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Tortonian
11.63 – 7.246 Ma
An map of the world 10 million years ago, during the Tortonian Age.
Chronology
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North American prairie expands[2]
Subdivision of the Neogene according to the ICS, as of 2024.[3]
Vertical axis scale: Millions of years ago
Formerly part ofTertiary Period/System
Etymology
Name formalityFormal
Usage information
Celestial bodyEarth
Regional usageGlobal (ICS)
Time scale(s) usedICS Time Scale
Definition
Chronological unitAge
Stratigraphic unitStage
Time span formalityFormal
Lower boundary definitionLAD of the Haptophyte Discoaster kugleri
Lower boundary GSSPMonte dei Corvi Beach section, Ancona, Italy
43°35′12″N 13°34′10″E / 43.5867°N 13.5694°E / 43.5867; 13.5694
Lower GSSP ratified2003[4]
Upper boundary definition
Upper boundary GSSPOued Akrech section, Rabat, Morocco
33°56′13″N 6°48′45″W / 33.9369°N 6.8125°W / 33.9369; -6.8125
Upper GSSP ratifiedJanuary 2000[5]

The Tortonian is in the geologic time scale an age or stage of the late Miocene that spans the time between 11.608 ± 0.005 Ma and 7.246 ± 0.005 Ma (million years ago).[3] It follows the Serravallian and is followed by the Messinian.

The Tortonian roughly overlaps with the regional Pannonian Stage of the Paratethys timescale of Central Europe. It also overlaps the upper Astaracian, Vallesian and lower Turolian European land mammal ages, the upper Clarendonian and lower Hemphillian North American land mammal ages and the upper Chasicoan and lower Huayquerian South American land mammal ages.

Definition

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The Tortonian was introduced by Swiss stratigrapher Karl Mayer-Eymar in 1858. It was named after the Italian city of Tortona in the Piedmont region.

The base of the Tortonian Stage is at the last common appearance of calcareous nanoplankton Discoaster kugleri and planktonic foram Globigerinoides subquadratus. It is also associated with the short normal polarized magnetic chronozone C5r.2n. A GSSP for the Tortonian has been established in the Monte dei Corvi section near Ancona (Italy).[6]

The top of the Tortonian (the base of the Messinian) is at the first appearance of the planktonic foram species Globorotalia conomiozea and is stratigraphically in the middle of magnetic chronozone C3Br.1r.

Geologic history

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In 2020, geologists reported two newly-identified supervolcano eruptions associated with the Yellowstone hotspot track, including the region's largest and most cataclysmic event – the Grey's Landing super-eruption – which had a volume of at least 2,800 cubic kilometres (670 cubic miles) and occurred around 8.72 Ma.[7][8]

Around 10 Ma, the inflow of North Atlantic Deep Water (NADW) into the Indian Ocean increased significantly.[9]

References

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Revisions and contributorsEdit on WikipediaRead on Wikipedia
from Grokipedia
The Tortonian is a stage in the Late Miocene Epoch of the Neogene Period, spanning from 11.63 to 7.246 million years ago (Ma).[1] It represents a key interval in Earth's geologic history, defined by its Global Boundary Stratotype Section and Point (GSSP) at the Monte dei Corvi Beach section on the Adriatic coast of central Italy (43°35'12" N, 13°34'10" E), where the base is placed at the midpoint of the sapropel layer within sedimentary cycle 76.[2] This boundary coincides with the last common occurrence of the nannofossil Discoaster kugleri and the foraminifer Globigerinoides subquadratus, as well as the base of magnetic subchron C5r.2n and oxygen isotope event Mi-5, corresponding to an astronomically tuned age of 11.608 Ma.[3] During the Tortonian, significant paleogeographic and climatic changes shaped global environments, including the continued uplift of the Tibetan Plateau and Himalayan ranges, which influenced monsoon intensification and atmospheric circulation patterns.[4] The stage is marked by the Tortonian Thermal Maximum (TTM) around 10.5–9 Ma, a transient episode of global warming driven by orbital forcing and elevated CO₂ levels, leading to warmer sea surface temperatures, expanded tropical belts, and increased precipitation in mid-latitude regions such as the "Late Miocene washhouse climate" in Europe.[5] This warming facilitated the widespread expansion of C₄ grasslands across low- to mid-latitude continents, particularly in Africa, Asia, and the Americas, as fire-adapted C₄ grasses gained competitive advantages over C₃ vegetation under declining atmospheric CO₂ and heightened aridity or seasonality.[6] Biologically, the Tortonian witnessed the diversification of modern mammalian faunas, reflecting the ecological shift toward savannas and woodlands. In marine realms, Mediterranean biodiversity underwent transformations due to tectonic restriction of Atlantic connections, promoting endemic species while global oceans supported diverse planktic and benthic communities.[7] Marine records from this period show fluctuating sea levels tied to glacio-eustatic cycles, with the stage culminating in cooling trends toward the Messinian boundary, setting the stage for the Messinian Salinity Crisis.[7] These developments underscore the Tortonian as a pivotal transition in Cenozoic evolution, bridging warmer Miocene climates with the cooler Pliocene.

Definition and nomenclature

Etymology and historical context

The Tortonian stage derives its name from the town of Tortona in the Piedmont region of northern Italy, where distinctive blue marls containing the fossil mollusks Conus canaliculatus and Ancillaria glandiformis were first identified and described.[3] These sediments, characteristic of marine deposits from the late Miocene, provided the basis for the stage's initial definition.[3] The stage was formally established in 1858 by Swiss stratigrapher and paleontologist Karl Mayer-Eymar (1826–1907) as part of his pioneering subdivision of the Miocene into chronostratigraphic units.[3] Mayer-Eymar, known for his extensive work on Cenozoic mollusks and regional stratigraphy, introduced the Tortonian as a regional stage primarily within the Paratethys and Mediterranean basins, drawing on correlations across Europe to delineate its extent.[8] His classification emphasized biostratigraphic markers, particularly mollusk assemblages, to link disparate sections and establish a coherent framework for Miocene correlations.[3] The original stratotype was designated at the Rio Mazzapiedi-Castellania section by Gianotti in 1953, though it was later superseded by the GSSP due to correlation issues.[3] Early recognition of the Tortonian centered on fossil mollusks from key European localities, including the Piedmont Basin in Italy, the Vienna Basin, and the Cabrières d'Aigues area in France, where marine facies were correlated with continental deposits bearing Hipparion fauna.[3] These correlations highlighted the stage's utility in integrating litho- and biostratigraphy across the Paratethys and Mediterranean realms, though initial definitions extended it upward to the base of the Piacenzian until refinements in the 1860s.[3] Over the subsequent decades, the Tortonian transitioned from a predominantly local lithostratigraphic unit—rooted in Mayer-Eymar's regional observations—to a standardized chronostratigraphic stage through iterative international collaborations and boundary adjustments, such as the 1867–1868 definition of the overlying Messinian.[3] By the late 20th century, it achieved global recognition as a key Miocene division, with subsequent formalization via a Global Stratotype Section and Point (detailed elsewhere).[3]

Global Stratotype Section and Point (GSSP)

The Global Stratotype Section and Point (GSSP) for the base of the Tortonian Stage is located at the Monte dei Corvi Beach section on the Adriatic coast, approximately 5 km southeast of Ancona in the Ancona Province of central Italy, at coordinates 43°35′12″N 13°34′10″E.[2] This site was selected due to its continuous, well-exposed sedimentary record that allows precise correlation using multiple stratigraphic tools, including cyclostratigraphy, biostratigraphy, magnetostratigraphy, and chemostratigraphy. The section is accessible via a coastal trail and spans the Serravallian-Tortonian boundary within the upper part of the Schlier Formation.[3] The base of the Tortonian is defined at the midpoint of the sapropel layer within the lithological cycle known as basic cycle 76 (BC 76), which is astronomically tuned to a precession-related cycle and correlated to the base of magnetic subchron C5r.2n.[2] This definition provides a primary criterion based on the rhythmic sedimentation driven by orbital forcing, with secondary markers including the last common occurrence (LCO) of the calcareous nannofossil Discoaster kugleri (marking the base of standard nannofossil zone NN9) and the LCO of the planktonic foraminifer Globigerinoides subquadratus.[3] Additional supporting events at or near the boundary include oxygen isotope event Mi-5, a glacio-eustatic lowstand (sequence boundary T3.1), and a deep-sea hiatus (NH4). The section's lithology consists of cyclic alternations of greenish-grey marls, whitish marly limestones, and organic-rich sapropels, reflecting Milankovitch-band orbital cycles of eccentricity, obliquity, and precession, with two volcanic ash layers (Respighi and Ancona) aiding tephrostratigraphic correlation.[2] The GSSP was proposed by the Subcommission on Neogene Stratigraphy and ratified by the International Union of Geological Sciences (IUGS) in 2003, following international voting, with formal publication in 2005.[2] An auxiliary reference section at Monte Gibliscemi in Sicily complements the primary site by offering better preservation of microfossils for biostratigraphic correlation. The astronomically calibrated age of the boundary is 11.63 Ma, with refinements to this estimate continuing through updated astronomical models and radioisotopic dating of ash layers.[3]

Stratigraphy and chronology

Temporal boundaries

The Tortonian stage encompasses the interval from its base at 11.63 million years ago (Ma) to its top at 7.246 Ma, yielding a duration of about 4.38 million years.[9] This places the Tortonian within the late Miocene epoch of the Neogene period. The base of the Tortonian, defining the boundary with the underlying Serravallian stage, is calibrated through astronomically tuned cyclostratigraphy integrated with magnetostratigraphy, as established at the Global Stratotype Section and Point (GSSP).[10] Similarly, the top boundary with the Messinian stage relies on astronomical tuning of sedimentary cycles.[11] These absolute ages result from the integration of orbital forcing related to Milankovitch cycles, radioisotopic dating via the ⁴⁰Ar/³⁹Ar method on intercalated volcanic ash layers, and supporting biostratigraphic correlations.[10][12] Refinements in the 2020 Geologic Time Scale (GTS2020) incorporate these techniques to provide the current precise boundaries, with minor adjustments from prior broader estimates of 11.6–7.2 Ma that lacked such high-resolution tuning.[13]

Subdivisions and biozonation

The Tortonian is informally subdivided into three parts based on lithological cycles observed in astronomically tuned Mediterranean sections and associated faunal turnovers in marine microfossils: the lower Tortonian (11.63–10.5 Ma), middle Tortonian (10.5–8.6 Ma), and upper Tortonian (8.6–7.25 Ma).[14] These divisions reflect periodic alternations in sedimentation patterns driven by orbital forcing, with transitions marked by changes in carbonate content and biostratigraphic events.[3] Biostratigraphic correlation within the Tortonian relies primarily on planktonic foraminiferal zones N17 and N18 of the standard low-latitude scheme. Zone N17 is defined by the first appearance and evolution of the Globorotalia menardii lineage, spanning much of the lower and middle Tortonian, while Zone N18 is characterized by the dominance of Neogloboquadrina acostaensis (formerly N. dutertrei acostaensis) and extends into the upper Tortonian, with subzones refined by accessory events such as the coiling change in Globorotalia menardii. These zones provide high-resolution markers for global correlation, particularly in open-ocean settings.[15] Calcareous nannofossil biozonation complements foraminiferal schemes, with the Tortonian encompassing zones CN8 to CN9 (Okada and Bukry, 1980). Zone CN8, the Discoaster hamatus zone, covers the lower Tortonian and is bounded above by the last common occurrence of D. hamatus, while CN9, the Discoaster quinquelobus zone, dominates the middle to upper Tortonian and is defined by the first regular occurrence of D. quinquelobus, highlighting evolutionary shifts in discoasterids.[16] These zones are particularly useful in hemipelagic to pelagic carbonates for pinpointing faunal turnovers.[17] Regional variations in biozonation occur due to provincialism. In the Central Paratethys, the Tortonian aligns with the upper Badenian stage, where equivalent assemblages show N17–N18 foraminiferal markers alongside local nannofossil events adjusted for restricted marine conditions. In the Indo-Pacific, correlations emphasize N17–N18 zones with enhanced diversity in globorotaliids, facilitating links to Tethyan sections.[18] Global correlation of Tortonian subdivisions integrates magnetostratigraphy, with the stage spanning chrons C5r through C3An (including subchrons like C5r.2n at the base and C3Br.1r near the top).[3] Sapropel cycles in Mediterranean sequences, such as cycles 66–76, provide additional tuning points tied to precession-driven eccentricity, enhancing precision when combined with bioevents.[19]

Paleoenvironment

Paleogeography

During the Tortonian stage (11.63–7.246 Ma), global continental configurations approached their modern positions, as reconstructed from plate tectonic models integrating paleomagnetic, stratigraphic, and geophysical data.[20] Key differences persisted in the Mediterranean region, where the Tethys Ocean remnants were undergoing final closure, and in the Red Sea-Gulf of Aden area, which saw progressive rifting. These configurations reflect ongoing subduction along convergent margins and continental collisions driven by plate motions at rates of 2–5 cm/year.[21] The closure of the Tethys Ocean progressed significantly during the Tortonian, with the eastern connection to the Indian Ocean having terminated by the preceding Serravallian due to northward migration of the African-Arabian plates and associated sea-level fluctuations.[22] In the western Mediterranean, this narrowing set the stage for precursors to the Messinian Salinity Crisis, as the Rifian Corridor in Morocco and the Betic Corridor in southern Spain increasingly restricted Atlantic inflow, reducing exchange to episodic pulses by around 7.16 Ma.[23] Concurrently, the African-Arabian plates continued their northward drift at approximately 1–2 cm/year, intensifying collision with Eurasia and contributing to the uplift of the Anatolian Plateau and the Zagros Mountains through crustal shortening and subduction-related compression.[21] This motion also affected the East African Rift system, initiated in the early Miocene around 25 Ma, which continued to experience rifting and basaltic volcanism during the Tortonian, marking ongoing divergence between the Nubian and Somalian plates.[24] The expansion of the Antarctic Ice Sheet during the late Miocene, including toward the end of the Tortonian, contributed to global sea-level lowering of approximately 20–50 m relative to mid-Miocene highs, though levels remained close to or slightly below present during much of the stage, exposing additional continental shelves and influencing marginal marine sedimentation worldwide.[25] Regionally, the uplift of the Tibetan Plateau accelerated in the early Tortonian, with the northern sector (Qaidam Basin) rising by 1,650 ± 450 meters between 11.3 and 10.8 Ma due to mantle lithosphere detachment, which mechanically enhanced Asian monsoon circulation by altering atmospheric dynamics over the plateau.[26] In the Pacific, ongoing subduction along the Ring of Fire generated island arcs, as evidenced by volcanic folding and magmatism in the Ryukyu Arc during the Serravallian-Tortonian transition, where the Philippine Sea Plate resumed subduction beneath the Eurasian margin.[27] Similarly, the North American Cordilleran orogeny persisted through Miocene subduction of the Farallon Plate remnants, driving extension in the Basin and Range Province and associated volcanism from 12–8 Ma.[28]

Climate and oceanography

The Tortonian stage was characterized by a globally warmer climate than present, with mean surface temperatures estimated at 3–6°C higher, accompanied by a reduced equator-to-pole temperature gradient that facilitated poleward heat transport.[29] This period marked a transition from the mid-Miocene Climatic Optimum toward late Miocene cooling, driven by declining atmospheric CO₂ levels from approximately 400 ppm to 300 ppm, which contributed to stepwise global temperature decreases of up to 2–4°C in tropical and high-latitude regions.[29] The onset of significant Antarctic ice sheet volume around 10 Ma further amplified this cooling trend, leading to eustatic sea-level falls of 20–50 m relative to earlier Miocene highs.[29][30] Precipitation patterns during the Tortonian showed enhanced monsoonal activity in Asia, intensified around 8 Ma due to the uplift of the Tibetan Plateau, which strengthened summer rainfall and associated riverine discharge.[29] Tropics and subtropics generally experienced wetter conditions, with increased humidity supporting expansive vegetation belts, while mid-latitudes underwent progressive aridification, evidenced by expanding desert margins in Eurasia and North Africa.[29] These shifts were modulated by orbital precession, which amplified seasonal contrasts and monsoon variability.[31] Oceanic conditions featured a global thermohaline circulation broadly similar to modern patterns but with warmer polar surface waters, promoting reduced sea ice extent and enhanced meridional overturning that emerged around 13.9–9 Ma.[29] In the Mediterranean, deep-water formation was increasingly disrupted by the restriction of Atlantic gateways during the late Tortonian (around 7.5 Ma), leading to stratified water columns, oxygen depletion, and periodic anoxic events marked by sapropel deposition in deeper basins.[31] This isolation fostered bottom-water stagnation and elevated salinities, as indicated by benthic foraminiferal assemblages shifting toward low-oxygen tolerant species.[31] Globally, the expansion of Pacific oxygen-deficient zones began around 9–8 Ma, linked to rising deep-ocean nutrients and cooling-induced circulation changes.[32] Key paleoclimatic proxies underscore these trends: benthic and planktic foraminiferal δ¹⁸O records reveal a cooling signal of 0.5–1‰ over the stage, reflecting both temperature drops and growing ice volume.[29] Alkenone-based sea surface temperature (SST) estimates indicate low-latitude values of 25–30°C, with a gradual decline toward the stage's end, while high-latitude SSTs cooled by up to 8°C.[29][32] These data, corroborated by TEX₈₆ and Mg/Ca ratios, highlight the Tortonian's role in establishing modern-like ocean- atmosphere coupling amid ongoing cooling.[29]

Paleontology

Marine biota

During the Tortonian stage of the Late Miocene, marine plankton communities were dominated by siliceous diatoms and calcareous coccolithophores, which formed the primary basis of oceanic food webs and contributed significantly to silica and carbonate export to the seafloor.[33][34] Diatoms, such as those in the genus Thalassiosira, exhibited high abundances in nutrient-rich waters, reflecting enhanced primary productivity linked to regional upwelling systems.[35] Coccolithophores, including species like Helicosphaera carteri var. wallichii, showed increased diversity and first occurrences, underscoring their ecological expansion in subtropical to temperate surface waters. Calcareous nannoplankton underwent a notable radiation, with Discoaster species such as D. brouweri, D. pentaradiatus, and D. challengeri becoming prominent markers of this interval, indicating stable warm-water conditions favorable for their calcification.[36][37] Planktonic foraminifera experienced significant turnover, characterized by the last occurrence of Paragloborotalia mayeri and the first appearance of Neogloboquadrina acostaensis, alongside keeled forms like Globorotalia continuosa and G. mayeri, signaling shifts in thermocline depth and water mass stratification.[38][39] These changes reflect a transition to more oligotrophic conditions in parts of the Tethyan realm, with neogloboquadrinids adapting to deeper habitats.[40] Benthic communities in shelf environments featured high diversity among mollusks, particularly bivalves and gastropods, which thrived in shallow, oxygenated settings with soft to firm substrates.[41] In the Mediterranean, pectinid bivalves such as Gigantopecten species formed dense assemblages on hardgrounds, often bioeroded and associated with rhodoliths, indicating stable, low-energy depositional regimes.[42] Echinoids, including clypeasteroids like Clypeaster, and bryozoans were abundant in temperate seas, contributing to encrusting and infaunal biofacies on carbonate platforms. Coral reefs in the Indo-Pacific region underwent a decline during the Tortonian, attributed to cooling sea surface temperatures and associated environmental stressors (such as monsoon belt shifts and changes in ocean currents) that reduced framework-building capacity.[43] Symbiont-bearing colonial corals shifted southward in distribution, with limited biodiversity and increased heterotrophy in marginal settings, marking the transition from peak Miocene reef development.[44][45] Marine mammals diversified notably, with early radiation among baleen whales (Mysticeti) including basal balaenopterids like those from the Stirone River deposits in Italy, which exhibit archaic skeletal features adapted for filter-feeding in productive coastal waters. Recent discoveries include an archaic balaenid whale from late Tortonian sediments in Japan, filling gaps in right whale evolution.[46][47][48] Toothed whales (Odontoceti) also expanded, with species showing increased body sizes and predatory adaptations in the Paratethys and Mediterranean basins; a new beaked whale species from the Pisco Formation in Peru, revised to Tortonian age as of 2024, highlights odontocete diversity in the Pacific.[49][50] Sirenians, particularly dugong-like forms such as Metaxytherium, were widespread in coastal Tethys environments, grazing on seagrasses in shallow, warm embayments.[51][52] Teleost fishes underwent a significant radiation, with otolith assemblages from turbiditic deposits in northern Italy revealing diverse families like Myctophidae (lanternfishes), which increased in size and ecological dominance in mid-water niches.[53][54] Nautiloids persisted as relict cephalopods in offshore settings, while ammonites remained absent following their end-Cretaceous extinction, highlighting the post-K/Pg recovery patterns among shelled mollusks.[55] Regional Mediterranean assemblages included pectinid bivalves in hardground biofacies, supporting infaunal communities. Evolutionary events included heightened productivity in upwelling zones, such as those off the Iberian Margin and in the Indian Ocean, where nutrient influx drove diatom blooms and phosphogenesis in benthic hardgrounds.[56][57] Localized anoxic events impacted benthic diversity, reducing foraminiferal and macrofaunal abundances in restricted basins while favoring opportunistic taxa in oxygenated refugia.[58][59] These dynamics contributed to overall marine ecosystem restructuring, with increased turnover among echinoids and mollusks (up to 85% in the Mediterranean).[7]

Terrestrial biota

During the Tortonian, terrestrial flora underwent significant transformations driven by global cooling and aridification, with the expansion of C4 grasslands becoming prominent in Africa and Asia, where these ecosystems began replacing C3-dominated woodlands around 10 million years ago.[60] In sub-Saharan Africa, C4 grasses, including members of the Panicoideae and Andropogoneae subfamilies, spread widely, forming savannas that supported diverse herbivore communities by the late Tortonian.[61] This shift was particularly evident in eastern and northwestern Africa, where isotopic evidence from soil carbonates and plant waxes indicates C4 dominance by approximately 7.2 Ma in regions like Anatolia and southern Asia, contemporaneous with similar developments elsewhere.[62] Concurrently, tropical rainforests in Southeast Asia flourished, characterized by dense stands of dipterocarp trees that had migrated from African origins via the Indian subcontinent earlier in the Cenozoic, creating multilayered canopies in humid lowlands.[63] In the Mediterranean region, sclerophyllous vegetation began to emerge more distinctly, with evergreen shrubs and small trees adapted to seasonal aridity, such as elements of the modern maquis, increasing in abundance as mesic forests declined.[64] Mammalian faunas diversified markedly on land during the Tortonian, with Old World monkeys (Cercopithecidae) undergoing adaptive radiations in forested and woodland habitats across Eurasia and Africa, adapting to folivorous and frugivorous diets.[65] Early hominoids, including the genus Dryopithecus in Europe, exemplified this trend, with fossils from sites like Rudabánya in Hungary dating to around 11-10 Ma, featuring suspensory locomotion suited to arboreal environments.[66] Proboscideans, particularly deinotheres such as Deinotherium, roamed open woodlands in Africa and Eurasia, using their downward-curving tusks for foraging on soft vegetation, persisting through much of the stage until the Messinian transition.[67] Rodent communities in Africa experienced a notable radiation, with cricetids like Afrocricetodon and early cane rats (Thryonomys) appearing in rift valley deposits, exploiting emerging grassland seeds and contributing to soil aeration in savanna ecosystems.[68] Other vertebrate groups reflected the heterogeneous terrestrial landscapes of the Tortonian. Reptiles, including varanid lizards (Varanus sp.), inhabited subtropical woodlands in the Southern Caucasus, preying on small vertebrates in semi-arid settings as evidenced by osteoderms and vertebrae from late Miocene localities.[69] Testudinid tortoises thrived in open, grassy habitats of the South Balkans, with species like those from Pikermian faunas adapting to herbivory in Mediterranean-like environments.[70] Among birds, passerines continued their diversification, with oscine lineages spreading into woodland edges and grasslands, as inferred from comparative phylogenies indicating Miocene radiations tied to insect abundance.[71] Amphibians occupied wetland margins and riverine habitats, such as those in the Caujarao Formation of Venezuela, where anurans and caudates utilized ephemeral ponds for breeding amid tropical vegetation.[68] Insects and other invertebrates played crucial ecological roles, with complex termite and ant societies facilitating nutrient cycling in expanding grasslands, thereby supporting herbivore populations through decomposed organic matter. Eusocial hymenopterans, including attine ants that domesticated fungi for agriculture around 8-12 Ma, enhanced plant decomposition in Neotropical analogs, paralleling Old World trends where such societies influenced ungulate foraging efficiency.[72] Regional faunas, like the Hemphillian of North America, featured specialized mammals such as teleoceratine rhinoceroses (Teleoceras), which grazed in floodplain grasslands, exemplifying local adaptations to open habitats.[73] Key evolutionary trends included the proliferation of open habitats, which spurred ungulate migrations across Eurasia and into Africa, as aridification fragmented forests and created migratory corridors for grazers like equids and bovids.[74] In Africa, mammal exchanges intensified via rift valleys of the East African Plateau, allowing faunal mixing between northern savannas and southern woodlands, as tectonic uplift provided dispersal routes amid climatic shifts favoring grasslands.[75] These dynamics, briefly linked to broader cooling that promoted C4 photosynthesis, underscored the Tortonian as a pivotal interval for continental biome restructuring.[62]

References

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