Carboniferous rainforest collapse

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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. Alethopteris serli and Neuropteris sp., Carboniferous (Pennsylvanian), Llewellyn Formation, St. Clair, Schuykill County, Pennsylvania, USA - Houston Museum of Natural Science - DSC01757.JPG
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] The event occurred at the end of the Moscovian and continued into the early Kasimovian stages of the Pennsylvanian (Upper Carboniferous).

Contents

It altered the vast coal forests that covered the equatorial region of Euramerica (Europe and North 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.

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 griffinflies, millipedes, blattopterans, 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 2011 study showing that the presence of meandering and anabranching streams, 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. Little mixing of different plant assemblages occurred throughout this transition; floral assemblages were highly discrete and conservative and gave way to new ones without any transitional floras intermediate in composition with regards to the preceding one and succeeding one. [5]

Invertebrates

The fossil record of insects can be difficult to study, due 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–Triassic extinction event. 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 and the transition of lycopod to tree fern-dominated land floras. [6]

Vertebrates

Before the collapse, vertebrate animal species distribution was very cosmopolitan, with the same species existing across tropical Pangaea. 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 CRC, 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. [7]

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. [8] By the Asselian, many families of seed ferns that characterized the Moscovian tropical wetlands had disappeared including Flemingitaceae, Diaphorodendraceae, Tedeleaceae, Urnatopteridaceae, Cyclopteridaceae, and Neurodontopteridaceae. [8]

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. [9] 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. [10] [11] [12] This means that rainforest collapse and 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. Archaeothyris BW.jpg
Terrestrially adapted synapsids, the predecessors of the mammal lineage, like Archaeothyris were among the groups who quickly recovered after the collapse.

The 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. [13] 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. [14] [15] [16] After the late Bashkirian glacial maximum of the Late Paleozoic Ice Age  I, around 318  Ma, frequent shifts in seasonality from humid to arid times began. [17]

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, coinciding with abrupt faunal changes in marine and terrestrial species. [18] 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. [18] [19] [20]

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. [21] 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. [17] [21] As the climate aridified through the Late Paleozoic, rainforests were eventually replaced by seasonally dry biomes. [22]

Volcanism

After restoring the middle of the Skagerrak-Centered Large Igneous Province using a new reference frame, it has been shown that the Skagerrak plume rose from the core–mantle boundary to its ~300 Ma position. [23] 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. [24]

Geography

While the CRC affected 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 persisted until the end of the Permian, around 252 million years ago.

Fossil sites

Fossil lycopsid, probably Sigillaria, from Joggins, with attached stigmarian roots Lycopsid joggins mcr1.JPG
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 . [25]

Related Research Articles

<span class="mw-page-title-main">Carboniferous</span> Fifth period of the Paleozoic Era, 359–299 million years ago

The Carboniferous is a geologic period and system of the Paleozoic that spans 60 million years from the end of the Devonian Period 358.9 Ma to the beginning of the Permian Period, 298.9 Ma. In North America, the Carboniferous is often treated as two separate geological periods, the earlier Mississippian and the later Pennsylvanian.

The Pennsylvanian is, on the ICS geologic timescale, the younger of two subperiods of the Carboniferous Period. It lasted from roughly 323.2 million years ago to 298.9 million years ago. As with most other geochronologic units, the rock beds that define the Pennsylvanian are well identified, but the exact date of the start and end are uncertain by a few hundred thousand years. The Pennsylvanian is named after the U.S. state of Pennsylvania, where the coal-producing beds of this age are widespread.

<span class="mw-page-title-main">Permian</span> Sixth and last period of the Paleozoic Era 299–252 million years ago

The Permian is a geologic period and stratigraphic system which spans 47 million years from the end of the Carboniferous Period 298.9 million years ago (Mya), to the beginning of the Triassic Period 251.902 Mya. It is the last period of the Paleozoic Era; the following Triassic Period belongs to the Mesozoic Era. The concept of the Permian was introduced in 1841 by geologist Sir Roderick Murchison, who named it after the region of Perm in Russia.

The PaleozoicEra is the first of three geological eras of the Phanerozoic Eon. Beginning 538.8 million years ago (Ma), it succeeds the Neoproterozoic and ends 251.9 Ma at the start of the Mesozoic Era. The Paleozoic is subdivided into six geologic periods :

The Phanerozoic is the current and the latest of the four geologic eons in the Earth's geologic time scale, covering the time period from 538.8 million years ago to the present. It is the eon during which abundant animal and plant life has proliferated, diversified and colonized various niches on the Earth's surface, beginning with the Cambrian period when animals first developed hard shells that can be clearly preserved in the fossil record. The time before the Phanerozoic, collectively called the Precambrian, is now divided into the Hadean, Archaean and Proterozoic eons.

<span class="mw-page-title-main">Labyrinthodontia</span> Paraphyletic group of tetrapodomorphs

"Labyrinthodontia" is an informal grouping of extinct predatory amphibians which were major components of ecosystems in the late Paleozoic and early Mesozoic eras. Traditionally considered a subclass of the class Amphibia, modern classification systems recognize that labyrinthodonts are not a formal natural group (clade) exclusive of other tetrapods. Instead, they consistute an evolutionary grade, ancestral to living tetrapods such as lissamphibians and amniotes. "Labyrinthodont"-grade vertebrates evolved from lobe-finned fishes in the Devonian, though a formal boundary between fish and amphibian is difficult to define at this point in time.

<span class="mw-page-title-main">Diadectomorpha</span> Extinct clade of tetrapods

Diadectomorpha is a clade of large tetrapods that lived in Euramerica during the Carboniferous and Early Permian periods and in Asia during Late Permian (Wuchiapingian), They have typically been classified as advanced reptiliomorphs positioned close to, but outside of the clade Amniota, though some recent research has recovered them as the sister group to the traditional Synapsida within Amniota, based on inner ear anatomy and cladistic analyses. They include both large carnivorous and even larger herbivorous forms, some semi-aquatic and others fully terrestrial. The diadectomorphs seem to have originated during late Mississippian times, although they only became common after the Carboniferous rainforest collapse and flourished during the Late Pennsylvanian and Early Permian periods.

<span class="mw-page-title-main">Joggins</span> Community in Nova Scotia, Canada

Joggins is a rural community located in western Cumberland County, Nova Scotia, Canada. On July 7, 2008 a 15-km length of the coast constituting the Joggins Fossil Cliffs was officially inscribed on the World Heritage List.

<i>Arthropleura</i> Extinct genus of many-legged arthropods

Arthropleura is an extinct genus of massive millipedes that lived in what is now North America and Europe around 345 to 290 million years ago, from the Viséan stage of the lower Carboniferous Period to the Sakmarian stage of the lower Permian Period. The species of the genus are the largest known land invertebrates of all time, and would have had few, if any, predators.

<span class="mw-page-title-main">Late Paleozoic icehouse</span> Ice age

The late Paleozoic icehouse, also known as the Late Paleozoic Ice Age (LPIA) and formerly known as the Karoo ice age, was an ice age that began in the Late Devonian and ended in the Late Permian, occurring from 360 to 255 million years ago (Mya), and large land-based ice-sheets were then present on Earth's surface. It was the second major icehouse period of the Phanerozoic.

The Kasimovian is a geochronologic age or chronostratigraphic stage in the ICS geologic timescale. It is the third stage in the Pennsylvanian, lasting from 307 to 303.7 Ma. The Kasimovian Stage follows the Moscovian and is followed by the Gzhelian. The Kasimovian saw an extinction event which occurred around 305 mya, referred to as the Carboniferous Rainforest Collapse. It roughly corresponds to the Missourian in North American geochronology and the Stephanian in western European geochronology.

<i>Limnoscelis</i> Genus of diadectomorphs

Limnoscelis was a genus of large diadectomorph tetrapods from the Late Carboniferous to early Permian of western North America. It includes two species: the type species Limnoscelis paludis from New Mexico, and Limnoscelis dynatis from Colorado, both of which are thought to have lived concurrently. No specimens of Limnoscelis are known from outside of North America. Limnoscelis was carnivorous, and likely semiaquatic, though it may have spent a significant portion of its life on land. Limnoscelis had a combination of derived amphibian and primitive reptilian features, and its placement relative to Amniota has significant implications regarding the origins of the first amniotes.

<span class="mw-page-title-main">Allegheny Group</span> Pennsylvanian-age geological unit

The Allegheny Group, often termed the Allegheny Formation, is a Pennsylvanian-age geological unit in the Appalachian Plateau. It is a major coal-bearing unit in the eastern United States, extending through western and central Pennsylvania, western Maryland and West Virginia, and southeastern Ohio. Fossils of fishes such as Bandringa are known from the Kittaning Formation, which is part of the Allegheny Group.

Macromerion is an extinct genus of non-mammalian synapsids, specifically Pelycosaurs, in the family Sphenacodontidae from Late Carboniferous deposits in the Czech Republic. It was named as a species of Labyrinthodon in 1875 and as its own genus in 1879.

<span class="mw-page-title-main">Medullosales</span> Extinct order of Late Carboniferous seed ferns

The Medullosales is an extinct order of pteridospermous seed plants characterised by large ovules with circular cross-section and a vascularised nucellus, complex pollen-organs, stems and rachides with a dissected stele, and frond-like leaves. Their nearest still-living relatives are the cycads.

The Westphalian is a regional stage or age in the regional stratigraphy of northwest Europe, with an age between roughly 315 and 307 Ma. It is a subdivision of the Carboniferous System or Period and the regional Silesian Series. The Westphalian is named for the region of Westphalia in western Germany where strata of this age occur. The Coal Measures of England and Wales are also largely of Westphalian age, though they also extend into the succeeding Stephanian.

<span class="mw-page-title-main">Callistophytaceae</span> Extinct family of seed ferns

The Callistophytaceae was a family of seed ferns (pteridosperms) from the Carboniferous and Permian periods. They first appeared in late Middle Pennsylvanian (Moscovian) times, 306.5–311.7 million years ago (Ma) in the tropical coal forests of Euramerica, and became an important component of Late Pennsylvanian vegetation of clastic soils and some peat soils. The best known callistophyte was documented from Late Pennsylvanian coal ball petrifactions in North America.

Olson's Extinction was a mass extinction that occurred 273 million years ago in the late Cisuralian or early Guadalupian epoch of the Permian period, predating the much larger Permian–Triassic extinction event. The event is named after American paleontologist Everett C. Olson, who first identified the gap in fossil record indicating a sudden change between the early Permian and middle/late Permian faunas. Some authors also place a hiatus in the continental fossil record around that time, but others disagree. This event has been argued by some authors to have affected many taxa, including embryophytes, marine metazoans, and tetrapods.

<i>Macroneuropteris</i> Extinct genus of plants

Macroneuropteris is a genus of Carboniferous seed plants in the order Medullosales. The genus is best known for the species Macroneuropteris scheuchzeri, a medium-size tree that was common throughout the late Carboniferous Euramerica. Three similar species, M. macrophylla, M. britannica and M. subauriculata are also included in the genus.

This timeline of Permian research is a chronological listing of events in the history of geology and paleontology focused on the study of earth during the span of time lasting from 298.9–252.17 million years ago and the legacies of this period in the rock and fossil records.

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Further reading