Carboniferous rainforest collapse

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 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 'islands', which in turn caused 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.

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

Extinction patterns on land

Ferns and treeferns from Mount Field National Park, giving an impression of how a Carboniferous rainforest might have looked.

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, cockroaches, 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.

Animals

Before the collapse, 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 reptiles 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 can be applied equally to any other ecosystem that is fragmented, only existing in small patches, surrounded by another 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 ecologists term "endemism".

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.[5] By the Asselian many families that characterized the Moscovian tropical wetlands had disappeared including Flemingitaceae, Diaphorodendraceae, Tedeleaceae, Urnatopteridaceae, Alethopteridaceae, Cyclopteridaceae, Neurodontopteridaceae.[5]

Invertebrates

The depletion of the plant life contributed to the deteriorating levels of oxygen in the atmosphere, which had facilitated the enormous arthropods of the time. Due to the decreasing oxygen, these sizes could no longer be accommodated, and thus between this and the loss of habitat, the giant arthropods were wiped out in this event, most notably the giant dragonflies (Meganeura and millipedes (Arthropleura).

Vertebrates

Terrestrially adapted early mammal-like reptiles 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 first reptiles fared better, being physiologically better adapted to the drier conditions that followed. Amphibians must return to water to lay eggs; in contrast, reptiles – whose amniote eggs have a membrane ensuring gas exchange out of water and can therefore be laid on land – cound better exploit the new conditions. Reptiles acquired new niches at a faster rate than before the collapse and at a much faster rate than amphibians. They acquired new feeding strategies including herbivory and carnivory, previously only having been insectivores and piscivores.[1]

This extinction event had long term implications for the evolution of amphibians. Under prolonged cold conditions, amphibians can survive 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 only a short term strategy and not an effective way of dealing with prolonged unfavourable conditions, especially desiccation. Since 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.[6]

Possible causes

Climate and atmosphere

There are several hypotheses about the nature and cause of the Carboniferous Rainforest Collapse, some of which include climate change.[7][8][9] Following a late Bashkirian interval of glaciation, high-frequency shifts in seasonality from humid to arid times began.[10]

Beginning 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 into a short, intense ice age. Sea levels dropped by 100 m, and glacial ice covered most of the southern continent of Gondwana.[11] The cooler, drier climate conditions were not favourable to the growth of rainforests, and much of the biodiversity within them. Rainforests shrank into isolated patches, the islands of rainforest were 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.[10][11]

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.

As the climate aridified again through the later Paleozoic, the rainforests were eventually replaced by seasonally dry biomes.[12] 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.

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.[13] 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.[14]

Multiple causes

In recent years, scientists have put forth the idea that many of Earth's largest extinction events were due to multiple causes that coincided in time. Proponents suggest multiple causes because they see no single cause as sufficient in strength to cause the mass extinctions or believe that a single cause is likely to produce the taxonomic pattern of the extinction. Two of Earth's largest extinction events have been hypothesized to be multicausal in nature.

The cause of the Permo-Triassic extinction is unclear, and some authors have indicated that it may be best explained by a "Murder on the Orient Express Scenario" in which multiple causes contributed to a devastating impact on life. Possible causes supported by strong evidence include the large scale volcanism at the Siberian Traps, the release of noxious gases, global warming, and anoxia (oxygen depletion).[15]

David Archibald and David E. Fastovsky discussed a scenario combining possibly three major causes to the K-T extinction: volcanism, marine regression, and extraterrestrial impact, together wiping out the non-avian dinosaurs 66 million years ago.[16] The specific cause of the CRC is not known but certainly a multiple cause scenario is a possibility.

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.[17] 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.[17][18][19]

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, Hylonomus lyelli, and since then hundreds more skeletons have been found.[20]

References

  1. 1 2 3 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.
  2. 1 2 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.
  5. 1 2 Borja Cascales-Miñana; Christopher J. Cleal (2013). "The plant fossil record reflects just two great extinction events". Terra Nova. 26 (3): 195–200. doi:10.1111/ter.12086.
  6. ↑ 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.
  7. ↑ 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: 129–140.
  8. ↑ Heckel, P.H. (1991). "Lost Branch Formation and revision of upper Desmoinesian stratigraphy along midcontinent Pennsylvanian outcrop belt". Geological Survey Geology Series. 4.
  9. ↑ DiMichele, W.A.; Cecil, B.; Montanez, I.P. & Falcon-Lang, H.J. (2010). "Cyclic changes in Pennsylvanian paleoclimate and it 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.
  10. 1 2 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. doi:10.1016/j.palaeo.2014.10.029.
  11. 1 2 Polly, D.P (2011). "The Carboniferous Crisis". http://www.indiana.edu/~g404/Lectures/Lecture%201%20-%20Carboniferous%20Crisis%20and%20Introduction%20to%20Vertebrate%20Geobiology.pdf.
  12. ↑ 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.
  13. ↑ 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.
  14. ↑ 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.
  15. ↑ Benton M J (2005). When life nearly died: the greatest mass extinction of all time. London: Thames & Hudson. ISBN 0-500-28573-X.
  16. ↑ David, Archibald; David Fastovsky (2004). "Dinosaur Extinction" (PDF). In Weishampel David B; Dodson Peter; Osmólska Halszka. The Dinosauria (2nd ed.). Berkeley: University of California Press. pp. 672–684. ISBN 0-520-24209-2.
  17. 1 2 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.
  18. ↑ 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. doi:10.1016/j.palaeo.2013.09.020.
  19. ↑ 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.
  20. ↑ 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. doi:10.1144/0016-764905-063.

Further reading


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Millions of years before present

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