North Atlantic. During the cooling period, benthic oxygen isotopes show the possibility of ice creation and ice increase during this later cooling.

During the early middle Eocene, forests covered most of the Earth including the poles. Tropical forests extended across much of modern Africa, South America, Central America, India, South east Asia and China. Paratropical forests grew over North America, Europe and Russia, with broad leafed evergreen and broad leafed deciduous forests at higher latitudes. Polar forests were quite extensive. Fossils and even preserved remains of trees such as swamp cypress and dawn redwood from the Eocene have been found on Ellesmere Island in the Arctic. Even at that time, Ellesmere Island was only a few degrees in latitude further south than it is today. Fossils of subtropical and even tropical trees and plants from the Eocene also have been found in Greenland and Alaska. Tropical rainforests grew as far north as northern North America and Europe. Palm trees were growing as far north as Alaska and northern Europe during the early Eocene, although they became less abundant as the climate cooled. Dawn redwoods were far more extensive as well. The earliest definitive Eucalyptus fossils were dated from 51.9 Ma, and were found in the Laguna del Hunco deposit in Chubut province in Argentina. Cooling began mid period, and by the end of the Eocene continental interiors had begun to dry, with forests thinning considerably in some areas. The newly evolved grasses were still confined to river banks and lake shores, and had not yet expanded into plains and savannas. The cooling also brought seasonal changes. Deciduous trees, better able to cope with large temperature changes, began to overtake evergreen tropical species. By the end of the period, deciduous forests covered large parts of the northern continents, including North America, Eurasia and the Arctic, and rainforests held on only in equatorial South America, Africa, India and Australia. Antarctica began the Eocene fringed with a warm temperate to sub tropical rainforest. Pollen found in Prydz Bay from the Eocene suggest taiga forest existed there. It became much colder as the period progressed; the heat loving tropical flora was wiped out, and by the beginning of the Oligocene, the continent hosted deciduous forests and vast stretches of tundra.

During the Eocene, plants and marine faunas became quite modern. Many modern bird orders first appeared in the Eocene. The Eocene oceans were warm and teeming with fish and other sea life.

The oldest known fossils of most of the modern mammal orders appear within a brief period during the early Eocene. At the beginning of the Eocene, several new mammal groups arrived in North America. These modern mammals, like artiodactyls, perissodactyls, and primates, had features like long, thin legs, feet, and hands capable of grasping, as well as differentiated teeth adapted for chewing. Dwarf forms reigned. All the members of the new mammal orders were small, under 10 kg; based on comparisons of tooth size, Eocene mammals were only 60% of the size of the primitive Palaeocene mammals that preceded them. They were also smaller than the mammals that followed them. It is assumed that the hot Eocene temperatures favored smaller animals that were better able to manage the heat. Both groups of modern ungulates (hoofed animals) became prevalent because of a major radiation between Europe and North America, along with carnivorous ungulates like Mesonyx. Early forms of many other modern mammalian orders appeared, including horses (most notably the Eohippus), bats, proboscidians (elephants), primates, and rodents. Older primitive forms of mammals declined in variety and importance. Important Eocene land fauna fossil remains have been found in western North America, Europe, Patagonia, Egypt, and southeast Asia. Marine fauna are best known from South Asia and the southeast United States. Established megafauna of the Eocene include the Uintatherium, Arsinoitherium, and brontotheres, in which the former two, unlike the latter, did not belong to ungulates but groups that became extinct shortly after their establishments. Large terrestrial mammalian predators had already existed since the Paleocene, but new forms now arose like Hyaenodon and Daphoenus (the earliest lineage of a once successful predatory family known as bear dogs). Entelodonts meanwhile established themselves as some of the largest omnivores. The first nimravids, including Dinictis, established themselves as amongst the first feliforms to appear. Their groups became highly successful and continued to live past the Eocene. Basilosaurus is a very well known Eocene whale, but whales as a group had become very diverse during the Eocene, which is when the major transitions from being terrestrial to fully aquatic in cetaceans occurred. The first sirenians were evolving at this time, and would eventually evolve into the extant manatees and dugongs. It is thought that millions of years after the Cretaceous Paleogene extinction event, brain sizes of mammals now started to increase, "likely driven by a need for greater cognition in increasingly complex environments".

Eocene birds include some enigmatic groups with resemblances to modern forms, some of which continued from the Paleocene. Bird taxa of the Eocene include carnivorous psittaciforms, such as Messelasturidae, Halcyornithidae, large flightless forms such as Gastornis and Eleutherornis, long legged falcon Masillaraptor, ancient galliforms such as Gallinuloides, putative rail relatives of the family Songziidae, various pseudotooth birds such as Gigantornis, the ibis relative Rhynchaeites, primitive swifts of the genus Aegialornis, and primitive penguins such as Archaeospheniscus and Inkayacu.

Reptile fossils from this time, such as fossils of pythons and turtles, are abundant.

Several rich fossil insect faunas are known from the Eocene, notably the Baltic amber found mainly along the south coast of the Baltic Sea, amber from the Paris Basin, France, the Fur Formation, Denmark, and the Bembridge Marls from the Isle of Wight, England. Insects found in Eocene deposits mostly belong to genera that exist today, though their range has often shifted since the Eocene. For instance the bibionid genus Plecia is common in fossil faunas from presently temperate areas, but only lives in the tropics and subtropics today.