Hangenberg event - Knowledge

79: 931:. This study found that >45% of genera were lost during the Famennian, lowered to ~28% considering only “multiple interval” genera which appeared prior to the stage. The Famennian extinction(s) would be the eighth worst mass extinction by the latter metric. He also found that the percentage loss of “well-preserved” (hard tissue) marine genera in the last substage of the Famennian was around 21%, nearly as large as the rate in the last substage of the Frasnian. The Famennian-wide extinction rate for “multiple interval” marine animal families was around 16%. All of these estimates approached, but did not surpass, the end-Frasnian extinction, and the 1446:(ray-finned fish), which took over in diversity and relative abundance during the Early Carboniferous. These survivors were generally small and fast-breeding, resulting in a decrease in average vertebrate body size across the extinction. Even so, few Devonian chondrichthyan and actinopterygian species survived into the Carboniferous, indicating that these groups also experienced extinctions. Sharks that survived the extinction were greatly reduced in size; only sharks less than a meter long remained, and it would take 40 million years before they started to increase in size again. 939: 1119: 141: 1128: 435:) define the D–C boundary, but difficulty in finding reliable and universal index taxa has complicated study of the boundary in many areas. The sea level fluctuated during the upper crisis interval, as several minor regressions and transgressions continued to occur around the D–C boundary. Nevertheless, the general trend was sea level rise, with the melting of the glaciers which formed in the middle crisis interval. In the early Tournaisian, the crisis finally ends at the base of the 1316: 265: 1108: 1099: 78: 1305: 1260:, were strongly affected. The order Phacopida completely died out during the event. Deep-water phacopids were eradicated at the start of the crisis, while widespread shallow-water phacopids went extinct slightly later, alongside the cymaclymeniid ammonoids. Proetids were also hit hard, but several families in the group survived and rediversified quickly in the Tournaisian. 1505:“tetrapods” has helped to close in this gap, suggesting that the Hangenberg Event affected some vertebrates less severely than previously thought. Coprolitic evidence from early Tournaisian deposits in eastern Greenland has also supported the notion that tetrapods were not as affected by the Hangenberg Event as previously thought. 905:'s 1982 landmark paper on mass extinctions. However, late Famennian extinction rates were typically considered to be of lesser taxonomic severity than those in the Kellwasser Event, one of the “big five” mass extinctions. Depending on the method used, the Hangenberg Event typically falls between the fifth and tenth deadliest post- 970:. They found a significantly higher extinction rate, with 50% of marine genera lost during the event. This estimate would rank the end-Famennian extinction as the fourth-deadliest mass extinction, ahead of the end-Frasnian extinction. They also ranked the end-Famennian mass extinction as the seventh most 1528:

was abundant in most spore zones used to define the terrestrial ecosystems of the Famennian. The major marine extinction pulse of the Hangenberg Event occurred at the boundary between the LE and LN zones, the third- and second-to-last spore zones of the Devonian, respectively. Plants were unaffected

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Chemical analysis of cores in the Bakken shale suggest that while it was being formed, successive eras of higher sea level corresponded with euxinic (high in toxic hydrogen sulfide and low in oxygen) water in the shallow ocean basin, which could kill animals in the ocean and near the shoreline. As

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were moderately affected by the event, with different regions varying in the number of species lost. Pelagic conodonts had a total species extinction rate of about 40%, with some areas have a local rate as high as 72%. About 50% of neritic conodont species died out, with survivors characterized by

916:(1995) estimated that 20–23.7% of all families went extinct in the Famennian, with marine families at a proportion of 1.2–20.4%. About 27.4–28.6% of continental families appear to have died out, but the early and low-diversity nature of Devonian continental life makes this estimate very imprecise. 1370:

extinctions, and far exceeds the impact of the Kellwasser Event. Furthermore, because the vertebrate fossil record of the Famennian is sparse, many extinctions attributed to the Kellwasser event may have actually been caused by the Hangenberg event. Among vertebrates, 44% of high-level clades and

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genozone (UD VI-F), indicating that post-Devonian ammonoids were beginning to diversify after the main extinction pulse. A major marine regression occurred during the middle crisis interval, as indicated by the increased amount of erosion and river-supplied siliciclastic material. Some areas even

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Evidence such as glacial deposits in northern Brazil (near the Devonian South Pole) suggests widespread glaciation at the end of the Devonian, as a broad continental mass covered the polar region. The Hangenberg event has been associated with sea-level rise followed swiftly by glaciation-related

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were filled rather quickly, but these communities were exceptions. By comparison, the end-Frasnian extinction was ranked as the fourth most ecologically-severe mass extinction, and the Givetian crisis was ranked as the eighth. Even in areas with oxygenated seabeds, such as parts of Morocco, the

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mass extinctions, in terms of marine genera lost. Most estimates of proportional extinction have low resolution, only as fine as the stages in which the extinctions occur. This can lead to uncertainty in differentiating between the Hangenberg Event and other Famennian extinctions in broad-scale

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The Hangenberg Event can be recognized by its unique multi-phase sequence of sedimentary layers, representing a relatively short interval of time with extreme fluctuations in the climate, sea level, and diversity of life. The entire event had an estimated duration of 100,000 to several hundred

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maintained high rates of both speciation and extinction during the late Famennian, with only a small drop in overall richness. The Hangenberg event effectively "refreshed" bryozoan diversity, eliminating old clades and allowing new forms to radiate and eventually reach a diversity peak in the

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Using an updated biodiversity database, Bambach (2006) estimated that a total of 31% of marine genera died out in the last substage of the Famennian. By this metric, the Hangenberg Event was the joint seventh-worst post-Cambrian mass extinction, tied with the poorly-studied early

128:. This layer and its surrounding geological units define the "classic" Rhenish succession, one of the most well-studied geological examples of the extinction. Sequences equivalent to the Rhenish succession have been found at over 30 other sites on every continent except 1903:"Kaiser, S.I., Steuber, T., Becker, R.T. & Joachimski, M.M. 2006. Geochemical evidence for major environmental change at the Devonian–Carboniferous boundary in the Carnic Alps and the Rhenish Massif, Palaeogeography, Palaeoclimatology, Palaeoecology 240, 146–160" 1681:

near the Devonian–Carboniferous boundary has led some researchers to hypothesise a volcanic cause for the Hangenberg event. The activities of the Kola and Timan-Pechora magmatic provinces have been proposed as other hypothetical causes for the Hangenberg event.

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their wide distribution and versatile ecology. Species diversity rebounded soon afterwards, returning close to pre-extinction levels by the middle of the Tournaisian. The Hangenberg Event has also been implicated in the final extinction of several

347:(which would have been a tropical alpine environment). These are known to have been deposited within the LE and/or LN spore zones, which are difficult to distinguish outside of Europe. Less well-constrained glacial deposits have also been found in 1060:

reefs which were previously unscathed. Nevertheless, in the absence of pressures from metazoan communities, there was a brief resurgence of microbial carbonate in the early Tournaisian, a similar pattern to other mass extinctions.

164:(sea level fall). This may have been caused by a small glacial phase, but other evidence suggests a warm and wet climate at the time. The uppermost part of the Wocklum Limestone and the Drewer Sandstone occupy the LE 1268:

dying out. At least 50% of pelagic ostracod species went extinct, with some areas having extinction rates up to 66%. Shallow-water species were less affected, with newer taxa replacing older ones late in the crisis.

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floras, is correlated with the extinction of ‘survivor’ faunas in the latter part of the Hangenberg Event. Spore taxa that went extinct include specialized forms with divided spines (likely from an early form of

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Algeo TJ, Scheckler SE, Maynard JB (2001). "Effects of the middle to late Devonian spread of vascular land plants on weathering regimes, marine biotas, and global climate". In Gensel PG, Edwards D (eds.).

1056:) reefs had already been devastated by the Frasnian–Famennian event, and were still recovering during the Famennian. The end of the Famennian not only eliminated the metazoan reef community, but also many 1641:, an ozone-depleting compound. However, this mechanism has been criticized for its slow and weak effect on ozone concentrations, as well as its suspect rejection of volcanic influences. Alternatively, 1493:

disappearing from the fossil record. A distinct gap in time traditionally separated the Famennian “tetrapod” faunas from their successors in the Early Carboniferous. This fossil hiatus, known as “

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oceans flooded terrestrial basins, water would have entered areas with high nutrient levels, leading to an algal bloom, removing oxygen and then creating hydrogen sulfide as the algae decayed.

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Kaufmann, B.; Trapp, E.; Mezger, K. (2004). "The numerical age of the Upper Frasnian (Upper Devonian) Kellwasser horizons: A new U-Pb zircon date from Steinbruch Schmidt(Kellerwald, Germany)".

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Below the Hangenberg Event strata is the Wocklum Limestone, a pelagic unit rich in fossils (especially ammonoids). In some places the Wocklum Limestone grades into the Drewer Sandstone, a thin

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Becker, Ralph Thomas; Kaiser, Sandra Isabella; Aretz, Markus (2016). "Review of chrono-, litho- and biostratigraphy across the global Hangenberg Crisis and Devonian–Carboniferous Boundary".

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apparently experienced a major ecological turnover across the Devonian-Carboniferous boundary. The Hangenberg Event's impact on vertebrate evolution approaches “Big Five” events such as the

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in fossils, but these tests have yet to be published. Ozone depletion could just as easily be explained by an increase in greenhouse gas concentrations resulting from an intense period of

3123: 1237:(shallow-water) taxa is less clear-cut. Some neritic taxa expanded after the initial extinction pulse but died out at the end of the crisis with other members of the ‘survivor’ fauna. 47:

Period (roughly 358.9 ± 0.4 million years ago). It is usually considered the second-largest extinction in the Devonian Period, having occurred approximately 13 million years after the

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Global Events and Event Stratigraphy in the Phanerozoic: Results of the International Interdisciplinary Cooperation in the IGCP-Project 216 “Global Biological Events in Earth History”

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One hypothesis for the cause of the last pulse of the extinction notes the abundance of malformed plant spores at the Devonian-Carboniferous boundary. This could implicate increased

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provide dates of 358.97 ± 0.11 Ma and 358.89 ± 0.20 Ma below and above the black shale. This constrains the main marine extinction pulse to a duration of 50,000 to 190,000 years.

313: 2877:"Oldest known naiaditid bivalve from the high-latitude Late Devonian (Famennian) of South Africa offers clues to survival strategies following the Hangenberg mass extinction" 4182:

Fields, Brian D.; Melott, Adrian L.; Ellis, John; Ertel, Adrienne F.; Fry, Brian J.; Lieberman, Bruce S.; Liu, Zhenghai; Miller, Jesse A.; Thomas, Brian C. (2020-09-01).

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sea-level fall, and thus a cause of the extinctions may have been an episode of severe global cooling and glaciation at the end of the Famennian, marking the dawn of the

1661:. The spore malformations may not even be related to UV radiation in the first place, and could simply be a result of volcanism-related environmental pressures such as 1283:

also experienced very high extinction rates which devastated their formerly high diversity. Surviving forms were low in diversity and small in size, an example of the ‘

3659:"Coprolite diversity reveals a cryptic ecosystem in an early Tournaisian lake in East Greenland: Implications for ecosystem recovery after the end-Devonian extinction" 2254:

Algeo, T. J. (1998). "Terrestrial-marine teleconnections in the Devonian: links between the evolution of land plants, weathering processes, and marine anoxic events".

1583:, and has been suggested to have been linked to an increase in terrestrial plant cover. That would have led to increased nutrient supply in rivers and may have led to 4014:

Sandberg, C.A.; Morrow, J.R.; Ziegler, W. (2002). "Late Devonian sea-level changes, catastrophic events, and mass extinctions". In Koeberl, C.; MacLeod, K.G. (eds.).

399:, spans the Devonian–Carboniferous (D–C) boundary. Foraminifera reappear in the fossil record within the Stockum Limestone, forming the DFZ8 zone characterized by 205: 3391: 981:. This was justified by the fact that two whole communities within an ecological megaguild went extinct with no replacements. For the end-Famennian, these were 1974: 1221:

apparently took advantage of the Famennian glaciation to expand from polar regions towards the equator, sparking diversification in the Carboniferous tropics.

1182: 436: 297: 277: 4015: 1975:"Water column euxinia and wildfire evidence during deposition of the Upper Famennian Hangenberg event horizon from the Holy Cross Mountains (central Poland)" 491: 3189:"Five hundred million years of extinction and recovery: a phanerozoic survey of large-scale diversity patterns in fishes: EXTINCTION AND RECOVERY IN FISHES" 4034: 3911: 396: 214: 427:

trilobites, also die out at this time, making it the second largest extinction pulse of the Hangenberg Crisis. Conodont zones (usually characterized by

2088:"Climate-controlled mass extinctions, facies, and sea-level changes around the Devonian–Carboniferous boundary in the eastern Anti-Atlas (SE Morocco)" 3972: 3809: 3663: 1265: 1187: 420: 1371:

over 96% of species were lost during the Hangenberg Event, which occurred globally and did not discriminate between freshwater and marine species.

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zone (UD VI-E), an ammonoid genozone based on massive extinctions within the group, rather than new occurrences. This is also the case for the

232: 4266:"Mercury spikes as evidence of extended arc-volcanism around the Devonian–Carboniferous boundary in the South Tian Shan (southern Uzbekistan)" 2609: 1176:, were already suffering smaller extinctions just prior to the event. Although clymeniids survived the extinction event itself, they became a 1690:

An asteroid impact has been suggested as a possible cause of the Hangenberg event. However, most impact craters, such as the Hangenberg-aged

1218: 962:. 2013 attempted to tackle extinction rates via a new resampling protocol designed to counter biases in biodiversity estimates, such as the 443:

superficially similar to the pre-crisis Wocklum Limestone. The base of the Hangenberg Limestone is characterized by the first occurrence of

2660:"Late Devonian and Early Carboniferous alpha diversity, ecospace occupation, vertebrate assemblages and bio-events of southeastern Morocco" 2715:"Global microbial carbonate proliferation after the end-Devonian mass extinction: Mainly controlled by demise of skeletal bioconstructors" 1902: 897:

Along with the Givetian and Frasnian stages, the Famennian was qualitatively acknowledged as having elevated extinction rates as early as

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indicates the beginning of the VI spore zone shortly before the D–C boundary. ‘Survivor’ faunas of marine invertebrates, such as the last

4371: 2782:"Richness of Famennian–Tournaisian (Late Devonian-early Carboniferous) bryozoans in shallow areas of Palaeotethys and Palaeoasian oceans" 1194:) briefly expanded into a cosmopolitan ‘survivor’ fauna, but ultimately died out at the end of the crisis. Only one ammonoid family, the 1241:

survived relatively unscathed, and instead used the extinction as an opportunity to drastically increase their diversity and body size.

218:. However, in some areas the boundary between the LE and LN zones is unclear and possibly based on geography more than chronology. The 120:. It is named after the Hangenberg Black Shale, a distinctive layer of anoxic sediment originally found along the northern edge of the 1172:

were nearly wiped out by the Hangenberg Event, a fact noted very early in the study of the extinction. One major Famennian group, the

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Rakociński, Michał; Pisarzowska, Agnieszka; Corradini, Carlo; Narkiewicz, Katarzyna; Dubicka, Zofia; Abdiyev, Nuriddin (2021-03-11).

3935:"Famennian glaciation in the eastern side of Parnaíba Basin, Brazil: evidence of advance and retreat of glacier in Cabeças Formation" 3417:"Confirmation of Romer's Gap as a low oxygen interval constraining the timing of initial arthropod and vertebrate terrestrialization" 3337:"Newly recognized Famennian lungfishes from East Greenland reveal tooth plate diversity and blur the Devonian–Carboniferous boundary" 1363: 978: 484: 996:

within the attached epifaunal (seabed-living) filter-feeder megaguild. Other taxa impacted by the extinction rediversified or their

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Otoo, Benjamin K. A.; Clack, Jennifer A.; Smithson, Timothy R.; Bennett, Carys E.; Kearsey, Timothy I.; Coates, Michael I. (2019).

1076:, which were already fairly rare, experienced a large extinction and ecological turnover before rediversifying in the Tournaisian. 884: 946:

extinctions tracked via proportional genera extinctions by Bambach (2006). The Hangenberg Event is labelled at "8 Late Famennian".

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at the start of the VI zone, the last spore zone of the Devonian. This land plant extinction, which wiped out most or all of the

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zone, in the early part of the crisis. 75% of remaining families, 86% of genera, and 87% of species died out at this time. A few

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of Hangenberg event communities was restricted to only a few ecological groups, particularly slow-moving pelagic predators (i.e.

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extinction(s) also surpassed the Famennian extinction(s) in the “multiple-interval” and “well-preserved” full-stage categories.

2571:"Devonian–Carboniferous Hangenberg mass extinction event, widespread organic-rich mudrock and anoxia: causes and consequences" 4389: 4333:

Kravchinsky, V. A. (2012). "Paleozoic large igneous provinces of Northern Eurasia: Correlation with mass extinction events".

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Sallan, L.; Galimberti, A. K. (2015-11-13). "Body-size reduction in vertebrates following the end-Devonian mass extinction".

2431: 1367: 55:-Famennian boundary. The event is named after the Hangenberg Shale, which is part of a sequence that straddles the Devonian- 477: 4023:. Special Paper. Vol. 356. Geological Society of America. pp. 473–487. Archived from the original on 2010-03-26. 1513:

During the Famennian, the world was covered by a fairly homogenous and low-diversity land plant flora, dominated by giant

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deep-water environments. This is correlated with the beginning of the LN spore zone, indicated by the first occurrence of

2135:"Greenhouse to icehouse: a biostratigraphic review of latest Devonian–Mississippian glaciations and their global effects" 1426:, survived but failed to significantly increase their ecological disparity, eventually dying out later in the Paleozoic. 304:

I conodont zone and LN spore zone, and foraminifera are still absent. However, ammonoid fossils switch over to the lower

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Barbosa, Roberto Cesar de Mendonça; Nogueira, Afonso César Rodrigues; Domingos, Fábio Henrique Garcia (August 2015).

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Myrow, Paul M.; Ramezani, Jahandar; Hanson, Anne E.; Bowring, Samuel A.; Racki, Grzegorz; Rakociński, Michał (2014).

1694:, cannot be dated precisely enough to determine any causal relationship between the impact and the extinction event. 1044:

ecosystems disappeared from the fossil record during the Hangenberg Event, not returning until the late Tournaisian.

2190:"A high-precision U–Pb age constraint on the Rhynie Chert Konservat-Lagerstätte: time scale and other implications" 3754:"Late Frasnian–Famennian climates based on palynomorph analyses and the question of the Late Devonian glaciations" 1434:

from marine environments. Among the most major ecological changes associated with the extinction are the rise of

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thousand years, occupying the upper third of the ‘Strunian’ (latest Famennian), and a small portion of the early

3149: 2033:"High-precision U–Pb age and duration of the latest Devonian (Famennian) Hangenberg event, and its implications" 1430:(lungfish) persisted through the extinction more easily than other sarcopterygians, though they were apparently 4430: 3657:

Byrne, Hannah M.; Niedźwiedzki, Grzegorz; Blom, Henning; Kear, Benjamin P.; Ahlberg, Per E. (1 November 2022).

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as the kill mechanism, at least for terrestrial organisms. Intense warming may lead to increased convection of

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Isaacson, P. E.; Díaz-Martínez, E.; Grader, G. W.; Kalvoda, J.; Babek, O.; Devuyst, F. X. (24 October 2008).

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McGhee, George R.; Clapham, Matthew E.; Sheehan, Peter M.; Bottjer, David J.; Droser, Mary L. (2013-01-15).

4165:"Comment on "UV-B radiation was the Devonian–Carboniferous boundary terrestrial extinction kill mechanism"" 1779:"The global Hangenberg Crisis (Devonian–Carboniferous transition): review of a first-order mass extinction" 1068:

sponges, a major group of Devonian reef builders, completely died out in the Hangenberg Event. Conversely,

4367: 3967: 3753: 2570: 2300: 2087: 1927: 4335: 4049: 3335:

Clack, Jennifer Alice; Challands, Thomas James; Smithson, Timothy Richard; Smithson, Keturah Zoe (2019).

285: 2837:"The youngest African clymeniids (Ammonoidea, Late Devonian) – failed survivors of the Hangenberg Event" 2876: 2659: 324:

suggest that the sea level fell by more than 100 meters (328 feet) during the middle crisis interval.

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in the Carboniferous. However, no known Famennian "tetrapod" persisted into the Carboniferous, with "

3968:"Late Devonian–earliest Mississippian glaciation in Gondwanaland and its biogeographic consequences" 2836: 2835:

Korn, Dieter; Belka, Zdzislaw; Fröhlich, Sebastian; Rücklin, Martin; Wendt, Jobst (2 January 2007).

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Kaiser, Sandra Isabella; Becker, Ralf Thomas; Steuber, Thomas; Aboussalam, Sarah Zhor (2011-10-01).

4420: 1703: 831: 226:(sea level rise), as indicated by flooding reducing the input of terrestrial spores and increasing 3610:"A fish and tetrapod fauna from Romer's Gap preserved in Scottish Tournaisian floodplain deposits" 3541:

Anderson, Jason S.; Smithson, Tim; Mansky, Chris F.; Meyer, Taran; Clack, Jennifer (2015-04-27).

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diversity was somewhat impacted by the event, with survival largely based on ecology. Deep-water

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I) conodont zone. Foraminifera disappear from the fossil record during the black shale interval.

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Streel, Maurice; Caputo, Mário V.; Loboziak, Stanislas; Melo, José Henrique G. (November 2000).

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Smithson, Timothy R.; Wood, Stanley P.; Marshall, John E. A.; Clack, Jennifer A. (2012-03-20).

3128: 1909: 1595:. However, support for a rapid increase in plant cover at the end of the Famennian is lacking. 1529:

at this time. However, they started to decline near the end of the LN zone and the terrestrial

4095:"UV-B radiation was the Devonian-Carboniferous boundary terrestrial extinction kill mechanism" 2460: 1562:

Plants were significantly more affected by the Hangenberg event than by the Kellwasser event.

1457:" in the broad sense of the term) evidently survived, eventually leading to the earliest true 3758: 3055:"End-Devonian extinction and a bottleneck in the early evolution of modern jawed vertebrates" 1524: 3805:"Evidence for long-term climate change in Upper Devonian strata of the central Appalachians" 3336: 2281: 2189: 2032: 1497:”, has been linked to the Hangenberg Event. However, recent and continued discovery of many 4340: 4279: 4207: 4106: 3981: 3912:"Major North American Oil Source Yields Clues to One of Earth's Deadliest Mass Extinctions" 3869: 3818: 3767: 3713: 3672: 3621: 3554: 3487: 3428: 3348: 3299: 3244: 3137: 3068: 3002: 2935: 2888: 2793: 2726: 2671: 2636: 2624: 2582: 2530: 2475: 2359: 2312: 2228: 2146: 2099: 2044: 1986: 1939: 1860: 1790: 1650: 1180:

and died out shortly after it. Ammonoid extinction rates were highest near the base of the

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Stratigraphy and biostratigraphy of the Hangenberg Event in the classic Rhenish succession

8: 3476:"Earliest Carboniferous tetrapod and arthropod faunas from Scotland populate Romer's Gap" 4344: 4283: 4211: 4110: 3985: 3873: 3822: 3771: 3717: 3676: 3625: 3558: 3491: 3432: 3352: 3303: 3248: 3141: 3072: 3006: 2939: 2892: 2797: 2730: 2675: 2628: 2586: 2534: 2479: 2363: 2316: 2232: 2150: 2103: 2048: 1990: 1943: 1864: 1794: 197:

genozone (also known as the UD VI-D zone). A very short subzone (UD VI-D2) diagnosed by

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The main marine extinction pulse begins abruptly with the subsequent deposition of the

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Marshall, John E. A.; Lakin, Jon; Troth, Ian; Wallace-Johnson, Sarah M. (2020-05-01).

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diversity had already decreased in the Kellwasser Event, and all remaining subgroups (

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Yao, Le; Aretz, Markus; Chen, Jitao; Webb, Gregory E.; Wang, Xiangdong (2016-12-23).

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were barely affected, even in anoxic deep-water environments. Bivalves in the family

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In the middle crisis interval, the black shale grades into a thicker deposit of more

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Bault, Valentin; Balseiro, Diego; Monnet, Claude; Crônier, Catherine (2022-07-01).

2961: 2943: 2896: 2850: 2801: 2752: 2734: 2679: 2632: 2590: 2538: 2483: 2419: 2367: 2320: 2263: 2236: 2201: 2154: 2134: 2107: 2052: 1994: 1947: 1868: 1798: 1691: 1065: 997: 993: 924: 289: 3993: 3830: 3802: 3685: 3658: 3567: 2900: 2683: 2543: 2487: 2111: 1630: 1474: 1419: 1334: 1284: 1245: 1195: 938: 687: 444: 269: 32: 2515: 2423: 140: 4368:"Toward understanding Late Devonian global events: few answers, many questions" 4292: 4189:

Proceedings of the National Academy of Sciences of the United States of America

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Catastrophic Events and Mass Extinctions: Impacts and Beyond: Boulder, Colorado

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Proceedings of the National Academy of Sciences of the United States of America

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Proceedings of the National Academy of Sciences of the United States of America

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declined strongly in the late Famennian and were very rare in the Tournaisian.

1226: 1153: 1118: 913: 678: 407: 360: 305: 227: 121: 60: 2854: 2411: 2205: 1998: 1198:, survived the full extinction interval and went on to rediversify into later 4404: 4301: 4229: 4128: 3735: 3643: 3576: 3509: 3368: 3321: 3090: 3024: 2957: 2908: 2813: 2781: 2748: 2691: 2644: 2552: 2495: 2379: 2332: 2166: 2119: 2064: 2006: 1959: 1880: 1810: 1654: 1515: 1450: 1396: 1392: 1388: 1338: 1017: 989: 902: 898: 696: 669: 209: 157: 56: 4220: 3500: 3441: 3288:"Lungfish diversity in Romer's Gap: reaction to the end-Devonian extinction" 3256: 3081: 2922:

Brom, Krzysztof R.; Salamon, Mariusz A.; Gorzelak, Przemysław (2018-06-25).

2461:"A new ecological-severity ranking of major Phanerozoic biodiversity crises" 2371: 1127: 4373:

Understanding Late Devonian And Permian-Triassic Biotic and Climatic Events

4319: 4247: 4146: 4119: 3889: 3594: 3527: 3460: 3264: 3108: 2975: 2924:"Body-size increase in crinoids following the end-Devonian mass extinction" 2766: 2267: 1658: 1576: 1489: 1483: 1415: 1408: 1384: 1329: 1280: 1234: 1160: 1057: 986: 952: 788: 416: 356: 182: 177: 3862:"Basin-scale reconstruction of euxinia and Late Devonian mass extinctions" 2658:

Frey, Linda; Rücklin, Martin; Korn, Dieter; Klug, Christian (2018-05-01).

2412:"Patterns of Phanerozoic Extinction: a Perspective from Global Data Bases" 2387: 2133:

Lakin, J. A.; Marshall, J. E. A.; Troth, I.; Harding, I. C. (2016-01-01).

1637:

in the atmosphere, reacting to inorganic chlorine compounds and producing

264: 3702:"A terrestrial Devonian-Carboniferous boundary section in East Greenland" 1708: 1649:

would be capable of a similar degree of ozone depletion. The impact of a

1634: 1626: 1592: 1580: 1520: 1502: 1315: 1288: 1230: 982: 943: 928: 379:

which stretched across much of the Late Carboniferous and Early Permian.

368: 219: 117: 4263: 3414: 110:– marls and limestones (equivalent to Stockum Limestone and later units) 3634: 3609: 3312: 3287: 1678: 1642: 1431: 1423: 1376: 1372: 1359: 1222: 1210: 1173: 1169: 1149: 1010: 1006: 975: 704: 317: 190: 160:

input indicates that the Drewer Sandstone was deposited during a minor

129: 3360: 2739: 2256:

Philosophical Transactions of the Royal Society B: Biological Sciences

2056: 327:

This regression was caused by a cooling episode, and time-constrained

3286:

Smithson, Timothy R.; Richards, Kelly R.; Clack, Jennifer A. (2016).

2158: 1872: 1777:

Kaiser, Sandra Isabella; Aretz, Markus; Becker, Ralph Thomas (2016).

1662: 1646: 1466: 1458: 1380: 1324: 1276: 1253: 1249: 1206: 1199: 746: 639: 424: 293: 149: 135: 91:– cephalopod-rich nodular limestone (equivalent to Wocklum Limestone) 36: 4259: 4257: 3847:

Plants invade the land - evolutionary and environmental perspectives

1802: 1677:

and mercury spikes occurring in the Tien Shan Mountains of southern

1498: 1082: 411:

conodont zone and further ammonoid diversification within the upper

372: 4202: 2301:"Strength, timing, setting and cause of mid-Palaeozoic extinctions" 2240: 2188:

Parry, S. F.; Noble, S. R.; Crowley, Q. G.; Wellman, C. H. (2011).

1674: 1470: 1454: 1427: 1347: 1272: 1261: 1257: 1214: 1146: 1136: 1107: 1069: 1045: 1021: 1002: 932: 906: 739: 732: 725: 659: 649: 405:. The base of the Stockum Limestone also sees the beginning of the 340: 172: 52: 44: 3965: 4254: 4092: 2841: 1462: 1352: 1238: 1213:

is poorly studied, but appears to have been significant as well.

1156: 1098: 1077: 1025: 971: 711: 364: 332: 328: 321: 300:), and fossils are still rare. These layers are still within the 153: 125: 64: 3803:

Brezinski, D.K.; Cecil, C.B.; Skema, V.W.; Kertis, C.A. (2009).

3747: 3745: 1073: 1053: 718: 336: 253: 3334: 387:

The upper crisis interval begins with the return of prominent

3742: 2991:"Post-Ordovician trilobite diversity and evolutionary faunas" 2085: 1439: 1304: 1049: 920: 415:

genozone (LC I-A1). A major extinction among land plants and

352: 201:

occurs in the first few layers of the lower crisis interval.

165: 152:

deposit which initiates the lower crisis interval. Increased

1653:

can be supported or refuted by testing for trace amounts of

3540: 3392:"After mass extinctions, the meek (fish) inherit the earth" 3150:

10.1666/0094-8373(2005)031<0006:POAEIT>2.0.CO;2

2458: 1145:, top left) nearly died out in the Hangenberg Event, while 1041: 919:

Sepkoski (1996) plotted extinction rates for marine animal

392: 348: 273: 4370:. In Over, D. J.; Morrow, J. R.; Wignall, Paul B. (eds.). 2988: 2030: 382: 375:, acted as a prelude to the far larger and more prolonged 259: 3656: 3473: 2132: 1264:

experienced notable faunal turnover, with groups such as

3798: 3796: 3751: 3607: 2834: 2187: 288:

shallow-water sediment. It may be represented by shale (

132:, confirming the global nature of the Hangenberg Event. 3932: 3849:. New York: Columbia University Press. pp. 213–36. 3543:"A Diverse Tetrapod Fauna at the Base of 'Romer's Gap'" 3415:

Ward, P.; Labandeira, C.; Laurin, M; Berner, R (2006).

3124:"Pulsed origination and extinction in the marine realm" 2348:"Diversification and extinction in the history of life" 4181: 3053:

Sallan, Lauren Cole; Coates, Michael I. (2010-06-01).

1558:

which were probably from fast-growing fern-like plants

1287:’ often seen after mass extinctions. The flask-shaped 4013: 3843: 3793: 3285: 96:– marly shale, including the Hangenberg Black Shale ( 85:

The Hangenberg succession at Kowala Quarry in Poland:

2921: 2657: 2514:Bond, David P.G.; Grasby, Stephen E. (2017-07-15). 2410:Sepkoski, J. John (1996), Walliser, Otto H. (ed.), 2218: 460: 453:genozone, LC I-A2) and the MFZ1 foraminifera zone. 193:faunas are dominated by wocklumeriids, forming the 105:– marly limestone (equivalent to Stockum Limestone) 4056:(2nd ed.). Academic Press. pp. 534–545. 2779: 1973:Marynowski, Leszek; Filipiak, Paweł (2007-05-01). 1972: 136:Prelude and extinction – the lower crisis interval 4184:"Supernova triggers for end-Devonian extinctions" 3973:Palaeogeography, Palaeoclimatology, Palaeoecology 3810:Palaeogeography, Palaeoclimatology, Palaeoecology 3664:Palaeogeography, Palaeoclimatology, Palaeoecology 2881:Palaeogeography, Palaeoclimatology, Palaeoecology 2780:Tolokonnikova, Zoya; Ernst, Andrej (2021-12-01). 2664:Palaeogeography, Palaeoclimatology, Palaeoecology 2575:Palaeogeography, Palaeoclimatology, Palaeoecology 2523:Palaeogeography, Palaeoclimatology, Palaeoecology 2468:Palaeogeography, Palaeoclimatology, Palaeoecology 2305:Palaeogeography, Palaeoclimatology, Palaeoecology 2092:Palaeogeography, Palaeoclimatology, Palaeoecology 1850: 1776: 1333:, top) went extinct in the Hangenberg Event, and 1291:completely died out during the Hangenberg Event. 4402: 4033:: CS1 maint: bot: original URL status unknown ( 3234: 2712: 2418:, Berlin, Heidelberg: Springer, pp. 35–51, 2139:Geological Society, London, Special Publications 1853:Geological Society, London, Special Publications 1783:Geological Society, London, Special Publications 1395:(lobe-finned fish) were also strongly affected: 1391:) died out abruptly at the end of the Devonian. 230:. The Hangenberg Black Shale corresponds to the 3480:Proceedings of the National Academy of Sciences 2026: 2024: 367:, along with other short glacial phases in the 4052:. In Alderton, David; Elias, Scott A. (eds.). 3186: 2875:Scholze, Frank; Gess, Robert W. (2017-04-01). 4048:Rosa, Eduardo L. M.; Isbell, John L. (2021). 3230: 3228: 3226: 3224: 2617:Annual Review of Earth and Planetary Sciences 2247: 1925: 883:Subdivision of the Devonian according to the 485: 3868:. Nature volume 615 (2023) (7953): 640–645. 3187:Friedman, Matt; Sallan, Lauren Cole (2012). 3052: 2568: 2212: 2021: 4332: 3383: 2874: 2610:"Phanerozoic Biodiversity Mass Extinctions" 2569:Caplan, Mark L; Bustin, R.Mark (May 1999). 2288:. International Commission on Stratigraphy. 890:Vertical axis scale: millions of years ago. 208:, a layer of organic material deposited in 4047: 4041: 3408: 3221: 2513: 2200:(4). London: Geological Society: 863–872. 1461:, who gave rise to the fully terrestrial ( 1233:completely died out, but extinction among 492: 478: 316:, where rivers have cut into their former 276:, a valley fill deposit equivalent to the 16:Mass extinction at the end of the Devonian 4309: 4291: 4237: 4219: 4201: 4136: 4118: 3950: 3725: 3706:Palaeobiodiversity and Palaeoenvironments 3684: 3633: 3584: 3566: 3517: 3499: 3450: 3440: 3311: 3204: 3098: 3080: 3014: 2965: 2947: 2786:Palaeobiodiversity and Palaeoenvironments 2756: 2738: 2542: 4326: 4007: 3699: 2409: 1928:"Mass extinctions and sea-level changes" 1926:Hallam, A.; Wignall, P.B. (1999-12-01). 937: 263: 139: 3389: 2607: 1072:were apparently not strongly impacted. 383:Aftershocks – the upper crisis interval 260:Glaciation – the middle crisis interval 178:Siphonodella/Eosiphonodella praesulcata 4403: 2637:10.1146/annurev.earth.33.092203.122654 2345: 1088: 880: 456: 70: 4365: 4162: 4158: 4156: 3859: 3182: 3180: 3178: 3176: 3174: 3121: 3048: 3046: 3044: 3042: 2564: 2562: 2509: 2507: 2505: 2454: 2452: 2450: 2448: 2405: 2403: 2401: 2399: 2397: 2298: 2253: 1772: 1770: 1768: 1766: 1764: 1762: 1760: 1758: 1756: 1754: 1752: 1750: 1748: 1546:) as well as widespread tiny spores ( 1341:, bottom) also suffered heavy losses. 470: 2181: 1846: 1844: 1842: 1840: 1838: 1836: 1834: 1832: 1830: 1828: 1746: 1744: 1742: 1740: 1738: 1736: 1734: 1732: 1730: 1728: 858: 840: 817: 796: 778: 463: 3700:Marshall, John E. A. (2021-06-01). 2516:"On the causes of mass extinctions" 433:Siphonodella/Eosiphonodella sulcata 13: 4153: 4062:10.1016/B978-0-08-102908-4.00063-1 3171: 3039: 2559: 2502: 2445: 2394: 14: 4442: 2194:Journal of the Geological Society 1825: 1725: 1607: 1031: 974:severe extinction, tied with the 955:extinction in the Carboniferous. 3206:10.1111/j.1475-4983.2012.01165.x 2299:House, Michael R. (2002-06-20). 1314: 1303: 1163:(bottom right) all went extinct. 1126: 1117: 1106: 1097: 1036: 979:(end-Ordovician) mass extinction 77: 35:that occurred at the end of the 4359: 4353:10.1016/j.gloplacha.2012.01.007 4175: 4086: 3959: 3926: 3904: 3853: 3837: 3693: 3650: 3601: 3534: 3467: 3328: 3279: 3115: 3016:10.1016/j.earscirev.2022.104035 2982: 2915: 2868: 2828: 2773: 2706: 2651: 2601: 2339: 2292: 2274: 1685: 1591:seas and could have stimulated 3122:Foote, Michael (Winter 2005). 2126: 2079: 1966: 1919: 1895: 777: 168:zone. They also belong to the 1: 4382:10.1016/S0920-5446(05)80002-0 3780:10.1016/S0012-8252(00)00026-X 2595:10.1016/S0031-0182(98)00218-1 2325:10.1016/S0031-0182(01)00471-0 1952:10.1016/S0012-8252(99)00055-0 1719: 222:was deposited during a large 49:Late Devonian mass extinction 3994:10.1016/j.palaeo.2008.03.047 3939:Brazilian Journal of Geology 3831:10.1016/j.palaeo.2009.10.010 3686:10.1016/j.palaeo.2022.111215 3568:10.1371/journal.pone.0125446 2901:10.1016/j.palaeo.2017.01.018 2684:10.1016/j.palaeo.2017.12.028 2608:Bambach, Richard K. (2006). 2544:10.1016/j.palaeo.2016.11.005 2488:10.1016/j.palaeo.2012.12.019 2346:Benton, M. J. (1995-04-07). 2112:10.1016/j.palaeo.2011.07.026 1668: 1620: 1575:The Hangenberg event was an 1294: 413:Acutimitoceras (Stockumites) 339:(which would have been high- 331:deposits have been found in 314:incised valley fill deposits 7: 4376:. Elsevier. pp. 5–36. 4336:Global and Planetary Change 4050:"Late Paleozoic Glaciation" 2424:10.1007/978-3-642-79634-0_4 1697: 1205:Extinction in non-ammonoid 881: 634: 465:Devonian graphical timeline 307:Acutimitoceras (Stockumites 189:). The last pre-extinction 10: 4447: 4293:10.1038/s41598-021-85043-6 3952:10.1590/2317-4889201530147 3882:10.1038/s41586-023-05716-2 3727:10.1007/s12549-020-00448-x 3390:Feltman, R. (2015-11-13). 2949:10.1038/s41598-018-27986-x 2806:10.1007/s12549-020-00478-5 1598: 187:Quasiendothyra kobeitusana 51:(Kellwasser event) at the 4416:Late Devonian extinctions 2855:10.1080/00241160410002054 2206:10.1144/0016-76492010-043 1999:10.1017/S0016756807003317 1570: 1565: 1508: 1449:Four-limbed vertebrates ( 447:ammonoids (making up the 402:Tournayellina pseudobeata 215:Verrucosisporites nitidus 4366:Racki, Grzegorz (2005). 4163:Racki, Grzegorz (2021). 3914:. University of Maryland 1704:Late Devonian extinction 1481:stegocephalians such as 1414:Some large fish, namely 1275:suffered severe losses. 832:Late Devonian extinction 502: 4221:10.1073/pnas.2013774117 4054:Encyclopedia of Geology 3501:10.1073/pnas.1117332109 3442:10.1073/pnas.0607824103 3341:Papers in Palaeontology 3257:10.1126/science.aac7373 3082:10.1073/pnas.0914000107 2372:10.1126/science.7701342 1615:Late Palaeozoic Ice Age 1355:(jawless fish) groups. 625:−360 — 615:−365 — 605:−370 — 595:−375 — 585:−380 — 575:−385 — 565:−390 — 555:−395 — 545:−400 — 535:−405 — 525:−410 — 515:−415 — 505:−420 — 441:fossiliferous limestone 343:areas), as well as the 185:zone (characterized by 29:end-Devonian extinction 4120:10.1126/sciadv.aba0768 3860:Sahoo, Swapan (2023). 2268:10.1098/rstb.1998.0195 2221:The Journal of Geology 947: 408:Protognathodus kockeli 377:Late Paleozoic Ice Age 281: 206:Hangenberg Black Shale 145: 4431:Famennian extinctions 3759:Earth-Science Reviews 2995:Earth-Science Reviews 1932:Earth-Science Reviews 1579:marked by a layer of 1525:Retispora lepidophyta 941: 910:extinction trackers. 429:Protognathodus kuehni 363:. The Late Famennian 267: 143: 2286:www.stratigraphy.org 1183:Postclymenia evoluta 811:Famennian glaciation 437:Hangenberg Limestone 298:Hangenberg Sandstone 278:Hangenberg Sandstone 224:marine transgression 23:, also known as the 4345:2012GPC....86...31K 4284:2021NatSR..11.5708R 4212:2020PNAS..11721008F 4196:(35): 21008–21010. 4111:2020SciA....6..768M 3986:2008PPP...268..126I 3874:2023Natur.615..640S 3823:2009PPP...284..315B 3772:2000ESRv...52..121S 3718:2021PdPe..101..541M 3677:2022PPP...60511215B 3626:2019Palgy..62..225O 3559:2015PLoSO..1025446A 3492:2012PNAS..109.4532S 3433:2006PNAS..10316818W 3427:(45): 16818–16822. 3353:2019PPal....5..261C 3304:2016Palgy..59...29S 3249:2015Sci...350..812S 3142:2005Pbio...31....6F 3073:2010PNAS..10710131S 3067:(22): 10131–10135. 3007:2022ESRv..23004035B 2940:2018NatSR...8.9606B 2893:2017PPP...471...31S 2798:2021PdPe..101..885T 2731:2016NatSR...639694Y 2676:2018PPP...496....1F 2629:2006AREPS..34..127B 2587:1999PPP...148..187C 2535:2017PPP...478....3B 2480:2013PPP...370..260M 2364:1995Sci...268...52B 2317:2002PPP...181....5H 2233:2004JG....112..495K 2151:2016GSLSP.423..439L 2104:2011PPP...310..340K 2049:2014TeNov..26..222M 1991:2007GeoM..144..569M 1979:Geological Magazine 1944:1999ESRv...48..217H 1865:2016GSLSP.423..355B 1795:2016GSLSP.423..387K 1587:of semi-restricted 1531:ecosystem collapsed 1159:(bottom left), and 1089:Other invertebrates 964:Signor–Lipps effect 457:Extinction severity 250:Uranium-Lead dating 71:Geological evidence 4271:Scientific Reports 3635:10.1111/pala.12395 3313:10.1111/pala.12203 2928:Scientific Reports 2719:Scientific Reports 2282:"Chart/Time Scale" 1407:, and most other “ 1244:The two remaining 1178:dead clade walking 968:Pull of the Recent 948: 852:shrubs & trees 282: 175:zone (named after 146: 4391:978-0-444-52127-9 4071:978-0-08-102909-1 3486:(12): 4532–4537. 3361:10.1002/spp2.1242 3243:(6262): 812–815. 2740:10.1038/srep39694 2433:978-3-642-79634-0 2262:(1365): 113–130. 2057:10.1111/ter.12090 1915:on July 18, 2009. 895: 894: 876: 875: 857: 856: 839: 838: 816: 815: 795: 794: 397:Stockum Limestone 345:Appalachian Basin 162:marine regression 25:Hangenberg crisis 4438: 4426:Shale formations 4411:Devonian Germany 4396: 4395: 4363: 4357: 4356: 4339:. 86–87: 31–36. 4330: 4324: 4323: 4313: 4295: 4261: 4252: 4251: 4241: 4223: 4205: 4179: 4173: 4172: 4169:Science Advances 4160: 4151: 4150: 4140: 4122: 4105:(22): eaba0768. 4099:Science Advances 4090: 4084: 4083: 4045: 4039: 4038: 4032: 4024: 4022: 4011: 4005: 4004: 4002: 4000: 3980:(3–4): 126–142. 3963: 3957: 3956: 3954: 3930: 3924: 3923: 3921: 3919: 3908: 3902: 3901: 3857: 3851: 3850: 3841: 3835: 3834: 3817:(3–4): 315–325. 3800: 3791: 3790: 3788: 3786: 3766:(1–3): 121–173. 3749: 3740: 3739: 3729: 3697: 3691: 3690: 3688: 3654: 3648: 3647: 3637: 3605: 3599: 3598: 3588: 3570: 3538: 3532: 3531: 3521: 3503: 3471: 3465: 3464: 3454: 3444: 3412: 3406: 3405: 3403: 3402: 3387: 3381: 3380: 3332: 3326: 3325: 3315: 3283: 3277: 3276: 3232: 3219: 3218: 3208: 3184: 3169: 3168: 3166: 3164: 3119: 3113: 3112: 3102: 3084: 3050: 3037: 3036: 3018: 2986: 2980: 2979: 2969: 2951: 2919: 2913: 2912: 2872: 2866: 2865: 2863: 2861: 2832: 2826: 2825: 2777: 2771: 2770: 2760: 2742: 2710: 2704: 2703: 2655: 2649: 2648: 2614: 2605: 2599: 2598: 2566: 2557: 2556: 2546: 2520: 2511: 2500: 2499: 2465: 2456: 2443: 2442: 2441: 2440: 2407: 2392: 2391: 2343: 2337: 2336: 2296: 2290: 2289: 2278: 2272: 2271: 2251: 2245: 2244: 2216: 2210: 2209: 2185: 2179: 2178: 2159:10.1144/SP423.12 2130: 2124: 2123: 2098:(3–4): 340–364. 2083: 2077: 2076: 2028: 2019: 2018: 1970: 1964: 1963: 1923: 1917: 1916: 1914: 1908:. Archived from 1907: 1899: 1893: 1892: 1873:10.1144/SP423.10 1848: 1823: 1822: 1774: 1692:Woodleigh Crater 1651:nearby supernova 1475:ichthyostegalian 1444:actinopterygians 1411:” went extinct. 1405:tristichopterids 1318: 1307: 1130: 1121: 1110: 1101: 1013:) and epifaunal 864: 859: 846: 841: 828:Kellwasser event 823: 818: 807:Hangenberg event 802: 797: 784: 779: 699: 690: 681: 662: 642: 631: 626: 621: 616: 611: 606: 601: 596: 591: 586: 581: 576: 571: 566: 561: 556: 551: 546: 541: 536: 531: 526: 521: 516: 511: 506: 494: 487: 480: 474: 461: 290:Hangenberg Shale 81: 59:boundary in the 39:stage, the last 21:Hangenberg event 4446: 4445: 4441: 4440: 4439: 4437: 4436: 4435: 4421:Devonian events 4401: 4400: 4399: 4392: 4364: 4360: 4331: 4327: 4262: 4255: 4180: 4176: 4161: 4154: 4091: 4087: 4072: 4046: 4042: 4026: 4025: 4020: 4012: 4008: 3998: 3996: 3964: 3960: 3931: 3927: 3917: 3915: 3910: 3909: 3905: 3858: 3854: 3842: 3838: 3801: 3794: 3784: 3782: 3750: 3743: 3698: 3694: 3655: 3651: 3606: 3602: 3553:(4): e0125446. 3539: 3535: 3472: 3468: 3413: 3409: 3400: 3398: 3396:Washington Post 3388: 3384: 3333: 3329: 3284: 3280: 3233: 3222: 3185: 3172: 3162: 3160: 3120: 3116: 3051: 3040: 2987: 2983: 2920: 2916: 2873: 2869: 2859: 2857: 2833: 2829: 2778: 2774: 2711: 2707: 2656: 2652: 2612: 2606: 2602: 2567: 2560: 2518: 2512: 2503: 2463: 2457: 2446: 2438: 2436: 2434: 2408: 2395: 2358:(5207): 52–58. 2344: 2340: 2297: 2293: 2280: 2279: 2275: 2252: 2248: 2217: 2213: 2186: 2182: 2131: 2127: 2084: 2080: 2029: 2022: 1971: 1967: 1924: 1920: 1912: 1905: 1901: 1900: 1896: 1849: 1826: 1803:10.1144/SP423.9 1775: 1726: 1722: 1700: 1688: 1671: 1631:ozone depletion 1623: 1610: 1601: 1573: 1568: 1511: 1451:stegocephalians 1436:chondrichthyans 1397:onychodontidans 1393:Sarcopterygians 1345: 1344: 1343: 1342: 1335:sarcopterygians 1321: 1320: 1319: 1310: 1309: 1308: 1297: 1285:Lilliput effect 1266:leperditicopids 1196:Prionoceratidae 1167: 1166: 1165: 1164: 1133: 1132: 1131: 1123: 1122: 1113: 1112: 1111: 1103: 1102: 1091: 1070:tabulate corals 1039: 1034: 994:stromatoporoids 992:megaguild, and 927:throughout the 891: 889: 872: 871: 862: 853: 851: 844: 835: 821: 812: 810: 800: 791: 782: 775: 774: 770: 769: 765: 764: 760: 759: 755: 754: 750: 749: 743: 742: 736: 735: 729: 728: 722: 721: 715: 714: 708: 707: 701: 700: 695: 692: 691: 686: 683: 682: 677: 674: 673: 665: 664: 658: 654: 653: 645: 644: 638: 632: 629: 627: 624: 622: 619: 617: 614: 612: 609: 607: 604: 602: 599: 597: 594: 592: 589: 587: 584: 582: 579: 577: 574: 572: 569: 567: 564: 562: 559: 557: 554: 552: 549: 547: 544: 542: 539: 537: 534: 532: 529: 527: 524: 522: 519: 517: 514: 512: 509: 507: 504: 498: 472: 466: 459: 385: 270:Berea Sandstone 262: 252:of ash beds in 181:) and the DFZ7 138: 111: 106: 101: 92: 87: 82: 73: 33:mass extinction 17: 12: 11: 5: 4444: 4434: 4433: 4428: 4423: 4418: 4413: 4398: 4397: 4390: 4358: 4325: 4253: 4174: 4152: 4085: 4070: 4040: 4006: 3958: 3925: 3903: 3852: 3836: 3792: 3741: 3712:(2): 541–559. 3692: 3649: 3620:(2): 225–253. 3600: 3533: 3466: 3407: 3382: 3347:(2): 261–279. 3327: 3278: 3220: 3199:(4): 707–742. 3170: 3114: 3038: 2981: 2914: 2867: 2849:(3): 307–315. 2827: 2792:(4): 885–906. 2772: 2705: 2650: 2623:(1): 127–155. 2600: 2581:(4): 187–207. 2558: 2501: 2444: 2432: 2393: 2338: 2291: 2273: 2246: 2241:10.1086/421077 2227:(4): 495–501. 2211: 2180: 2145:(1): 439–464. 2125: 2078: 2043:(3): 222–229. 2020: 1985:(3): 569–595. 1965: 1938:(4): 217–250. 1918: 1894: 1859:(1): 355–386. 1824: 1789:(1): 387–437. 1723: 1721: 1718: 1717: 1716: 1714:Taghanic event 1711: 1706: 1699: 1696: 1687: 1684: 1670: 1667: 1645:from a nearby 1629:radiation and 1622: 1619: 1609: 1608:Global cooling 1606: 1600: 1597: 1589:epicontinental 1585:eutrophication 1572: 1569: 1567: 1564: 1539:R. lepidophyta 1510: 1507: 1420:megalichthyids 1364:end-Cretaceous 1323: 1322: 1313: 1312: 1311: 1302: 1301: 1300: 1299: 1298: 1296: 1293: 1227:rhynchonellids 1188:cymaclymeniids 1154:stromatoporoid 1135: 1134: 1125: 1124: 1116: 1115: 1114: 1105: 1104: 1096: 1095: 1094: 1093: 1092: 1090: 1087: 1066:stromatoporoid 1064:The last true 1058:calcimicrobial 1038: 1035: 1033: 1032:Impact on life 1030: 1018:filter feeders 893: 892: 878: 877: 874: 873: 869:Hunsrück fauna 867: 865: 855: 854: 849: 847: 837: 836: 826: 824: 814: 813: 805: 803: 793: 792: 787: 785: 776: 772: 771: 767: 766: 762: 761: 757: 756: 752: 751: 745: 744: 738: 737: 731: 730: 724: 723: 717: 716: 710: 709: 703: 702: 694: 693: 685: 684: 676: 675: 667: 666: 656: 655: 647: 646: 636: 635: 633: 628: 623: 618: 613: 608: 603: 598: 593: 588: 583: 578: 573: 568: 563: 558: 553: 548: 543: 538: 533: 528: 523: 518: 513: 508: 503: 500: 499: 497: 496: 489: 482: 471: 468: 467: 464: 458: 455: 445:gattendorfiine 423:ammonoids and 384: 381: 361:central Africa 261: 258: 228:eutrophication 199:Epiwocklumeria 137: 134: 122:Rhenish Massif 113: 112: 83: 76: 72: 69: 61:Rhenish Massif 15: 9: 6: 4: 3: 2: 4443: 4432: 4429: 4427: 4424: 4422: 4419: 4417: 4414: 4412: 4409: 4408: 4406: 4393: 4387: 4383: 4379: 4375: 4374: 4369: 4362: 4354: 4350: 4346: 4342: 4338: 4337: 4329: 4321: 4317: 4312: 4307: 4303: 4299: 4294: 4289: 4285: 4281: 4277: 4273: 4272: 4267: 4260: 4258: 4249: 4245: 4240: 4235: 4231: 4227: 4222: 4217: 4213: 4209: 4204: 4199: 4195: 4191: 4190: 4185: 4178: 4170: 4166: 4159: 4157: 4148: 4144: 4139: 4134: 4130: 4126: 4121: 4116: 4112: 4108: 4104: 4100: 4096: 4089: 4081: 4077: 4073: 4067: 4063: 4059: 4055: 4051: 4044: 4036: 4030: 4019: 4018: 4010: 3995: 3991: 3987: 3983: 3979: 3975: 3974: 3969: 3962: 3953: 3948: 3944: 3940: 3936: 3929: 3913: 3907: 3899: 3895: 3891: 3887: 3883: 3879: 3875: 3871: 3867: 3863: 3856: 3848: 3840: 3832: 3828: 3824: 3820: 3816: 3812: 3811: 3806: 3799: 3797: 3781: 3777: 3773: 3769: 3765: 3761: 3760: 3755: 3748: 3746: 3737: 3733: 3728: 3723: 3719: 3715: 3711: 3707: 3703: 3696: 3687: 3682: 3678: 3674: 3670: 3666: 3665: 3660: 3653: 3645: 3641: 3636: 3631: 3627: 3623: 3619: 3615: 3614:Palaeontology 3611: 3604: 3596: 3592: 3587: 3582: 3578: 3574: 3569: 3564: 3560: 3556: 3552: 3548: 3544: 3537: 3529: 3525: 3520: 3515: 3511: 3507: 3502: 3497: 3493: 3489: 3485: 3481: 3477: 3470: 3462: 3458: 3453: 3448: 3443: 3438: 3434: 3430: 3426: 3422: 3418: 3411: 3397: 3393: 3386: 3378: 3374: 3370: 3366: 3362: 3358: 3354: 3350: 3346: 3342: 3338: 3331: 3323: 3319: 3314: 3309: 3305: 3301: 3297: 3293: 3292:Palaeontology 3289: 3282: 3274: 3270: 3266: 3262: 3258: 3254: 3250: 3246: 3242: 3238: 3231: 3229: 3227: 3225: 3216: 3212: 3207: 3202: 3198: 3194: 3193:Palaeontology 3190: 3183: 3181: 3179: 3177: 3175: 3159: 3155: 3151: 3147: 3143: 3139: 3135: 3131: 3130: 3125: 3118: 3110: 3106: 3101: 3096: 3092: 3088: 3083: 3078: 3074: 3070: 3066: 3062: 3061: 3056: 3049: 3047: 3045: 3043: 3034: 3030: 3026: 3022: 3017: 3012: 3008: 3004: 3000: 2996: 2992: 2985: 2977: 2973: 2968: 2963: 2959: 2955: 2950: 2945: 2941: 2937: 2933: 2929: 2925: 2918: 2910: 2906: 2902: 2898: 2894: 2890: 2886: 2882: 2878: 2871: 2856: 2852: 2848: 2844: 2843: 2838: 2831: 2823: 2819: 2815: 2811: 2807: 2803: 2799: 2795: 2791: 2787: 2783: 2776: 2768: 2764: 2759: 2754: 2750: 2746: 2741: 2736: 2732: 2728: 2724: 2720: 2716: 2709: 2701: 2697: 2693: 2689: 2685: 2681: 2677: 2673: 2669: 2665: 2661: 2654: 2646: 2642: 2638: 2634: 2630: 2626: 2622: 2618: 2611: 2604: 2596: 2592: 2588: 2584: 2580: 2576: 2572: 2565: 2563: 2554: 2550: 2545: 2540: 2536: 2532: 2528: 2524: 2517: 2510: 2508: 2506: 2497: 2493: 2489: 2485: 2481: 2477: 2473: 2469: 2462: 2455: 2453: 2451: 2449: 2435: 2429: 2425: 2421: 2417: 2413: 2406: 2404: 2402: 2400: 2398: 2389: 2385: 2381: 2377: 2373: 2369: 2365: 2361: 2357: 2353: 2349: 2342: 2334: 2330: 2326: 2322: 2318: 2314: 2311:(1–3): 5–25. 2310: 2306: 2302: 2295: 2287: 2283: 2277: 2269: 2265: 2261: 2257: 2250: 2242: 2238: 2234: 2230: 2226: 2222: 2215: 2207: 2203: 2199: 2195: 2191: 2184: 2176: 2172: 2168: 2164: 2160: 2156: 2152: 2148: 2144: 2140: 2136: 2129: 2121: 2117: 2113: 2109: 2105: 2101: 2097: 2093: 2089: 2082: 2074: 2070: 2066: 2062: 2058: 2054: 2050: 2046: 2042: 2038: 2034: 2027: 2025: 2016: 2012: 2008: 2004: 2000: 1996: 1992: 1988: 1984: 1980: 1976: 1969: 1961: 1957: 1953: 1949: 1945: 1941: 1937: 1933: 1929: 1922: 1911: 1904: 1898: 1890: 1886: 1882: 1878: 1874: 1870: 1866: 1862: 1858: 1854: 1847: 1845: 1843: 1841: 1839: 1837: 1835: 1833: 1831: 1829: 1820: 1816: 1812: 1808: 1804: 1800: 1796: 1792: 1788: 1784: 1780: 1773: 1771: 1769: 1767: 1765: 1763: 1761: 1759: 1757: 1755: 1753: 1751: 1749: 1747: 1745: 1743: 1741: 1739: 1737: 1735: 1733: 1731: 1729: 1724: 1715: 1712: 1710: 1707: 1705: 1702: 1701: 1695: 1693: 1683: 1680: 1676: 1666: 1664: 1660: 1659:arc volcanism 1656: 1655:Plutonium-244 1652: 1648: 1644: 1640: 1636: 1632: 1628: 1618: 1616: 1605: 1596: 1594: 1590: 1586: 1582: 1578: 1563: 1561: 1557: 1553: 1549: 1545: 1540: 1536: 1535:Archaeopteris 1532: 1527: 1526: 1522: 1518: 1517: 1516:Archaeopteris 1506: 1504: 1500: 1496: 1492: 1491: 1486: 1485: 1480: 1476: 1472: 1468: 1464: 1460: 1456: 1452: 1447: 1445: 1442:and kin) and 1441: 1437: 1433: 1429: 1425: 1421: 1417: 1412: 1410: 1409:osteolepidids 1406: 1402: 1401:porolepiforms 1398: 1394: 1390: 1389:ptychodontids 1386: 1382: 1378: 1374: 1369: 1365: 1361: 1356: 1354: 1349: 1340: 1339:porolepiforms 1336: 1332: 1331: 1326: 1317: 1306: 1292: 1290: 1286: 1282: 1278: 1274: 1270: 1267: 1263: 1259: 1255: 1251: 1247: 1242: 1240: 1236: 1232: 1228: 1224: 1220: 1216: 1212: 1208: 1203: 1201: 1197: 1193: 1189: 1185: 1184: 1179: 1175: 1171: 1162: 1158: 1155: 1152:(top right), 1151: 1148: 1144: 1143: 1138: 1129: 1120: 1109: 1100: 1086: 1084: 1079: 1075: 1074:Rugose corals 1071: 1067: 1062: 1059: 1055: 1051: 1047: 1043: 1037:Reef builders 1029: 1027: 1023: 1019: 1016: 1012: 1008: 1004: 999: 995: 991: 990:filter-feeder 988: 984: 980: 977: 973: 969: 965: 961: 956: 954: 945: 940: 936: 934: 930: 926: 922: 917: 915: 911: 908: 904: 900: 888: 887:, as of 2021. 886: 879: 870: 866: 861: 860: 848: 843: 842: 833: 829: 825: 820: 819: 808: 804: 799: 798: 790: 786: 781: 780: 748: 741: 734: 727: 720: 713: 706: 698: 689: 680: 672: 671: 670:Carboniferous 663: 661: 652: 651: 643: 641: 501: 495: 490: 488: 483: 481: 476: 475: 469: 462: 454: 452: 451: 446: 442: 438: 434: 430: 426: 422: 421:cymaclymeniid 418: 414: 410: 409: 404: 403: 398: 394: 390: 380: 378: 374: 370: 366: 365:glacial phase 362: 358: 354: 350: 346: 342: 338: 334: 330: 325: 323: 319: 315: 310: 308: 303: 299: 295: 291: 287: 279: 275: 271: 266: 257: 255: 251: 247: 244:Interregnum ( 243: 239: 235: 234: 229: 225: 221: 217: 216: 211: 207: 202: 200: 196: 192: 188: 184: 180: 179: 174: 171: 167: 163: 159: 158:siliciclastic 155: 151: 142: 133: 131: 127: 123: 119: 109: 104: 100:) at its base 99: 95: 90: 86: 80: 75: 74: 68: 66: 62: 58: 57:Carboniferous 54: 50: 46: 42: 38: 34: 30: 26: 22: 4372: 4361: 4334: 4328: 4275: 4269: 4193: 4187: 4177: 4168: 4102: 4098: 4088: 4053: 4043: 4016: 4009: 3997:. Retrieved 3977: 3971: 3961: 3945:(1): 13–27. 3942: 3938: 3928: 3916:. Retrieved 3906: 3865: 3855: 3846: 3839: 3814: 3808: 3783:. Retrieved 3763: 3757: 3709: 3705: 3695: 3668: 3662: 3652: 3617: 3613: 3603: 3550: 3546: 3536: 3483: 3479: 3469: 3424: 3420: 3410: 3399:. Retrieved 3395: 3385: 3344: 3340: 3330: 3298:(1): 29–44. 3295: 3291: 3281: 3240: 3236: 3196: 3192: 3161:. Retrieved 3133: 3129:Paleobiology 3127: 3117: 3064: 3058: 2998: 2994: 2984: 2931: 2927: 2917: 2884: 2880: 2870: 2858:. Retrieved 2846: 2840: 2830: 2789: 2785: 2775: 2725:(1): 39694. 2722: 2718: 2708: 2667: 2663: 2653: 2620: 2616: 2603: 2578: 2574: 2526: 2522: 2471: 2467: 2437:, retrieved 2415: 2355: 2351: 2341: 2308: 2304: 2294: 2285: 2276: 2259: 2255: 2249: 2224: 2220: 2214: 2197: 2193: 2183: 2142: 2138: 2128: 2095: 2091: 2081: 2040: 2036: 1982: 1978: 1968: 1935: 1931: 1921: 1910:the original 1897: 1856: 1852: 1786: 1782: 1689: 1686:Impact event 1673:Evidence of 1672: 1624: 1611: 1602: 1593:algal blooms 1577:anoxic event 1574: 1559: 1555: 1551: 1547: 1538: 1534: 1523: 1514: 1512: 1490:Acanthostega 1488: 1484:Ichthyostega 1482: 1448: 1424:acanthodians 1422:, and a few 1413: 1385:phyllolepids 1357: 1346: 1330:Dunkleosteus 1328: 1289:chitinozoans 1281:Foraminifera 1271: 1243: 1204: 1192:Postclymenia 1191: 1181: 1168: 1161:chitinozoans 1142:Cymaclymenia 1140: 1063: 1040: 1011:acanthodians 983:chitinozoans 972:ecologically 959: 957: 953:Serpukhovian 949: 918: 912: 896: 882: 806: 789:Rhynie chert 668: 657: 648: 637: 450:Gattendorfia 448: 432: 428: 417:palynomorphs 412: 406: 400: 386: 357:South Africa 326: 320:. Strata in 306: 301: 283: 245: 241: 237: 233:Postclymenia 231: 213: 203: 198: 194: 186: 183:foraminifera 176: 169: 147: 114: 107: 102: 97: 93: 88: 84: 28: 24: 20: 18: 4278:(1): 5708. 3136:(1): 6–20. 2934:(1): 9606. 2474:: 260–270. 1709:Kacak Event 1643:cosmic rays 1635:water vapor 1581:black shale 1521:palynomorph 1519:trees. The 1503:Tournaisian 1495:Romer’s Gap 1377:arthrodires 1368:end-Permian 1360:vertebrates 1219:Naiaditidae 1190:(including 985:within the 944:Phanerozoic 929:Phanerozoic 473:This box: 369:Tournaisian 318:floodplains 220:black shale 195:Wocklumeria 170:praesulcata 118:Tournaisian 4405:Categories 4203:2007.01887 3999:28 January 3785:28 January 3671:: 111215. 3401:2015-11-16 3163:28 January 3001:: 104035. 2860:28 January 2439:2021-05-09 2037:Terra Nova 1720:References 1679:Uzbekistan 1556:Rugospora) 1467:sauropsids 1459:amphibians 1453:, a.k.a. " 1432:extirpated 1416:rhizodonts 1325:Placoderms 1277:Acritarchs 1250:trilobites 1223:Brachiopod 1211:gastropods 1207:nautiloids 1174:clymeniids 1150:trilobites 976:Hirnantian 850:Widespread 705:Lochkovian 395:unit, the 312:show deep 286:oxygenated 130:Antarctica 4302:2045-2322 4230:0027-8424 4129:2375-2548 4080:226643402 4029:cite book 3918:March 28, 3898:257426134 3736:1867-1608 3644:1475-4983 3577:1932-6203 3510:0027-8424 3377:134074159 3369:2056-2802 3322:1475-4983 3273:206640186 3091:0027-8424 3033:248439050 3025:0012-8252 2958:2045-2322 2909:0031-0182 2887:: 31–39. 2822:232273339 2814:1867-1608 2749:2045-2322 2700:133740756 2692:0031-0182 2645:0084-6597 2553:0031-0182 2496:0031-0182 2380:0036-8075 2333:0031-0182 2175:130689152 2167:0305-8719 2120:0031-0182 2073:131251110 2065:1365-3121 2015:129306243 2007:0016-7568 1960:0012-8252 1889:131491081 1881:0305-8719 1819:131270834 1811:0305-8719 1669:Volcanism 1663:acid rain 1647:supernova 1621:Supernova 1552:Diducites 1548:Retispora 1471:synapsids 1455:tetrapods 1381:antiarchs 1373:Placoderm 1348:Conodonts 1337:(such as 1327:(such as 1295:Chordates 1262:Ostracods 1254:Phacopida 1231:chonetids 1200:goniatite 1170:Ammonoids 1139:(such as 1137:Ammonoids 1078:Bryozoans 1026:bryozoans 1007:ammonoids 747:Famennian 640:Paleozoic 391:rocks: a 389:carbonate 294:sandstone 150:turbidite 37:Famennian 4320:33707566 4248:32817482 4147:32937380 3890:36890233 3595:25915639 3547:PLOS ONE 3528:22393016 3461:17065318 3265:26564854 3215:59423401 3158:53469954 3109:20479258 2976:29942036 2767:28009013 2670:: 1–17. 2529:: 3–29. 1698:See also 1675:coronene 1428:Dipnoans 1353:agnathan 1273:Plankton 1258:Proetida 1239:Crinoids 1215:Bivalves 1202:groups. 1147:phacopid 1046:Metazoan 1022:bivalves 1003:ecospace 933:Givetian 925:families 907:Cambrian 903:Sepkoski 740:Frasnian 733:Givetian 726:Eifelian 660:Devonian 650:Silurian 425:phacopid 341:latitude 238:costatus 191:ammonoid 173:conodont 53:Frasnian 45:Devonian 4341:Bibcode 4311:7970954 4280:Bibcode 4239:7474607 4208:Bibcode 4138:7253167 4107:Bibcode 3982:Bibcode 3870:Bibcode 3819:Bibcode 3768:Bibcode 3714:Bibcode 3673:Bibcode 3622:Bibcode 3586:4411152 3555:Bibcode 3519:3311392 3488:Bibcode 3452:1636538 3429:Bibcode 3349:Bibcode 3300:Bibcode 3245:Bibcode 3237:Science 3138:Bibcode 3100:2890420 3069:Bibcode 3003:Bibcode 2967:6018515 2936:Bibcode 2889:Bibcode 2842:Lethaia 2794:Bibcode 2758:5180103 2727:Bibcode 2672:Bibcode 2625:Bibcode 2583:Bibcode 2531:Bibcode 2476:Bibcode 2388:7701342 2360:Bibcode 2352:Science 2313:Bibcode 2229:Bibcode 2147:Bibcode 2100:Bibcode 2045:Bibcode 1987:Bibcode 1940:Bibcode 1861:Bibcode 1791:Bibcode 1599:Euxinia 1544:lycopod 1463:amniote 1235:neritic 1157:sponges 1085:stage. 1015:sessile 987:pelagic 958:McGhee 863:← 845:← 822:← 801:← 783:← 712:Pragian 630:– 620:– 610:– 600:– 590:– 580:– 570:– 560:– 550:– 540:– 530:– 520:– 510:– 333:Bolivia 329:glacial 322:Morocco 242:kockeli 154:erosion 126:Germany 65:Germany 43:in the 31:, is a 4388: 4318: 4308: 4300: 4246: 4236: 4228: 4145: 4135: 4127: 4078: 4068: 3896: 3888: 3866:Nature 3734: 3642: 3593: 3583: 3575: 3526: 3516: 3508: 3459: 3449: 3375: 3367: 3320: 3271: 3263: 3213: 3156: 3107: 3097: 3089: 3031: 3023: 2974: 2964: 2956: 2907: 2820: 2812: 2765: 2755: 2747: 2698: 2690: 2643: 2551: 2494: 2430: 2386: 2378: 2331: 2173: 2165: 2118: 2071: 2063: 2013: 2005: 1958: 1887: 1879: 1817: 1809: 1571:Anoxia 1566:Causes 1509:Plants 1499:Visean 1440:sharks 1387:, and 1358:Other 1246:orders 1083:Visean 1054:sponge 1020:(i.e. 998:niches 942:Major 921:genera 914:Benton 773: 768: 763: 758: 753: 719:Emsian 373:Visean 359:, and 337:Brazil 254:Poland 210:anoxic 4198:arXiv 4076:S2CID 4021:(PDF) 3894:S2CID 3373:S2CID 3269:S2CID 3211:S2CID 3154:S2CID 3029:S2CID 2818:S2CID 2696:S2CID 2613:(PDF) 2519:(PDF) 2464:(PDF) 2171:S2CID 2069:S2CID 2011:S2CID 1913:(PDF) 1906:(PDF) 1885:S2CID 1815:S2CID 1479:grade 1050:coral 960:et al 679:Early 393:marly 353:Libya 292:) or 166:spore 41:stage 4386:ISBN 4316:PMID 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