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Carboniferous rainforest collapse

March 05, 2023

The Carboniferous rainforest collapse (CRC) was a minor extinction event that occurred around 305 million years ago in the Carboniferous period.[1] It altered the vast coal forests that covered the equatorial region of Euramerica (Europe and America). This event may have fragmented the forests into isolated refugia or ecological 'islands', which in turn encouraged dwarfism and, shortly after, extinction of many plant and animal species. Following the event, coal-forming tropical forests continued in large areas of the Earth, but their extent and composition were changed.

Coal forests continued after the Carboniferous rainforest collapse. These plant fossils are from one of those forests from about 5 million years after the CRC. However, the composition of the forests changed from a lepidodendron-dominated forest to one of predominantly tree ferns and seed ferns.

The event occurred at the end of the Moscovian and continued into the early Kasimovian stages of the Pennsylvanian (Upper Carboniferous).

Extinction patterns on land

In the Carboniferous, the great tropical rainforests of Euramerica supported towering lycopodiophyta, a heterogeneous mix of vegetation, as well as a great diversity of animal life: giant dragonflies, millipedes, blattopterans, and smaller amphibians and the first amniotes.

Plants

The rise of rainforests in the Carboniferous greatly altered the landscapes by eroding low-energy, organic-rich anastomosing (braided) river systems with multiple channels and stable alluvial islands. The continuing evolution of tree-like plants increased floodplain stability (less erosion and movement) by the density of floodplain forests, the production of woody debris, and an increase in complexity and diversity of root assemblages.[2]

Collapse occurred through a series of step changes. First there was a gradual rise in the frequency of opportunistic ferns in late Moscovian times.[3] This was followed in the earliest Kasimovian by a major, abrupt extinction of the dominant lycopsids and a change to tree fern-dominated ecosystems.[4] This is confirmed by a recent study showing that the presence of meandering and anabranching rivers, occurrences of large woody debris, and records of log jams decrease significantly at the Moscovian-Kasimovian boundary.[2] Rainforests were fragmented, forming shrinking 'islands' further and further apart, and in latest Kasimovian time, rainforests vanished from the fossil record.

Invertebrates

The fossil record of insects can be difficult to study to the generally smaller and more delicate nature of their bodies. One study tabulate the rates of origination and extinction of over 600 terrestrial and freshwater animal families. Their stratigraphic ranges spanned a geologic interval from the middle Paleozoic biotic invasion of the land to the Permian crisis. Insects comprise more than half of the sampled families, most of which are from tropical Euramerica. This study found a Late Pennsylvanian extinction pulse that reflects drying climates the transition of lycopod to tree fern-dominated land floras.[5]

Vertebrates

Before the collapse, vertebrate animal species distribution was very cosmopolitan—the same species existed everywhere across tropical Pangaea—but after the collapse, each surviving rainforest 'island' developed its own unique mix of species. Many amphibian species became extinct, while the ancestors of reptiles and mammals diversified into more species after the initial crisis.[1] These patterns are explained by the theory of insular biogeography, a concept that explains how evolution progresses when populations are restricted into isolated pockets. This theory was originally developed for oceanic islands, but it can be applied equally well to any other ecosystem that is fragmented, only existing in small patches and surrounded by another unsuitable habitat. According to this theory, the initial impact of habitat fragmentation is devastating, with most life dying out quickly from lack of resources. Then, as surviving plants and animals reestablish themselves, they adapt to their restricted environment to take advantage of the new allotment of resources, and diversify. After the Carboniferous Rainforest Collapse, each pocket of life evolved in its own way, resulting in a unique species mix that ecologists call "endemism". A 2018 paper challenged this theory, however, finding evidence for increased cosmopolitanism rather than endemism following the demise of Carboniferous rainforests.[6]

Biotic recovery and evolutionary consequences

Plants

The fragmentation of wetlands left a few isolated refugia in Europe. However, even these were unable to maintain the diversity of Moscovian flora.[7] By the Asselian many families of seed ferns that characterized the Moscovian tropical wetlands had disappeared including Flemingitaceae, Diaphorodendraceae, Tedeleaceae, Urnatopteridaceae, Alethopteridaceae,[verification needed] Cyclopteridaceae, and Neurodontopteridaceae.[7]

Invertebrates

Carboniferous rainforest collapse is sometimes treated as an extinction factor for large Carboniferous arthropods such as giant griffinfly Meganeura and millipede Arthropleura. It is common theory that high oxygen levels have led to larger arthropods, and these organisms have been thought to live in forests. It was said that rainforest collapse led to a decrease in oxygen concentration and a decrease in the habitat of these arthropods, leading them to extinction.[8] However, later study shows that both griffinflies and Arthropleura more likely lived a forest-independent life, and fossil records of both large griffinflies and Arthropleura are known after rainforest collapse.[9][10][11] This means that rainforest collapse, reduced oxygen levels were less involved in their extinction.

Vertebrates

 
Terrestrially adapted synapsids, the predecessors of the mammal lineage, like Archaeothyris were among the groups who quickly recovered after the collapse.

This sudden collapse affected several large groups. Labyrinthodont amphibians were particularly devastated, while the amniotes (the first members of the sauropsid and synapsid groups) fared better, being physiologically better adapted to the drier conditions.[1]

Amphibians can survive cold conditions by decreasing metabolic rates and resorting to overwintering strategies (i.e. spending most of the year inactive in burrows or under logs). However, this is not an effective way to deal with prolonged unfavourable conditions, especially desiccation. Amphibians must return to water to lay eggs, while amniotes have eggs that have a membrane that retains water and allows gas exchange out of water. Because amphibians had a limited capacity to adapt to the drier conditions that dominated Permian environments, many amphibian families failed to occupy new ecological niches and became extinct.[12] Amphibians also removed the scales of their aquatic ancestors, and breathed with both lungs and skin (as long as the skin was kept wet). But amniotes re-evolved scales, now more keratinized, allowing them to conserve water but losing their cutaneous respiration.

Synapsids and sauropsids acquired new niches faster than amphibians, and new feeding strategies, including herbivory and carnivory, previously only having been insectivores and piscivores.[1] Synapsids in particular became substantially larger than before and this trend would continue until the Permian-Triassic extinction event, after which their cynodont (mammal ancestors) descendants became smaller and nocturnal.

Possible causes

Atmosphere and climate

There are several hypotheses about the nature and cause of the Carboniferous Rainforest Collapse, some of which include climate change.[13][14][15] After a late Bashkirian interval of glaciation, high-frequency shifts in seasonality from humid to arid times began.[16]

The Carboniferous period is characterised by the formation of coal deposits which were formed within a context of the removal of atmospheric carbon. In the latest Middle Pennsylvanian (late Moscovian) a cycle of aridification began. At the time of the Carboniferous rainforest collapse, the climate became cooler and drier. This is reflected in the rock record as the Earth entered a short, intense ice age. Sea levels dropped by about 100 metres (330 ft), and glacial ice covered most of the southern continent of Gondwana.[17] The climate was unfavourable to rainforests and much of the biodiversity in them. Rainforests shrank into isolated patches mostly confined to wet valleys further and further apart. Little of the original lycopsid rainforest biome survived this initial climate crisis. The concentration of carbon dioxide in the atmosphere crashed to one of its all time global lows in the Pennsylvanian and early Permian.[16][17]

Then a succeeding period of global warming reversed the climatic trend; the remaining rainforests, unable to survive the rapidly changing conditions, were finally wiped out.[vague][citation needed]

As the climate aridified again through the later Paleozoic, rainforests were eventually replaced by seasonally dry biomes.[18] Though the exact speed and nature of the collapse is not clear, it is thought to have occurred relatively quickly in geologic terms, only a few thousand years at most.[citation needed]

Volcanism

After restoring the middle of the Skagerrak-Centered Large Igneous Province (SCLIP) using a new reference frame, it has been shown that the Skagerrak plume rose from the core–mantle boundary (CMB) to its ~300 Ma position.[19] The major eruption interval took place in very narrow time interval, of 297 Ma ± 4 Ma. The rift formation coincides with the Moskovian/Kasimovian boundary and the Carboniferous Rainforest Collapse.[20]

Geography

While the CRC effected the equatorial region of Euramerica, the collapse had no effect in the region of Cathaysia to the east (which mostly corresponds to modern China), where Carboniferous-like rainforests would persist until the very end of the Permian, around 252 million years ago.

Climate and geology

A paleoclimate change of global nature occurred during the Moscovian and Kasimovian. An atmospheric drying (aridification) happened in the Middle to Late Pennsylvanian, coinciding with abrupt faunal changes in marine and terrestrial species.[21] This change was recorded in paleosols, which reflect a period of overall decreased hydromorphy, increased free-drainage and landscape stability, and a shift in the overall regional climate to drier conditions in the Upper Pennsylvanian (Missourian). This is consistent with climate interpretations based on contemporaneous paleo-floral assemblages and geological evidence.[21][22][23]

Fossil sites

 
Fossil lycopsid, probably Sigillaria, from Joggins, with attached stigmarian roots

Many fossil sites around the world reflect the changing conditions of the Carboniferous Rainforest Collapse.

The Joggins Fossil Cliffs on Nova Scotia's Bay of Fundy, a UNESCO World Heritage Site, is a particularly well-preserved fossil site. Fossil skeletons embedded in the crumbling sea cliffs were discovered by Sir Charles Lyell in 1852. In 1859, his colleague William Dawson discovered the oldest known reptile-ancestor, Hylonomus lyelli, and since then hundreds more skeletons have been found, including the oldest synapsid, Protoclepsydrops.[24]

References

  1. ^ a b c d Sahney, S., Benton, M.J. & Falcon-Lang, H.J. (2010). "Rainforest collapse triggered Pennsylvanian tetrapod diversification in Euramerica" (PDF). Geology. 38 (12): 1079–1082. Bibcode:2010Geo....38.1079S. doi:10.1130/G31182.1.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  2. ^ a b Davies, N.S.; Gibling, M. R. (2011). "Evolution of fixed-channel alluvial plains in response to Carboniferous vegetation". Nature Geoscience. 21 (9): 629–633. Bibcode:2011NatGe...4..629D. doi:10.1038/ngeo1237.
  3. ^ Pfefferkorn, H.W.; Thomson, M.C. (1982). "Changes in dominance patterns in Upper Carboniferous plant fossil assemblages". Geology. 10 (12): 641. Bibcode:1982Geo....10..641P. doi:10.1130/0091-7613(1982)10<641:CIDPIU>2.0.CO;2.
  4. ^ DiMichele, W.A.; Phillips, T.L. (1996). "Climate change, plant extinctions and vegetational recovery during the Middle-Late Pennsylvanian transition: The case of tropical peat-forming environments in North America". Biotic Recovery from Mass Extinction Events: Geological Society of London Special Publication. 102 (1): 201–221. Bibcode:1996GSLSP.102..201D. doi:10.1144/GSL.SP.1996.001.01.14. S2CID 53419701.
  5. ^ Rivera, Alexei A. (2017-07-16). "Sluggish Rates of Evolution for Land Animals During the Late Paleozoic Ice Age: A Geobiological Interpretation". The Paleontological Society Special Publications. 13: 91–91. doi:10.1017/S2475262200011965. ISSN 2475-2622.
  6. ^ Dunne, Emma M.; Close, Roger A.; Button, David J.; Brocklehurst, Neil; Cashmore, Daniel D.; Lloyd, Graeme T.; Butler, Richard J. (7 February 2018). "Diversity change during the rise of tetrapods and the impact of the 'Carboniferous rainforest collapse'". Proceedings of the Royal Society B. 285 (1872): 1–8. doi:10.1098/rspb.2017.2730. Retrieved 31 December 2022.
  7. ^ a b Borja Cascales-Miñana; Christopher J. Cleal (2013). "The plant fossil record reflects just two great extinction events". Terra Nova. 26 (3): 195–200. Bibcode:2014TeNov..26..195C. doi:10.1111/ter.12086.
  8. ^ Holmes, Thom (2008). March Onto Land: The Silurian Period to the Middle Triassic Epoch. Infobase Publishing. ISBN 978-0-8160-5959-1.
  9. ^ Nel, André; Prokop, Jakub; Pecharová, Martina; Engel, Michael S.; Garrouste, Romain (2018-08-14). "Palaeozoic giant dragonflies were hawker predators". Scientific Reports. 8 (1): 12141. Bibcode:2018NatSR...812141N. doi:10.1038/s41598-018-30629-w. ISSN 2045-2322. PMC 6092361. PMID 30108284.
  10. ^ Nel, Andre; Fleck, Gunther; GARROUSTE, Romain; Gand, Georges (2008-08-10). "The Odonatoptera of the Late Permian Lodève Basin (Insecta)". Journal of Iberian Geology.
  11. ^ Davies, Neil S.; Garwood, Russell J.; McMahon, William J.; Schneider, Joerg W.; Shillito, Anthony P. (2022-05-01). "The largest arthropod in Earth history: insights from newly discovered Arthropleura remains (Serpukhovian Stainmore Formation, Northumberland, England)". Journal of the Geological Society. 179 (3). Bibcode:2022JGSoc.179..115D. doi:10.1144/jgs2021-115. ISSN 0016-7649. S2CID 245401499.
  12. ^ Miguel Á. Olalla-Tárraga1, Lynsey McInnes, Luis M. Bini, José A. F. Diniz-Filho, Susanne A. Fritz, Bradford A. Hawkins, Joaquín Hortal, C. David L. Orme1, Carsten Rahbek, Miguel Á. Rodríguez, Andy Purvis (2010). "Climatic niche conservatism and the evolutionary dynamics in species range boundaries: global congruence across mammals and amphibians". Journal of Biogeography. 38 (12): 2237–2247. doi:10.1111/j.1365-2699.2011.02570.x. S2CID 73556763.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  13. ^ Fielding, C.R.; Frank, T.D.; Birgenheier, L.P.; Rygel, M.C.; Jones, A.T. & Roberts, J. (2008). "Stratigraphic imprint of the Late Palaeozoic Ice Age in eastern Australia: A record of alternating glacial and nonglacial climate regime". Geological Society of London Journal. 165 (1): 129–140. Bibcode:2008JGSoc.165..129F. doi:10.1144/0016-76492007-036. S2CID 31953303.
  14. ^ Heckel, P.H. (1991). "Lost Branch Formation and revision of upper Desmoinesian stratigraphy along midcontinent Pennsylvanian outcrop belt". Geological Survey Geology Series. 4.
  15. ^ DiMichele, W.A.; Cecil, B.; Montanez, I.P. & Falcon-Lang, H.J. (2010). "Cyclic changes in Pennsylvanian paleoclimate and effects on floristic dynamics in tropical Pangaea". International Journal of Coal Geology. 83 (2–3): 329–344. doi:10.1016/j.coal.2010.01.007. S2CID 64358884.
  16. ^ a b Gulbransona, Montañezb; Taborc, Limarinod (2014). "Late Pennsylvanian aridification on the southwestern margin of Gondwana (Paganzo Basin, NW Argentina): A regional expression of a global climate perturbation". Palaeogeography, Palaeoclimatology, Palaeoecology. 417: 220–235. Bibcode:2015PPP...417..220G. doi:10.1016/j.palaeo.2014.10.029.
  17. ^ a b Polly, D.P. (2011). "The Carboniferous Crisis" (PDF). {{cite journal}}: Cite journal requires |journal= (help)
  18. ^ Montañez, I.P.; Tabor, N.J.; Niemeier, D.; DiMichele, W.A.; Frank, T.D.; Fielding, C.R.; Isbell, J.L.; Birgenheier, L.P. & Rygel, M.C. (2007). "CO2-forced climate and vegetation instability during late Paleozoic deglaciation". Science. 315 (5808): 87–91. Bibcode:2007Sci...315...87M. doi:10.1126/science.1134207. PMID 17204648. S2CID 5757323.
  19. ^ T.H. Torsvik; M.A. Smethurst; K. Burke; B. Steinberger (2008). "Long term stability in deep mantle structure: evidence from the 300 Ma Skagerrak-Centered Large Igneous Province (the SCLIP)". Earth and Planetary Science Letters. 267 (3–4): 444–452. Bibcode:2008E&PSL.267..444T. doi:10.1016/j.epsl.2007.12.004.
  20. ^ Vadim A. Kravchinsky (2012). "Paleozoic large igneous provinces of Northern Eurasia: Correlation with mass extinction events". Global and Planetary Change. 86–87: 31–36. Bibcode:2012GPC....86...31K. doi:10.1016/j.gloplacha.2012.01.007.
  21. ^ a b Gulbranson, EL; Montanez, IP; Tabor, NJ; Limarino, CO (2015). "Late Pennsylvanian aridification on the southwestern margin of Gondwana (Paganzo Basin, NW Argentina): A regional expression of a global climate perturbation". Palaeogeography, Palaeoclimatology, Palaeoecology. 417: 220–235. Bibcode:2015PPP...417..220G. doi:10.1016/j.palaeo.2014.10.029.
  22. ^ Rosenau, Nicholasd; Neil J. Tabor (2013). "Oxygen and hydrogen isotope composition of paleosol phyllosilicates: Differential burial histories and determination of Middle–Late Pennsylvanian low-latitude terrestrial paleotemperatures". Palaeogeography, Palaeoclimatology, Palaeoecology. 392: 382–397. Bibcode:2013PPP...392..382R. doi:10.1016/j.palaeo.2013.09.020.
  23. ^ Rosenau, Nicholas; Tabor, Neil J.; Elrick, Scott D.; Nelson, W. John (2013). "Polygenetic History of Paleosols In Middle–Upper Pennsylvanian Cyclothems of the Illinois Basin, U.S.A.: Part II. Integrating Geomorphology, Climate, and Glacioeustasy". Journal of Sedimentary Research. 83 (8): 637–668. Bibcode:2013JSedR..83..637R. doi:10.2110/jsr.2013.51. See "Dinosaur Extinction" chapter.
  24. ^ Falcon-Lang, H. J., Benton, M.J., Braddy, S. J. and Davies, S.J. (2006). "The Pennsylvanian tropical biome reconstructed from the Joggins Formation of Nova Scotia, Canada". Journal of the Geological Society, London. 163 (3): 561–576. Bibcode:2006JGSoc.163..561F. doi:10.1144/0016-764905-063. S2CID 129791363.{{cite journal}}: CS1 maint: multiple names: authors list (link)

Further reading

  • Polly, David (2011). (PDF). Department of Geological Sciences, Indiana University. Archived from the original (PDF) on 2012-01-11. Retrieved 2011-09-04.
  • Rincon, Paul (November 2010). "Rainforest collapse kickstarted reptile evolution". BBC News. Retrieved 30 June 2019.
  • Mirsky, Steve (November 2010). "Ancient Rainforest Collapse Increased Reptile Diversity". Scientific American Podcast. Retrieved 30 June 2019.
  • Falcon-Lang, Howard (December 2010). "Brave new reptilian world". Planet Earth Online. Retrieved 30 June 2019.
  • . Palaeobiology and Biodiversity Research Group, Department of Earth Sciences, University of Bristol. April 2011. Archived from the original on 25 March 2017. Retrieved 30 June 2019.
  • Pritchard, Hamish (August 2011). "Early forests tamed wild rivers". BBC News. Retrieved 30 June 2019.

March 05, 2023
carboniferous, rainforest, collapse, minor, extinction, event, that, occurred, around, million, years, carboniferous, period, altered, vast, coal, forests, that, covered, equatorial, region, euramerica, europe, america, this, event, have, fragmented, forests, . The Carboniferous rainforest collapse CRC was a minor extinction event that occurred around 305 million years ago in the Carboniferous period 1 It altered the vast coal forests that covered the equatorial region of Euramerica Europe and America This event may have fragmented the forests into isolated refugia or ecological islands which in turn encouraged dwarfism and shortly after extinction of many plant and animal species Following the event coal forming tropical forests continued in large areas of the Earth but their extent and composition were changed Coal forests continued after the Carboniferous rainforest collapse These plant fossils are from one of those forests from about 5 million years after the CRC However the composition of the forests changed from a lepidodendron dominated forest to one of predominantly tree ferns and seed ferns The event occurred at the end of the Moscovian and continued into the early Kasimovian stages of the Pennsylvanian Upper Carboniferous Contents 1 Extinction patterns on land 1 1 Plants 1 2 Invertebrates 1 3 Vertebrates 2 Biotic recovery and evolutionary consequences 2 1 Plants 2 2 Invertebrates 2 3 Vertebrates 3 Possible causes 3 1 Atmosphere and climate 3 2 Volcanism 4 Geography 5 Climate and geology 6 Fossil sites 7 References 8 Further readingExtinction patterns on land EditIn the Carboniferous the great tropical rainforests of Euramerica supported towering lycopodiophyta a heterogeneous mix of vegetation as well as a great diversity of animal life giant dragonflies millipedes blattopterans and smaller amphibians and the first amniotes Plants Edit The rise of rainforests in the Carboniferous greatly altered the landscapes by eroding low energy organic rich anastomosing braided river systems with multiple channels and stable alluvial islands The continuing evolution of tree like plants increased floodplain stability less erosion and movement by the density of floodplain forests the production of woody debris and an increase in complexity and diversity of root assemblages 2 Collapse occurred through a series of step changes First there was a gradual rise in the frequency of opportunistic ferns in late Moscovian times 3 This was followed in the earliest Kasimovian by a major abrupt extinction of the dominant lycopsids and a change to tree fern dominated ecosystems 4 This is confirmed by a recent study showing that the presence of meandering and anabranching rivers occurrences of large woody debris and records of log jams decrease significantly at the Moscovian Kasimovian boundary 2 Rainforests were fragmented forming shrinking islands further and further apart and in latest Kasimovian time rainforests vanished from the fossil record Invertebrates Edit The fossil record of insects can be difficult to study to the generally smaller and more delicate nature of their bodies One study tabulate the rates of origination and extinction of over 600 terrestrial and freshwater animal families Their stratigraphic ranges spanned a geologic interval from the middle Paleozoic biotic invasion of the land to the Permian crisis Insects comprise more than half of the sampled families most of which are from tropical Euramerica This study found a Late Pennsylvanian extinction pulse that reflects drying climates the transition of lycopod to tree fern dominated land floras 5 Vertebrates Edit Before the collapse vertebrate animal species distribution was very cosmopolitan the same species existed everywhere across tropical Pangaea but after the collapse each surviving rainforest island developed its own unique mix of species Many amphibian species became extinct while the ancestors of reptiles and mammals diversified into more species after the initial crisis 1 These patterns are explained by the theory of insular biogeography a concept that explains how evolution progresses when populations are restricted into isolated pockets This theory was originally developed for oceanic islands but it can be applied equally well to any other ecosystem that is fragmented only existing in small patches and surrounded by another unsuitable habitat According to this theory the initial impact of habitat fragmentation is devastating with most life dying out quickly from lack of resources Then as surviving plants and animals reestablish themselves they adapt to their restricted environment to take advantage of the new allotment of resources and diversify After the Carboniferous Rainforest Collapse each pocket of life evolved in its own way resulting in a unique species mix that ecologists call endemism A 2018 paper challenged this theory however finding evidence for increased cosmopolitanism rather than endemism following the demise of Carboniferous rainforests 6 Biotic recovery and evolutionary consequences EditPlants Edit The fragmentation of wetlands left a few isolated refugia in Europe However even these were unable to maintain the diversity of Moscovian flora 7 By the Asselian many families of seed ferns that characterized the Moscovian tropical wetlands had disappeared including Flemingitaceae Diaphorodendraceae Tedeleaceae Urnatopteridaceae Alethopteridaceae verification needed Cyclopteridaceae and Neurodontopteridaceae 7 Invertebrates Edit Carboniferous rainforest collapse is sometimes treated as an extinction factor for large Carboniferous arthropods such as giant griffinfly Meganeura and millipede Arthropleura It is common theory that high oxygen levels have led to larger arthropods and these organisms have been thought to live in forests It was said that rainforest collapse led to a decrease in oxygen concentration and a decrease in the habitat of these arthropods leading them to extinction 8 However later study shows that both griffinflies and Arthropleura more likely lived a forest independent life and fossil records of both large griffinflies and Arthropleura are known after rainforest collapse 9 10 11 This means that rainforest collapse reduced oxygen levels were less involved in their extinction Vertebrates Edit Terrestrially adapted synapsids the predecessors of the mammal lineage like Archaeothyris were among the groups who quickly recovered after the collapse This sudden collapse affected several large groups Labyrinthodont amphibians were particularly devastated while the amniotes the first members of the sauropsid and synapsid groups fared better being physiologically better adapted to the drier conditions 1 Amphibians can survive cold conditions by decreasing metabolic rates and resorting to overwintering strategies i e spending most of the year inactive in burrows or under logs However this is not an effective way to deal with prolonged unfavourable conditions especially desiccation Amphibians must return to water to lay eggs while amniotes have eggs that have a membrane that retains water and allows gas exchange out of water Because amphibians had a limited capacity to adapt to the drier conditions that dominated Permian environments many amphibian families failed to occupy new ecological niches and became extinct 12 Amphibians also removed the scales of their aquatic ancestors and breathed with both lungs and skin as long as the skin was kept wet But amniotes re evolved scales now more keratinized allowing them to conserve water but losing their cutaneous respiration Synapsids and sauropsids acquired new niches faster than amphibians and new feeding strategies including herbivory and carnivory previously only having been insectivores and piscivores 1 Synapsids in particular became substantially larger than before and this trend would continue until the Permian Triassic extinction event after which their cynodont mammal ancestors descendants became smaller and nocturnal Possible causes EditAtmosphere and climate Edit There are several hypotheses about the nature and cause of the Carboniferous Rainforest Collapse some of which include climate change 13 14 15 After a late Bashkirian interval of glaciation high frequency shifts in seasonality from humid to arid times began 16 The Carboniferous period is characterised by the formation of coal deposits which were formed within a context of the removal of atmospheric carbon In the latest Middle Pennsylvanian late Moscovian a cycle of aridification began At the time of the Carboniferous rainforest collapse the climate became cooler and drier This is reflected in the rock record as the Earth entered a short intense ice age Sea levels dropped by about 100 metres 330 ft and glacial ice covered most of the southern continent of Gondwana 17 The climate was unfavourable to rainforests and much of the biodiversity in them Rainforests shrank into isolated patches mostly confined to wet valleys further and further apart Little of the original lycopsid rainforest biome survived this initial climate crisis The concentration of carbon dioxide in the atmosphere crashed to one of its all time global lows in the Pennsylvanian and early Permian 16 17 Then a succeeding period of global warming reversed the climatic trend the remaining rainforests unable to survive the rapidly changing conditions were finally wiped out vague citation needed As the climate aridified again through the later Paleozoic rainforests were eventually replaced by seasonally dry biomes 18 Though the exact speed and nature of the collapse is not clear it is thought to have occurred relatively quickly in geologic terms only a few thousand years at most citation needed Volcanism Edit After restoring the middle of the Skagerrak Centered Large Igneous Province SCLIP using a new reference frame it has been shown that the Skagerrak plume rose from the core mantle boundary CMB to its 300 Ma position 19 The major eruption interval took place in very narrow time interval of 297 Ma 4 Ma The rift formation coincides with the Moskovian Kasimovian boundary and the Carboniferous Rainforest Collapse 20 Geography EditWhile the CRC effected the equatorial region of Euramerica the collapse had no effect in the region of Cathaysia to the east which mostly corresponds to modern China where Carboniferous like rainforests would persist until the very end of the Permian around 252 million years ago Climate and geology EditA paleoclimate change of global nature occurred during the Moscovian and Kasimovian An atmospheric drying aridification happened in the Middle to Late Pennsylvanian coinciding with abrupt faunal changes in marine and terrestrial species 21 This change was recorded in paleosols which reflect a period of overall decreased hydromorphy increased free drainage and landscape stability and a shift in the overall regional climate to drier conditions in the Upper Pennsylvanian Missourian This is consistent with climate interpretations based on contemporaneous paleo floral assemblages and geological evidence 21 22 23 Fossil sites Edit Fossil lycopsid probably Sigillaria from Joggins with attached stigmarian roots Many fossil sites around the world reflect the changing conditions of the Carboniferous Rainforest Collapse Hamilton Kansas US Jarrow Tyne amp Wear UK former Linton Mine in Saline Township Jefferson County Ohio US Nyrany Czech Republic Joggins Nova Scotia CanadaThe Joggins Fossil Cliffs on Nova Scotia s Bay of Fundy a UNESCO World Heritage Site is a particularly well preserved fossil site Fossil skeletons embedded in the crumbling sea cliffs were discovered by Sir Charles Lyell in 1852 In 1859 his colleague William Dawson discovered the oldest known reptile ancestor Hylonomus lyelli and since then hundreds more skeletons have been found including the oldest synapsid Protoclepsydrops 24 References Edit a b c d Sahney S Benton M J amp Falcon Lang H J 2010 Rainforest collapse triggered Pennsylvanian tetrapod diversification in Euramerica PDF Geology 38 12 1079 1082 Bibcode 2010Geo 38 1079S doi 10 1130 G31182 1 a href Template Cite journal html title Template Cite journal cite journal a CS1 maint multiple names authors list link a b Davies N S Gibling M R 2011 Evolution of fixed channel alluvial plains in response to Carboniferous vegetation Nature Geoscience 21 9 629 633 Bibcode 2011NatGe 4 629D doi 10 1038 ngeo1237 Pfefferkorn H W Thomson M C 1982 Changes in dominance patterns in Upper Carboniferous plant fossil assemblages Geology 10 12 641 Bibcode 1982Geo 10 641P doi 10 1130 0091 7613 1982 10 lt 641 CIDPIU gt 2 0 CO 2 DiMichele W A Phillips T L 1996 Climate change plant extinctions and vegetational recovery during the Middle Late Pennsylvanian transition The case of tropical peat forming environments in North America Biotic Recovery from Mass Extinction Events Geological Society of London Special Publication 102 1 201 221 Bibcode 1996GSLSP 102 201D doi 10 1144 GSL SP 1996 001 01 14 S2CID 53419701 Rivera Alexei A 2017 07 16 Sluggish Rates of Evolution for Land Animals During the Late Paleozoic Ice Age A Geobiological Interpretation The Paleontological Society Special Publications 13 91 91 doi 10 1017 S2475262200011965 ISSN 2475 2622 Dunne Emma M Close Roger A Button David J Brocklehurst Neil Cashmore Daniel D Lloyd Graeme T Butler Richard J 7 February 2018 Diversity change during the rise of tetrapods and the impact of the Carboniferous rainforest collapse Proceedings of the Royal Society B 285 1872 1 8 doi 10 1098 rspb 2017 2730 Retrieved 31 December 2022 a b Borja Cascales Minana Christopher J Cleal 2013 The plant fossil record reflects just two great extinction events Terra Nova 26 3 195 200 Bibcode 2014TeNov 26 195C doi 10 1111 ter 12086 Holmes Thom 2008 March Onto Land The Silurian Period to the Middle Triassic Epoch Infobase Publishing ISBN 978 0 8160 5959 1 Nel Andre Prokop Jakub Pecharova Martina Engel Michael S Garrouste Romain 2018 08 14 Palaeozoic giant dragonflies were hawker predators Scientific Reports 8 1 12141 Bibcode 2018NatSR 812141N doi 10 1038 s41598 018 30629 w ISSN 2045 2322 PMC 6092361 PMID 30108284 Nel Andre Fleck Gunther GARROUSTE Romain Gand Georges 2008 08 10 The 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Frank T D Birgenheier L P Rygel M C Jones A T amp Roberts J 2008 Stratigraphic imprint of the Late Palaeozoic Ice Age in eastern Australia A record of alternating glacial and nonglacial climate regime Geological Society of London Journal 165 1 129 140 Bibcode 2008JGSoc 165 129F doi 10 1144 0016 76492007 036 S2CID 31953303 Heckel P H 1991 Lost Branch Formation and revision of upper Desmoinesian stratigraphy along midcontinent Pennsylvanian outcrop belt Geological Survey Geology Series 4 DiMichele W A Cecil B Montanez I P amp Falcon Lang H J 2010 Cyclic changes in Pennsylvanian paleoclimate and effects on floristic dynamics in tropical Pangaea International Journal of Coal Geology 83 2 3 329 344 doi 10 1016 j coal 2010 01 007 S2CID 64358884 a b Gulbransona Montanezb Taborc Limarinod 2014 Late Pennsylvanian aridification on the southwestern margin of Gondwana Paganzo Basin NW Argentina A regional expression of a global climate perturbation Palaeogeography Palaeoclimatology Palaeoecology 417 220 235 Bibcode 2015PPP 417 220G doi 10 1016 j palaeo 2014 10 029 a b Polly D P 2011 The Carboniferous Crisis PDF a href Template Cite journal html title Template Cite journal cite journal a Cite journal requires journal help Montanez I P Tabor N J Niemeier D DiMichele W A Frank T D Fielding C R Isbell J L Birgenheier L P amp Rygel M C 2007 CO2 forced climate and vegetation instability during late Paleozoic deglaciation Science 315 5808 87 91 Bibcode 2007Sci 315 87M doi 10 1126 science 1134207 PMID 17204648 S2CID 5757323 T H Torsvik M A Smethurst K Burke B Steinberger 2008 Long term stability in deep mantle structure evidence from the 300 Ma Skagerrak Centered Large Igneous Province the SCLIP Earth and Planetary Science Letters 267 3 4 444 452 Bibcode 2008E amp PSL 267 444T doi 10 1016 j epsl 2007 12 004 Vadim A Kravchinsky 2012 Paleozoic large igneous provinces of Northern Eurasia Correlation with mass extinction events Global and Planetary Change 86 87 31 36 Bibcode 2012GPC 86 31K doi 10 1016 j gloplacha 2012 01 007 a b Gulbranson EL Montanez IP Tabor NJ Limarino CO 2015 Late Pennsylvanian aridification on the southwestern margin of Gondwana Paganzo Basin NW Argentina A regional expression of a global climate perturbation Palaeogeography Palaeoclimatology Palaeoecology 417 220 235 Bibcode 2015PPP 417 220G doi 10 1016 j palaeo 2014 10 029 Rosenau Nicholasd Neil J Tabor 2013 Oxygen and hydrogen isotope composition of paleosol phyllosilicates Differential burial histories and determination of Middle Late Pennsylvanian low latitude terrestrial paleotemperatures Palaeogeography Palaeoclimatology Palaeoecology 392 382 397 Bibcode 2013PPP 392 382R doi 10 1016 j palaeo 2013 09 020 Rosenau Nicholas Tabor Neil J Elrick Scott D Nelson W John 2013 Polygenetic History of Paleosols In Middle Upper Pennsylvanian Cyclothems of the Illinois Basin U S A Part II Integrating Geomorphology Climate and Glacioeustasy Journal of Sedimentary Research 83 8 637 668 Bibcode 2013JSedR 83 637R doi 10 2110 jsr 2013 51 See Dinosaur Extinction chapter Falcon Lang H J Benton M J Braddy S J and Davies S J 2006 The Pennsylvanian tropical biome reconstructed from the Joggins Formation of Nova Scotia Canada Journal of the Geological Society London 163 3 561 576 Bibcode 2006JGSoc 163 561F doi 10 1144 0016 764905 063 S2CID 129791363 a href Template Cite journal html title Template Cite journal cite journal a CS1 maint multiple names authors list link Further reading EditPolly David 2011 The Carboniferous Crises PDF Department of Geological Sciences Indiana University Archived from the original PDF on 2012 01 11 Retrieved 2011 09 04 Rincon Paul November 2010 Rainforest collapse kickstarted reptile evolution BBC News Retrieved 30 June 2019 Mirsky Steve November 2010 Ancient Rainforest Collapse Increased Reptile Diversity Scientific American Podcast Retrieved 30 June 2019 Falcon Lang Howard December 2010 Brave new reptilian world Planet Earth Online Retrieved 30 June 2019 Carboniferous climates and amniote origins Palaeobiology and Biodiversity Research Group Department of Earth Sciences University of Bristol April 2011 Archived from the original on 25 March 2017 Retrieved 30 June 2019 Pritchard Hamish August 2011 Early forests tamed wild rivers BBC News Retrieved 30 June 2019 Retrieved from https en wikipedia org w index php title Carboniferous rainforest collapse amp oldid 1136441693, wikipedia, wiki, book, books, library,

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