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The history of life on
Earth Earth is the third planet from the Sun and the only astronomical object known to harbor life. While large volumes of water can be found throughout the Solar System, only Earth sustains liquid surface water. About 71% of Earth's sur ...
traces the processes by which living and
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
organism In biology, an organism () is any living system that functions as an individual entity. All organisms are composed of cells ( cell theory). Organisms are classified by taxonomy into groups such as multicellular animals, plants, and fu ...
s evolved, from the earliest emergence of life to present day. Earth formed about 4.5 billion years ago (abbreviated as ''Ga'', for ''
gigaannum A year or annus is the orbital period of a planetary body, for example, the Earth, moving in its orbit around the Sun. Due to the Earth's axial tilt, the course of a year sees the passing of the seasons, marked by change in weather, the h ...
'') and evidence suggests that life emerged prior to 3.7 Ga. Although there is some evidence of life as early as 4.1 to 4.28 Ga, it remains controversial due to the possible non-biological formation of the purported fossils. The similarities among all known present-day
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriat ...
indicate that they have diverged through the process of
evolution Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation ...
from a common ancestor. Only a very small percentage of species have been identified: one estimate claims that Earth may have 1 trillion species. * However, only 1.75–1.8 million have been named "A version of this article appears in print on Nov. 9, 2014, Section SR, Page 6 of the New York edition with the headline: Prehistory's Brilliant Future." and 1.8 million documented in a central database. These currently living species represent less than one percent of all species that have ever lived on Earth. The earliest evidence of life comes from
biogenic A biogenic substance is a product made by or of life forms. While the term originally was specific to metabolite compounds that had toxic effects on other organisms, it has developed to encompass any constituents, secretions, and metabolites of p ...
carbon signatures and stromatolite fossils discovered in 3.7 billion-year-old metasedimentary rocks from western
Greenland Greenland ( kl, Kalaallit Nunaat, ; da, Grønland, ) is an island country in North America that is part of the Kingdom of Denmark. It is located between the Arctic and Atlantic oceans, east of the Canadian Arctic Archipelago. Greenland ...
. In 2015, possible "remains of biotic life" were found in 4.1 billion-year-old rocks in
Western Australia Western Australia (commonly abbreviated as WA) is a state of Australia occupying the western percent of the land area of Australia excluding external territories. It is bounded by the Indian Ocean to the north and west, the Southern Ocean to t ...
. In March 2017, putative evidence of possibly the oldest forms of life on Earth was reported in the form of fossilized
microorganism A microorganism, or microbe,, ''mikros'', "small") and ''organism'' from the el, ὀργανισμός, ''organismós'', "organism"). It is usually written as a single word but is sometimes hyphenated (''micro-organism''), especially in old ...
s discovered in
hydrothermal vent A hydrothermal vent is a fissure on the seabed from which geothermally heated water discharges. They are commonly found near volcanically active places, areas where tectonic plates are moving apart at mid-ocean ridges, ocean basins, and hotspo ...
precipitates in the Nuvvuagittuq Belt of Quebec, Canada, that may have lived as early as 4.28 billion years ago, not long after the oceans formed 4.4 billion years ago, and not long after the formation of the Earth 4.54 billion years ago. "A version of this article appears in print on March 2, 2017, Section A, Page 9 of the New York edition with the headline: Artful Squiggles in Rocks May Be Earth's Oldest Fossils." Microbial mats of coexisting
bacteria Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one biological cell. They constitute a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria were am ...
and
archaea Archaea ( ; singular archaeon ) is a domain of single-celled organisms. These microorganisms lack cell nuclei and are therefore prokaryotes. Archaea were initially classified as bacteria, receiving the name archaebacteria (in the Archaeba ...
were the dominant form of life in the early
Archean The Archean Eon ( , also spelled Archaean or Archæan) is the second of four geologic eons of Earth's history, representing the time from . The Archean was preceded by the Hadean Eon and followed by the Proterozoic. The Earth during the Arc ...
Epoch and many of the major steps in early evolution are thought to have taken place in this environment. The evolution of
photosynthesis Photosynthesis is a process used by plants and other organisms to convert light energy into chemical energy that, through cellular respiration, can later be released to fuel the organism's activities. Some of this chemical energy is stored in ...
, around 3.5 Ga, eventually led to a buildup of its waste product,
oxygen Oxygen is the chemical element with the symbol O and atomic number 8. It is a member of the chalcogen group in the periodic table, a highly reactive nonmetal, and an oxidizing agent that readily forms oxides with most elements ...
, in the atmosphere, leading to the
great oxygenation event The Great Oxidation Event (GOE), also called the Great Oxygenation Event, the Oxygen Catastrophe, the Oxygen Revolution, the Oxygen Crisis, or the Oxygen Holocaust, was a time interval during the Paleoproterozoic era when the Earth's atmospher ...
, beginning around 2.4 Ga. The earliest evidence of
eukaryote Eukaryotes () are organisms whose cells have a nucleus. All animals, plants, fungi, and many unicellular organisms, are Eukaryotes. They belong to the group of organisms Eukaryota or Eukarya, which is one of the three domains of life. Bacter ...
s (complex cells with
organelle In cell biology, an organelle is a specialized subunit, usually within a cell, that has a specific function. The name ''organelle'' comes from the idea that these structures are parts of cells, as organs are to the body, hence ''organelle,'' th ...
s) dates from 1.85 Ga, and while they may have been present earlier, their diversification accelerated when they started using oxygen in their
metabolism Metabolism (, from el, μεταβολή ''metabolē'', "change") is the set of life-sustaining chemical reactions in organisms. The three main functions of metabolism are: the conversion of the energy in food to energy available to run ...
. Later, around 1.7 Ga,
multicellular organism A multicellular organism is an organism that consists of more than one cell, in contrast to unicellular organism. All species of animals, land plants and most fungi are multicellular, as are many algae, whereas a few organisms are partially uni ...
s began to appear, with differentiated cells performing specialised functions.
Sexual reproduction Sexual reproduction is a type of reproduction that involves a complex life cycle in which a gamete ( haploid reproductive cells, such as a sperm or egg cell) with a single set of chromosomes combines with another gamete to produce a zygote th ...
, which involves the fusion of male and female reproductive cells (
gamete A gamete (; , ultimately ) is a haploid cell that fuses with another haploid cell during fertilization in organisms that reproduce sexually. Gametes are an organism's reproductive cells, also referred to as sex cells. In species that produce ...
s) to create a
zygote A zygote (, ) is a eukaryotic cell formed by a fertilization event between two gametes. The zygote's genome is a combination of the DNA in each gamete, and contains all of the genetic information of a new individual organism. In multicell ...
in a process called
fertilization Fertilisation or fertilization (see spelling differences), also known as generative fertilisation, syngamy and impregnation, is the fusion of gametes to give rise to a new individual organism or offspring and initiate its development. Pro ...
is, in contrast to
asexual reproduction Asexual reproduction is a type of reproduction that does not involve the fusion of gametes or change in the number of chromosomes. The offspring that arise by asexual reproduction from either unicellular or multicellular organisms inherit the ...
, the primary method of reproduction for the vast majority of macroscopic organisms, including almost all
eukaryotes Eukaryotes () are organisms whose cells have a nucleus. All animals, plants, fungi, and many unicellular organisms, are Eukaryotes. They belong to the group of organisms Eukaryota or Eukarya, which is one of the three domains of life. Bacter ...
(which includes
animal Animals are multicellular, eukaryotic organisms in the biological kingdom Animalia. With few exceptions, animals consume organic material, breathe oxygen, are able to move, can reproduce sexually, and go through an ontogenetic stage ...
s and
plant Plants are predominantly photosynthetic eukaryotes of the kingdom Plantae. Historically, the plant kingdom encompassed all living things that were not animals, and included algae and fungi; however, all current definitions of Plantae excl ...
s). However the origin and evolution of sexual reproduction remain a puzzle for biologists though it did evolve from a common ancestor that was a single celled eukaryotic species. * *
Bilateria The Bilateria or bilaterians are animals with bilateral symmetry as an embryo, i.e. having a left and a right side that are mirror images of each other. This also means they have a head and a tail (anterior-posterior axis) as well as a belly and ...
, animals having a left and a right side that are mirror images of each other, appeared by 555 Ma (million years ago). Algae-like multicellular land plants are dated back even to about 1 billion years ago, although evidence suggests that
microorganism A microorganism, or microbe,, ''mikros'', "small") and ''organism'' from the el, ὀργανισμός, ''organismós'', "organism"). It is usually written as a single word but is sometimes hyphenated (''micro-organism''), especially in old ...
s formed the earliest
terrestrial ecosystem Terrestrial ecosystems are ecosystems which are found on land. Examples include tundra, taiga, temperate deciduous forest, tropical rain forest, grassland, deserts. Terrestrial ecosystems differ from aquatic ecosystems by the predominant pres ...
s, at least 2.7 Ga. Microorganisms are thought to have paved the way for the inception of land plants in the
Ordovician The Ordovician ( ) is a geologic period and system, the second of six periods of the Paleozoic Era. The Ordovician spans 41.6 million years from the end of the Cambrian Period million years ago (Mya) to the start of the Silurian Period Mya. T ...
period. Land plants were so successful that they are thought to have contributed to the Late Devonian extinction event. (The long causal chain implied seems to involve (1) the success of early tree
archaeopteris ''Archaeopteris'' is an extinct genus of progymnosperm tree with fern-like leaves. A useful index fossil, this tree is found in strata dating from the Upper Devonian to Lower Carboniferous (), the oldest fossils being 385 million years old, ...
drew down CO2 levels, leading to global cooling and lowered sea levels, (2) roots of archeopteris fostered soil development which ''increased'' rock weathering, and the subsequent nutrient run-off may have triggered algal blooms resulting in anoxic events which caused marine-life die-offs. Marine species were the primary victims of the Late Devonian extinction.) Ediacara biota appear during the
Ediacaran The Ediacaran Period ( ) is a geological period that spans 96 million years from the end of the Cryogenian Period 635 million years ago (Mya), to the beginning of the Cambrian Period 538.8 Mya. It marks the end of the Proterozoic Eon, and t ...
period, while
vertebrate Vertebrates () comprise all animal taxa within the subphylum Vertebrata () ( chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, with ...
s, along with most other modern
phyla Phyla, the plural of ''phylum'', may refer to: * Phylum, a biological taxon between Kingdom and Class * by analogy, in linguistics, a large division of possibly related languages, or a major language family which is not subordinate to another Phy ...
originated about during the
Cambrian explosion The Cambrian explosion, Cambrian radiation, Cambrian diversification, or the Biological Big Bang refers to an interval of time approximately in the Cambrian Period when practically all major animal phyla started appearing in the fossil record. ...
. During the
Permian The Permian ( ) is a geologic period and stratigraphic system which spans 47 million years from the end of the Carboniferous Period million years ago (Mya), to the beginning of the Triassic Period 251.9 Mya. It is the last period of the Paleo ...
period, synapsids, including the ancestors of
mammal Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fur ...
s, dominated the land, but most of this group became extinct in the
Permian–Triassic extinction event The Permian–Triassic (P–T, P–Tr) extinction event, also known as the Latest Permian extinction event, the End-Permian Extinction and colloquially as the Great Dying, formed the boundary between the Permian and Triassic geologic periods, ...
. During the recovery from this catastrophe,
archosaur Archosauria () is a clade of diapsids, with birds and crocodilians as the only living representatives. Archosaurs are broadly classified as reptiles, in the cladistic sense of the term which includes birds. Extinct archosaurs include non-avia ...
s became the most abundant land vertebrates; one archosaur group, the
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the evolution of dinosaurs is t ...
s, dominated the
Jurassic The Jurassic ( ) is a geologic period and stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately Mya. The Jurassic constitutes the middle period of ...
and
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of ...
periods. After the
Cretaceous–Paleogene extinction event The Cretaceous–Paleogene (K–Pg) extinction event (also known as the Cretaceous–Tertiary extinction) was a sudden mass extinction of three-quarters of the plant and animal species on Earth, approximately 66 million years ago. With the ...
killed off the non-avian dinosaurs, mammals increased rapidly in size and diversity. Such
mass extinction An extinction event (also known as a mass extinction or biotic crisis) is a widespread and rapid decrease in the biodiversity on Earth. Such an event is identified by a sharp change in the diversity and abundance of multicellular organisms. I ...
s may have accelerated evolution by providing opportunities for new groups of organisms to diversify.


Earliest history of Earth

The oldest
meteorite A meteorite is a solid piece of debris from an object, such as a comet, asteroid, or meteoroid, that originates in outer space and survives its passage through the atmosphere to reach the surface of a planet or moon. When the original object ...
fragments found on Earth are about 4.54 billion years old; this, coupled primarily with the dating of ancient
lead Lead is a chemical element with the symbol Pb (from the Latin ) and atomic number 82. It is a heavy metal that is denser than most common materials. Lead is soft and malleable, and also has a relatively low melting point. When freshly cut, ...
deposits, has put the estimated age of Earth at around that time. * * The
Moon The Moon is Earth's only natural satellite. It is the fifth largest satellite in the Solar System and the largest and most massive relative to its parent planet, with a diameter about one-quarter that of Earth (comparable to the width of ...
has the same composition as Earth's crust but does not contain an
iron Iron () is a chemical element with symbol Fe (from la, ferrum) and atomic number 26. It is a metal that belongs to the first transition series and group 8 of the periodic table. It is, by mass, the most common element on Earth, right in ...
-rich core like the Earth's. Many scientists think that about 40 million years after the formation of Earth, it collided with a body the size of Mars, throwing crust material into the orbit that formed the Moon. Another
hypothesis A hypothesis (plural hypotheses) is a proposed explanation for a phenomenon. For a hypothesis to be a scientific hypothesis, the scientific method requires that one can test it. Scientists generally base scientific hypotheses on previous obse ...
is that the Earth and Moon started to coalesce at the same time but the Earth, having much stronger
gravity In physics, gravity () is a fundamental interaction which causes mutual attraction between all things with mass or energy. Gravity is, by far, the weakest of the four fundamental interactions, approximately 1038 times weaker than the stro ...
than the early Moon, attracted almost all the iron particles in the area. Until 2001, the oldest rocks found on Earth were about 3.8 billion years old, leading scientists to estimate that the Earth's surface had been molten until then. Accordingly, they named this part of Earth's history the
Hadean The Hadean ( ) is a geologic eon of Earth history preceding the Archean. On Earth, the Hadean began with the planet's formation about 4.54 billion years ago (although the start of the Hadean is defined as the age of the oldest solid materia ...
. However, analysis of
zircon Zircon () is a mineral belonging to the group of nesosilicates and is a source of the metal zirconium. Its chemical name is zirconium(IV) silicate, and its corresponding chemical formula is Zr SiO4. An empirical formula showing some of t ...
s formed 4.4 Ga indicates that Earth's crust solidified about 100 million years after the planet's formation and that the planet quickly acquired oceans and an
atmosphere An atmosphere () is a layer of gas or layers of gases that envelop a planet, and is held in place by the gravity of the planetary body. A planet retains an atmosphere when the gravity is great and the temperature of the atmosphere is low. A ...
, which may have been capable of supporting life. Evidence from the Moon indicates that from 4 to 3.8 Ga it suffered a Late Heavy Bombardment by debris that was left over from the formation of the
Solar System The Solar System Capitalization of the name varies. The International Astronomical Union, the authoritative body regarding astronomical nomenclature, specifies capitalizing the names of all individual astronomical objects but uses mixed "Solar ...
, and the Earth should have experienced an even heavier bombardment due to its stronger gravity. While there is no direct evidence of conditions on Earth 4 to 3.8 Ga, there is no reason to think that the Earth was not also affected by this late heavy bombardment. This event may well have stripped away any previous atmosphere and oceans; in this case gases and water from
comet A comet is an icy, small Solar System body that, when passing close to the Sun, warms and begins to release gases, a process that is called outgassing. This produces a visible atmosphere or coma, and sometimes also a tail. These phenomena ...
impacts may have contributed to their replacement, although outgassing from
volcano A volcano is a rupture in the crust of a planetary-mass object, such as Earth, that allows hot lava, volcanic ash, and gases to escape from a magma chamber below the surface. On Earth, volcanoes are most often found where tectonic plates ...
es on Earth would have supplied at least half. However, if subsurface microbial life had evolved by this point, it would have survived the bombardment.


Earliest evidence for life on Earth

The earliest identified organisms were minute and relatively featureless, and their fossils look like small rods that are very difficult to tell apart from structures that arise through abiotic physical processes. The oldest undisputed evidence of life on Earth, interpreted as fossilized bacteria, dates to 3 Ga. Other finds in rocks dated to about 3.5 Ga have been interpreted as bacteria, with
geochemical Geochemistry is the science that uses the tools and principles of chemistry to explain the mechanisms behind major geological systems such as the Earth's crust and its oceans. The realm of geochemistry extends beyond the Earth, encompassing the e ...
evidence also seeming to show the presence of life 3.8 Ga. However, these analyses were closely scrutinized, and non-biological processes were found which could produce all of the "signatures of life" that had been reported. While this does not prove that the structures found had a non-biological origin, they cannot be taken as clear evidence for the presence of life. Geochemical signatures from rocks deposited 3.4 Ga have been interpreted as evidence for life, although these statements have not been thoroughly examined by critics. Evidence for fossilized microorganisms considered to be 3.77 billion to 4.28 billion years old was found in the Nuvvuagittuq Greenstone Belt in Quebec, Canada, although the evidence is disputed as inconclusive.


Origins of life on Earth

Biologist A biologist is a scientist who conducts research in biology. Biologists are interested in studying life on Earth, whether it is an individual cell, a multicellular organism, or a community of interacting populations. They usually specialize ...
s reason that all living organisms on Earth must share a single last universal ancestor, because it would be virtually impossible that two or more separate lineages could have independently developed the many complex biochemical mechanisms common to all living organisms.


Independent emergence on Earth

Life on Earth is based on
carbon Carbon () is a chemical element with the symbol C and atomic number 6. It is nonmetallic and tetravalent—its atom making four electrons available to form covalent chemical bonds. It belongs to group 14 of the periodic table. Carbon ma ...
and
water Water (chemical formula ) is an inorganic, transparent, tasteless, odorless, and nearly colorless chemical substance, which is the main constituent of Earth's hydrosphere and the fluids of all known living organisms (in which it acts as ...
. Carbon provides stable frameworks for complex chemicals and can be easily extracted from the environment, especially from
carbon dioxide Carbon dioxide ( chemical formula ) is a chemical compound made up of molecules that each have one carbon atom covalently double bonded to two oxygen atoms. It is found in the gas state at room temperature. In the air, carbon dioxide is t ...
. There is no other chemical element whose properties are similar enough to carbon's to be called an analogue;
silicon Silicon is a chemical element with the symbol Si and atomic number 14. It is a hard, brittle crystalline solid with a blue-grey metallic luster, and is a tetravalent metalloid and semiconductor. It is a member of group 14 in the periodic ...
, the element directly below carbon on the
periodic table The periodic table, also known as the periodic table of the (chemical) elements, is a rows and columns arrangement of the chemical elements. It is widely used in chemistry, physics, and other sciences, and is generally seen as an icon of ch ...
, does not form very many complex stable molecules, and because most of its compounds are water-insoluble and because
silicon dioxide Silicon dioxide, also known as silica, is an oxide of silicon with the chemical formula , most commonly found in nature as quartz and in various living organisms. In many parts of the world, silica is the major constituent of sand. Silica is one ...
is a hard and abrasive solid in contrast to
carbon dioxide Carbon dioxide ( chemical formula ) is a chemical compound made up of molecules that each have one carbon atom covalently double bonded to two oxygen atoms. It is found in the gas state at room temperature. In the air, carbon dioxide is t ...
at temperatures associated with living things, it would be more difficult for organisms to extract. The elements
boron Boron is a chemical element with the symbol B and atomic number 5. In its crystalline form it is a brittle, dark, lustrous metalloid; in its amorphous form it is a brown powder. As the lightest element of the '' boron group'' it has t ...
and
phosphorus Phosphorus is a chemical element with the symbol P and atomic number 15. Elemental phosphorus exists in two major forms, white phosphorus and red phosphorus, but because it is highly reactive, phosphorus is never found as a free element on Ea ...
have more complex chemistries, but suffer from other limitations relative to carbon. Water is an excellent
solvent A solvent (s) (from the Latin '' solvō'', "loosen, untie, solve") is a substance that dissolves a solute, resulting in a solution. A solvent is usually a liquid but can also be a solid, a gas, or a supercritical fluid. Water is a solvent for ...
and has two other useful properties: the fact that ice floats enables aquatic organisms to survive beneath it in winter; and its molecules have electrically negative and positive ends, which enables it to form a wider range of compounds than other solvents can. Other good solvents, such as
ammonia Ammonia is an inorganic compound of nitrogen and hydrogen with the formula . A stable binary hydride, and the simplest pnictogen hydride, ammonia is a colourless gas with a distinct pungent smell. Biologically, it is a common nitrogenous ...
, are liquid only at such low temperatures that chemical reactions may be too slow to sustain life, and lack water's other advantages. Organisms based on
alternative biochemistry Hypothetical types of biochemistry are forms of biochemistry agreed to be scientifically viable but not proven to exist at this time. The kinds of living organisms currently known on Earth all use carbon compounds for basic structural and metabo ...
may, however, be possible on other planets. Research on how life might have emerged from non-living chemicals focuses on three possible starting points: self-replication, an organism's ability to produce offspring that are very similar to itself; metabolism, its ability to feed and repair itself; and external
cell membrane The cell membrane (also known as the plasma membrane (PM) or cytoplasmic membrane, and historically referred to as the plasmalemma) is a biological membrane that separates and protects the interior of all cells from the outside environment (t ...
s, which allow food to enter and waste products to leave, but exclude unwanted substances. Research on abiogenesis still has a long way to go, since theoretical and
empirical Empirical evidence for a proposition is evidence, i.e. what supports or counters this proposition, that is constituted by or accessible to sense experience or experimental procedure. Empirical evidence is of central importance to the sciences and ...
approaches are only beginning to make contact with each other.


Replication first: RNA world

Even the simplest members of the three modern domains of life use DNA to record their "recipes" and a complex array of RNA and
protein Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residues. Proteins perform a vast array of functions within organisms, including catalysing metabolic reactions, DNA replication, res ...
molecules to "read" these instructions and use them for growth, maintenance and self-replication. The discovery that some RNA molecules can catalyze both their own replication and the construction of proteins led to the hypothesis of earlier life-forms based entirely on RNA. These ribozymes could have formed an RNA world in which there were individuals but no species, as
mutation In biology, a mutation is an alteration in the nucleic acid sequence of the genome of an organism, virus, or extrachromosomal DNA. Viral genomes contain either DNA or RNA. Mutations result from errors during DNA or viral replication, m ...
s and horizontal gene transfers would have meant that offspring were likely to have different
genome In the fields of molecular biology and genetics, a genome is all the genetic information of an organism. It consists of nucleotide sequences of DNA (or RNA in RNA viruses). The nuclear genome includes protein-coding genes and non-coding ...
s from their parents, and
evolution Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation ...
occurred at the level of genes rather than organisms. RNA would later have been replaced by DNA, which can build longer, more stable genomes, strengthening heritability and expanding the capabilities of individual organisms. Ribozymes remain as the main components of
ribosome Ribosomes ( ) are macromolecular machines, found within all cells, that perform biological protein synthesis (mRNA translation). Ribosomes link amino acids together in the order specified by the codons of messenger RNA (mRNA) molecules to fo ...
s, the "protein factories" in modern cells. Evidence suggests the first RNA molecules formed on Earth prior to 4.17 Ga. Although short self-replicating RNA molecules have been artificially produced in laboratories, doubts have been raised about whether natural non-biological synthesis of RNA is possible. * * The earliest "ribozymes" may have been formed of simpler
nucleic acid Nucleic acids are biopolymers, macromolecules, essential to all known forms of life. They are composed of nucleotides, which are the monomers made of three components: a 5-carbon sugar, a phosphate group and a nitrogenous base. The two main ...
s such as PNA,
TNA TNA may refer to: Organisations * Tamil National Alliance, a political coalition in Sri Lanka * The National Alliance, a political party in Kenya * The National Archives (United Kingdom), a UK public body * Tonga Nurses' Association, a trade union ...
or GNA, which would have been replaced later by RNA. In 2003, it was proposed that porous metal sulfide precipitates would assist RNA synthesis at about and ocean-bottom pressures near
hydrothermal vent A hydrothermal vent is a fissure on the seabed from which geothermally heated water discharges. They are commonly found near volcanically active places, areas where tectonic plates are moving apart at mid-ocean ridges, ocean basins, and hotspo ...
s. Under this hypothesis,
lipid Lipids are a broad group of naturally-occurring molecules which includes fats, waxes, sterols, fat-soluble vitamins (such as vitamins A, D, E and K), monoglycerides, diglycerides, phospholipids, and others. The functions of lipids in ...
membranes would be the last major cell components to appear and, until then, the protocells would be confined to the pores.


Metabolism first: Iron–sulfur world

A series of experiments starting in 1997 showed that early stages in the formation of proteins from inorganic materials including
carbon monoxide Carbon monoxide ( chemical formula CO) is a colorless, poisonous, odorless, tasteless, flammable gas that is slightly less dense than air. Carbon monoxide consists of one carbon atom and one oxygen atom connected by a triple bond. It is the simpl ...
and
hydrogen sulfide Hydrogen sulfide is a chemical compound with the formula . It is a colorless chalcogen-hydride gas, and is poisonous, corrosive, and flammable, with trace amounts in ambient atmosphere having a characteristic foul odor of rotten eggs. The under ...
could be achieved by using
iron sulfide Iron sulfide or Iron sulphide can refer to range of chemical compounds composed of iron and sulfur. Minerals By increasing order of stability: * Iron(II) sulfide, FeS * Greigite, Fe3S4 (cubic) * Pyrrhotite, Fe1−xS (where x = 0 to 0.2) (mono ...
and nickel sulfide as
catalyst Catalysis () is the process of increasing the rate of a chemical reaction by adding a substance known as a catalyst (). Catalysts are not consumed in the reaction and remain unchanged after it. If the reaction is rapid and the catalyst recyc ...
s. Most of the steps required temperatures of about and moderate pressures, although one stage required and a pressure equivalent to that found under of rock. Hence it was suggested that self-sustaining synthesis of proteins could have occurred near hydrothermal vents.


Membranes first: Lipid world

It has been suggested that double-walled "bubbles" of lipids like those that form the external membranes of cells may have been an essential first step. Experiments that simulated the conditions of the early Earth have reported the formation of lipids, and these can spontaneously form liposomes, double-walled "bubbles," and then reproduce themselves. Although they are not intrinsically information-carriers as nucleic acids are, they would be subject to
natural selection Natural selection is the differential survival and reproduction of individuals due to differences in phenotype. It is a key mechanism of evolution, the change in the heritable traits characteristic of a population over generations. Cha ...
for longevity and reproduction. Nucleic acids such as RNA might then have formed more easily within the liposomes than outside.


The clay hypothesis

RNA is complex and there are doubts about whether it can be produced non-biologically in the wild. Some
clay Clay is a type of fine-grained natural soil material containing clay minerals (hydrous aluminium phyllosilicates, e.g. kaolin, Al2 Si2 O5( OH)4). Clays develop plasticity when wet, due to a molecular film of water surrounding the clay pa ...
s, notably montmorillonite, have properties that make them plausible accelerators for the emergence of an RNA world: they grow by self-replication of their
crystal A crystal or crystalline solid is a solid material whose constituents (such as atoms, molecules, or ions) are arranged in a highly ordered microscopic structure, forming a crystal lattice that extends in all directions. In addition, macro ...
line pattern; they are subject to an analogue of natural selection, as the clay "species" that grows fastest in a particular environment rapidly becomes dominant; and they can catalyze the formation of RNA molecules. Although this idea has not become the scientific consensus, it still has active supporters. Research in 2003 reported that montmorillonite could also accelerate the conversion of
fatty acid In chemistry, particularly in biochemistry, a fatty acid is a carboxylic acid with an aliphatic chain, which is either saturated or unsaturated. Most naturally occurring fatty acids have an unbranched chain of an even number of carbon atoms, f ...
s into "bubbles," and that the "bubbles" could encapsulate RNA attached to the clay. These "bubbles" can then grow by absorbing additional lipids and then divide. The formation of the earliest cells may have been aided by similar processes. A similar hypothesis presents self-replicating iron-rich clays as the progenitors of
nucleotide Nucleotides are organic molecules consisting of a nucleoside and a phosphate. They serve as monomeric units of the nucleic acid polymers – deoxyribonucleic acid (DNA) and ribonucleic acid (RNA), both of which are essential biomolecu ...
s, lipids and
amino acid Amino acids are organic compounds that contain both amino and carboxylic acid functional groups. Although hundreds of amino acids exist in nature, by far the most important are the alpha-amino acids, which comprise proteins. Only 22 alpha ...
s.


Prebiotic environments


Geothermal springs

Wet-dry cycles at geothermal springs are shown to solve the problem of hydrolysis and promote the polymerization and vesicle encapsulation of biopolymers. The temperatures of geothermal springs are suitable for biomolecules. Silica minerals and metal sulfides at these environments have photocatalytic properties to catalyze biomolecules. Solar UV exposure also promotes the synthesis of biomolecules like RNA nucleotides. An analysis of hydrothermal veins at a 3.5 Gya geothermal spring setting were found to have elements required for the origin of life, which are potassium, boron, hydrogen, sulfur, phosphorus, zinc, nitrogen, and oxygen. Mulkidjanian and colleagues find that such environments have identical ionic concentrations to the cytoplasm of modern cells. Fatty acids in acidic or slightly alkaline geothermal springs assemble into vesicles after wet-dry cycles as there is a lower concentration of ionic solutes at geothermal springs since they are freshwater environments, in contrast to seawater which has a higher concentration of ionic solutes. For organic compounds to be present at geothermal springs, they would have likely been transported by carbonaceous meteors. The molecules that fell from the meteors were then accumulated in geothermal springs. As for the evolutionary implications, freshwater heterotrophic cells that depended upon synthesized organic compounds later evolved photosynthesis because of the continuous exposure to sunlight as well as their cell walls with ion pumps to maintain their intracellular metabolism after they entered the oceans.


Deep sea hydrothermal vents

Catalytic mineral particles at these environments likely catalyzed organic compounds. Scientists simulated laboratory conditions that were identical to white smokers and successfully oligomerized RNA, measured to be 4 units long. Another experiment that replicated conditions also similar white smokers, with phospholipids present resulted in the assembly of vesicles. Exergonic reactions at hydrothermal vents are suggested to have been a source of free energy that promoted chemical reactions, synthesis of organic molecules, and are inducive to chemical gradients. In small rock pore systems, membranous structures between alkaline seawater and the acidic ocean would be conducive to proton gradients. Small mineral cavities or mineral gels could have been a compartment for abiogenic processes. A genomic analysis supports this hypothesis as they found 355 genes that likely traced to LUCA upon 6.1 million sequenced prokaryotic genes. They reconstruct LUCA as a thermophilic anaerobe with a Wood-Ljungdahl pathway, implying an origin of life at white smokers.


Life "seeded" from elsewhere

The Panspermia hypothesis does not explain how life arose originally, but simply examines the possibility of it coming from somewhere other than Earth. The idea that life on Earth was "seeded" from elsewhere in the Universe dates back at least to the Greek philosopher
Anaximander Anaximander (; grc-gre, Ἀναξίμανδρος ''Anaximandros''; ) was a pre-Socratic Greek philosopher who lived in Miletus,"Anaximander" in ''Chambers's Encyclopædia''. London: George Newnes, 1961, Vol. 1, p. 403. a city of Ionia (in mo ...
in the sixth century BCE. In the twentieth century it was proposed by the physical chemist Svante Arrhenius, by the
astronomer An astronomer is a scientist in the field of astronomy who focuses their studies on a specific question or field outside the scope of Earth. They observe astronomical objects such as stars, planets, moons, comets and galaxies – in either ...
s Fred Hoyle and
Chandra Wickramasinghe Nalin Chandra Wickramasinghe (born 20 January 1939) is a Sri Lankan-born British mathematician, astronomer and astrobiologist of Sinhalese ethnicity. His research interests include the interstellar medium, infrared astronomy, light scattering t ...
, and by
molecular biologist Molecular biology is the branch of biology that seeks to understand the molecular basis of biological activity in and between cells, including biomolecular synthesis, modification, mechanisms, and interactions. The study of chemical and physic ...
Francis Crick Francis Harry Compton Crick (8 June 1916 – 28 July 2004) was an English molecular biologist, biophysicist, and neuroscientist. He, James Watson, Rosalind Franklin, and Maurice Wilkins played crucial roles in deciphering the helical stru ...
and chemist Leslie Orgel. There are three main versions of the "seeded from elsewhere" hypothesis: from elsewhere in our Solar System via fragments knocked into space by a large
meteor A meteoroid () is a small rocky or metallic body in outer space. Meteoroids are defined as objects significantly smaller than asteroids, ranging in size from grains to objects up to a meter wide. Objects smaller than this are classified as mi ...
impact, in which case the most credible sources are
Mars Mars is the fourth planet from the Sun and the second-smallest planet in the Solar System, only being larger than Mercury. In the English language, Mars is named for the Roman god of war. Mars is a terrestrial planet with a thin at ...
and
Venus Venus is the second planet from the Sun. It is sometimes called Earth's "sister" or "twin" planet as it is almost as large and has a similar composition. As an interior planet to Earth, Venus (like Mercury) appears in Earth's sky never f ...
; by alien visitors, possibly as a result of accidental
contamination Contamination is the presence of a constituent, impurity, or some other undesirable element that spoils, corrupts, infects, makes unfit, or makes inferior a material, physical body, natural environment, workplace, etc. Types of contamination ...
by microorganisms that they brought with them; and from outside the Solar System but by natural means. Experiments in low Earth orbit, such as EXOSTACK, have demonstrated that some microorganism
spore In biology, a spore is a unit of sexual or asexual reproduction that may be adapted for dispersal and for survival, often for extended periods of time, in unfavourable conditions. Spores form part of the life cycles of many plants, algae, ...
s can survive the shock of being catapulted into space and some can survive exposure to outer space radiation for at least 5.7 years. Scientists are divided over the likelihood of life arising independently on Mars, or on other planets in our
galaxy A galaxy is a system of stars, stellar remnants, interstellar gas, dust, dark matter, bound together by gravity. The word is derived from the Greek ' (), literally 'milky', a reference to the Milky Way galaxy that contains the Solar System ...
.


Environmental and evolutionary impact of microbial mats

Microbial mats are multi-layered, multi-species colonies of bacteria and other organisms that are generally only a few millimeters thick, but still contain a wide range of chemical environments, each of which favors a different set of microorganisms. To some extent each mat forms its own food chain, as the by-products of each group of microorganisms generally serve as "food" for adjacent groups. Stromatolites are stubby pillars built as microorganisms in mats slowly migrate upwards to avoid being smothered by sediment deposited on them by water. There has been vigorous debate about the validity of alleged stromatolite fossils from before 3 Ga, with critics arguing that they could have been formed by non-biological processes. In 2006, another find of stromatolites was reported from the same part of Australia, in rocks dated to 3.5 Ga. In modern underwater mats the top layer often consists of photosynthesizing
cyanobacteria Cyanobacteria (), also known as Cyanophyta, are a phylum of gram-negative bacteria that obtain energy via photosynthesis. The name ''cyanobacteria'' refers to their color (), which similarly forms the basis of cyanobacteria's common name, bl ...
which create an oxygen-rich environment, while the bottom layer is oxygen-free and often dominated by hydrogen sulfide emitted by the organisms living there. Oxygen is toxic to organisms that are not adapted to it, but greatly increases the
metabolic Metabolism (, from el, μεταβολή ''metabolē'', "change") is the set of life-sustaining chemical reactions in organisms. The three main functions of metabolism are: the conversion of the energy in food to energy available to run cell ...
efficiency of oxygen-adapted organisms;
oxygenic photosynthesis Photosynthesis is a process used by plants and other organisms to convert light energy into chemical energy that, through cellular respiration, can later be released to fuel the organism's activities. Some of this chemical energy is stored in ...
by bacteria in mats increased biological productivity by a factor of between 100 and 1,000. The source of hydrogen atoms used by oxygenic photosynthesis is water, which is much more plentiful than the geologically produced reducing agents required by the earlier non-oxygenic photosynthesis. From this point onwards life itself produced significantly more of the resources it needed than did geochemical processes. Oxygen became a significant component of Earth's atmosphere about 2.4 Ga. Although eukaryotes may have been present much earlier, the oxygenation of the atmosphere was a prerequisite for the evolution of the most complex eukaryotic cells, from which all multicellular organisms are built. The boundary between oxygen-rich and oxygen-free layers in microbial mats would have moved upwards when photosynthesis shut down overnight, and then downwards as it resumed on the next day. This would have created selection pressure for organisms in this intermediate zone to acquire the ability to tolerate and then to use oxygen, possibly via endosymbiosis, where one organism lives inside another and both of them benefit from their association. Cyanobacteria have the most complete biochemical "toolkits" of all the mat-forming organisms. Hence they are the most self-sufficient, well-adapted to strike out on their own both as floating mats and as the first of the
phytoplankton Phytoplankton () are the autotrophic (self-feeding) components of the plankton community and a key part of ocean and freshwater ecosystems. The name comes from the Greek words (), meaning 'plant', and (), meaning 'wanderer' or 'drifter'. ...
, providing the basis of most marine food chains.


Diversification of eukaryotes

One possible family tree of eukaryotes *


Chromatin, nucleus, endomembrane system, and mitochondria

Eukaryotes may have been present long before the oxygenation of the atmosphere, but most modern eukaryotes require oxygen, which is used by their
mitochondria A mitochondrion (; ) is an organelle found in the cells of most Eukaryotes, such as animals, plants and fungi. Mitochondria have a double membrane structure and use aerobic respiration to generate adenosine triphosphate (ATP), which is used ...
to fuel the production of ATP, the internal energy supply of all known cells. In the 1970s, a vigorous debate concluded that eukaryotes emerged as a result of a sequence of endosymbiosis between
prokaryote A prokaryote () is a single-celled organism that lacks a nucleus and other membrane-bound organelles. The word ''prokaryote'' comes from the Greek πρό (, 'before') and κάρυον (, 'nut' or 'kernel').Campbell, N. "Biology:Concepts & Con ...
s. For example: a
predator Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill t ...
y microorganism invaded a large prokaryote, probably an
archaea Archaea ( ; singular archaeon ) is a domain of single-celled organisms. These microorganisms lack cell nuclei and are therefore prokaryotes. Archaea were initially classified as bacteria, receiving the name archaebacteria (in the Archaeba ...
n, but instead of killing its prey, the attacker took up residence and evolved into mitochondria; one of these chimeras later tried to swallow a photosynthesizing cyanobacterium, but the victim survived inside the attacker and the new combination became the ancestor of
plant Plants are predominantly photosynthetic eukaryotes of the kingdom Plantae. Historically, the plant kingdom encompassed all living things that were not animals, and included algae and fungi; however, all current definitions of Plantae excl ...
s; and so on. After each endosymbiosis, the partners eventually eliminated unproductive duplication of genetic functions by re-arranging their genomes, a process which sometimes involved transfer of genes between them. Another hypothesis proposes that mitochondria were originally
sulfur Sulfur (or sulphur in British English) is a chemical element with the symbol S and atomic number 16. It is abundant, multivalent and nonmetallic. Under normal conditions, sulfur atoms form cyclic octatomic molecules with a chemical formul ...
- or
hydrogen Hydrogen is the chemical element with the symbol H and atomic number 1. Hydrogen is the lightest element. At standard conditions hydrogen is a gas of diatomic molecules having the formula . It is colorless, odorless, tasteless, non-to ...
-metabolising endosymbionts, and became oxygen-consumers later. On the other hand, mitochondria might have been part of eukaryotes' original equipment. There is a debate about when eukaryotes first appeared: the presence of
sterane Sterane (cyclopentanoperhydrophenanthrenes) compounds are a class of tetracyclic compounds derived from steroids or sterols via diagenetic and catagenetic degradation and saturation. Steranes have an androstane skeleton with a side chain at carb ...
s in Australian
shale Shale is a fine-grained, clastic sedimentary rock formed from mud that is a mix of flakes of clay minerals (hydrous aluminium phyllosilicates, e.g. kaolin, Al2 Si2 O5( OH)4) and tiny fragments (silt-sized particles) of other minerals, especiall ...
s may indicate eukaryotes at 2.7 Ga; however, an analysis in 2008 concluded that these chemicals infiltrated the rocks less than 2.2 Ga and prove nothing about the origins of eukaryotes. Fossils of the
algae Algae (; singular alga ) is an informal term for a large and diverse group of photosynthetic eukaryotic organisms. It is a polyphyletic grouping that includes species from multiple distinct clades. Included organisms range from unicellular micr ...
'' Grypania'' have been reported in 1.85 billion-year-old rocks (originally dated to 2.1 Ga but later revised), indicating that eukaryotes with organelles had already evolved. A diverse collection of fossil algae were found in rocks dated between 1.5 and 1.4 Ga. The earliest known fossils of
fungi A fungus ( : fungi or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. These organisms are classified as a kingdom, separately fr ...
date from 1.43 Ga.


Plastids

Plastid The plastid (Greek: πλαστός; plastós: formed, molded – plural plastids) is a membrane-bound organelle found in the cells of plants, algae, and some other eukaryotic organisms. They are considered to be intracellular endosymbiotic cyan ...
s, the superclass of
organelle In cell biology, an organelle is a specialized subunit, usually within a cell, that has a specific function. The name ''organelle'' comes from the idea that these structures are parts of cells, as organs are to the body, hence ''organelle,'' th ...
s of which
chloroplast A chloroplast () is a type of membrane-bound organelle known as a plastid that conducts photosynthesis mostly in plant and algal cells. The photosynthetic pigment chlorophyll captures the energy from sunlight, converts it, and stores it i ...
s are the best-known exemplar, are thought to have originated from endosymbiotic cyanobacteria. The symbiosis evolved around 1.5 Ga and enabled eukaryotes to carry out
oxygenic photosynthesis Photosynthesis is a process used by plants and other organisms to convert light energy into chemical energy that, through cellular respiration, can later be released to fuel the organism's activities. Some of this chemical energy is stored in ...
. Three evolutionary lineages of photosynthetic plastids have since emerged: chloroplasts in
green algae The green algae (singular: green alga) are a group consisting of the Prasinodermophyta and its unnamed sister which contains the Chlorophyta and Charophyta/ Streptophyta. The land plants (Embryophytes) have emerged deep in the Charophyte alga ...
and plants, rhodoplasts in
red algae Red algae, or Rhodophyta (, ; ), are one of the oldest groups of eukaryotic algae. The Rhodophyta also comprises one of the largest phyla of algae, containing over 7,000 currently recognized species with taxonomic revisions ongoing. The majority ...
and cyanelles in the glaucophytes. Not long after this primary endosymbiosis of plastids, rhodoplasts and chloroplasts were passed down to other bikonts, establishing an eukaryotic assemblage of phytoplankton by the end of the Neoproterozoic Eon.


Sexual reproduction and multicellular organisms


Evolution of sexual reproduction

The defining characteristics of
sexual reproduction Sexual reproduction is a type of reproduction that involves a complex life cycle in which a gamete ( haploid reproductive cells, such as a sperm or egg cell) with a single set of chromosomes combines with another gamete to produce a zygote th ...
in eukaryotes are
meiosis Meiosis (; , since it is a reductional division) is a special type of cell division of germ cells in sexually-reproducing organisms that produces the gametes, such as sperm or egg cells. It involves two rounds of division that ultimately ...
and
fertilization Fertilisation or fertilization (see spelling differences), also known as generative fertilisation, syngamy and impregnation, is the fusion of gametes to give rise to a new individual organism or offspring and initiate its development. Pro ...
, resulting in
genetic recombination Genetic recombination (also known as genetic reshuffling) is the exchange of genetic material between different organisms which leads to production of offspring with combinations of traits that differ from those found in either parent. In eukaryo ...
, giving offspring 50% of their genes from each parent. By contrast, in
asexual reproduction Asexual reproduction is a type of reproduction that does not involve the fusion of gametes or change in the number of chromosomes. The offspring that arise by asexual reproduction from either unicellular or multicellular organisms inherit the ...
there is no recombination, but occasional horizontal gene transfer. Bacteria also exchange DNA by
bacterial conjugation Bacterial conjugation is the transfer of genetic material between bacterial cells by direct cell-to-cell contact or by a bridge-like connection between two cells. This takes place through a pilus. It is a parasexual mode of reproduction in bacter ...
, enabling the spread of resistance to
antibiotic An antibiotic is a type of antimicrobial substance active against bacteria. It is the most important type of antibacterial agent for fighting bacterial infections, and antibiotic medications are widely used in the treatment and prevention ...
s and other
toxin A toxin is a naturally occurring organic poison produced by metabolic activities of living cells or organisms. Toxins occur especially as a protein or conjugated protein. The term toxin was first used by organic chemist Ludwig Brieger (1849 ...
s, and the ability to utilize new
metabolite In biochemistry, a metabolite is an intermediate or end product of metabolism. The term is usually used for small molecules. Metabolites have various functions, including fuel, structure, signaling, stimulatory and inhibitory effects on enzymes, ...
s. However, conjugation is not a means of reproduction, and is not limited to members of the same species – there are cases where bacteria transfer DNA to plants and animals. On the other hand, bacterial transformation is clearly an adaptation for transfer of DNA between bacteria of the same species. This is a complex process involving the products of numerous bacterial genes and can be regarded as a bacterial form of sex. This process occurs naturally in at least 67 prokaryotic species (in seven different phyla). Sexual reproduction in eukaryotes may have evolved from bacterial transformation. The disadvantages of sexual reproduction are well-known: the genetic reshuffle of recombination may break up favorable combinations of genes; and since males do not directly increase the number of offspring in the next generation, an asexual population can out-breed and displace in as little as 50 generations a sexual population that is equal in every other respect. Nevertheless, the great majority of animals, plants, fungi and
protist A protist () is any eukaryotic organism (that is, an organism whose cells contain a cell nucleus) that is not an animal, plant, or fungus. While it is likely that protists share a common ancestor (the last eukaryotic common ancestor), the e ...
s reproduce sexually. There is strong evidence that sexual reproduction arose early in the history of eukaryotes and that the genes controlling it have changed very little since then. How sexual reproduction evolved and survived is an unsolved puzzle. The Red Queen hypothesis suggests that sexual reproduction provides protection against
parasite Parasitism is a close relationship between species, where one organism, the parasite, lives on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The entomologist E. O. Wilson h ...
s, because it is easier for parasites to evolve means of overcoming the defenses of genetically identical clones than those of sexual species that present moving targets, and there is some experimental evidence for this. However, there is still doubt about whether it would explain the survival of sexual species if multiple similar clone species were present, as one of the clones may survive the attacks of parasites for long enough to out-breed the sexual species. Furthermore, contrary to the expectations of the Red Queen hypothesis, Kathryn A. Hanley et al. found that the prevalence, abundance and mean intensity of mites was significantly higher in sexual geckos than in asexuals sharing the same habitat. In addition, biologist Matthew Parker, after reviewing numerous genetic studies on plant disease resistance, failed to find a single example consistent with the concept that pathogens are the primary selective agent responsible for sexual reproduction in the host. Alexey Kondrashov's ''deterministic mutation hypothesis'' (DMH) assumes that each organism has more than one harmful mutation and the combined effects of these mutations are more harmful than the sum of the harm done by each individual mutation. If so, sexual recombination of genes will reduce the harm that bad mutations do to offspring and at the same time eliminate some bad mutations from the
gene pool The gene pool is the set of all genes, or genetic information, in any population, usually of a particular species. Description A large gene pool indicates extensive genetic diversity, which is associated with robust populations that can surv ...
by isolating them in individuals that perish quickly because they have an above-average number of bad mutations. However, the evidence suggests that the DMH's assumptions are shaky because many species have on average less than one harmful mutation per individual and no species that has been investigated shows evidence of
synergy Synergy is an interaction or cooperation giving rise to a whole that is greater than the simple sum of its parts. The term ''synergy'' comes from the Attic Greek word συνεργία ' from ', , meaning "working together". History In Christian ...
between harmful mutations. The random nature of recombination causes the relative abundance of alternative traits to vary from one generation to another. This
genetic drift Genetic drift, also known as allelic drift or the Wright effect, is the change in the frequency of an existing gene variant (allele) in a population due to random chance. Genetic drift may cause gene variants to disappear completely and there ...
is insufficient on its own to make sexual reproduction advantageous, but a combination of genetic drift and natural selection may be sufficient. When chance produces combinations of good traits, natural selection gives a large advantage to lineages in which these traits become genetically linked. On the other hand, the benefits of good traits are neutralized if they appear along with bad traits. Sexual recombination gives good traits the opportunities to become linked with other good traits, and mathematical models suggest this may be more than enough to offset the disadvantages of sexual reproduction. Other combinations of hypotheses that are inadequate on their own are also being examined. The adaptive function of sex remains a major unresolved issue in biology. The competing models to explain it were reviewed by John A. Birdsell and Christopher Wills. The hypotheses discussed above all depend on the possible beneficial effects of random genetic variation produced by genetic recombination. An alternative view is that sex arose and is maintained as a process for repairing DNA damage, and that the genetic variation produced is an occasionally beneficial byproduct.


Multicellularity

The simplest definitions of "multicellular," for example "having multiple cells," could include colonial cyanobacteria like ''
Nostoc ''Nostoc'', also known as star jelly, troll’s butter, spit of moon, fallen star, witch's butter (not to be confused with the fungi commonly known as witches' butter), and witch’s jelly, is the most common genus of cyanobacteria found in var ...
''. Even a technical definition such as "having the same genome but different types of cell" would still include some
genera Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nomenclat ...
of the green algae Volvox, which have cells that specialize in reproduction. Multicellularity evolved independently in organisms as diverse as
sponge Sponges, the members of the phylum Porifera (; meaning 'pore bearer'), are a basal animal clade as a sister of the diploblasts. They are multicellular organisms that have bodies full of pores and channels allowing water to circulate throu ...
s and other animals, fungi, plants,
brown algae Brown algae (singular: alga), comprising the class Phaeophyceae, are a large group of multicellular algae, including many seaweeds located in colder waters within the Northern Hemisphere. Brown algae are the major seaweeds of the temperate and p ...
, cyanobacteria, slime molds and myxobacteria. For the sake of brevity, this article focuses on the organisms that show the greatest specialization of cells and variety of cell types, although this approach to the evolution of biological complexity could be regarded as "rather anthropocentric." The initial advantages of multicellularity may have included: more efficient sharing of nutrients that are digested outside the cell, increased resistance to predators, many of which attacked by engulfing; the ability to resist currents by attaching to a firm surface; the ability to reach upwards to filter-feed or to obtain sunlight for photosynthesis; the ability to create an internal environment that gives protection against the external one; and even the opportunity for a group of cells to behave "intelligently" by sharing information. These features would also have provided opportunities for other organisms to diversify, by creating more varied environments than flat microbial mats could. Multicellularity with differentiated cells is beneficial to the organism as a whole but disadvantageous from the point of view of individual cells, most of which lose the opportunity to reproduce themselves. In an asexual multicellular organism, rogue cells which retain the ability to reproduce may take over and reduce the organism to a mass of undifferentiated cells. Sexual reproduction eliminates such rogue cells from the next generation and therefore appears to be a prerequisite for complex multicellularity. The available evidence indicates that eukaryotes evolved much earlier but remained inconspicuous until a rapid diversification around 1 Ga. The only respect in which eukaryotes clearly surpass bacteria and archaea is their capacity for variety of forms, and sexual reproduction enabled eukaryotes to exploit that advantage by producing organisms with multiple cells that differed in form and function. By comparing the composition of transcription factor families and regulatory network motifs between unicellular organisms and multicellular organisms, scientists found there are many novel transcription factor families and three novel types of regulatory network motifs in multicellular organisms, and novel family transcription factors are preferentially wired into these novel network motifs which are essential for multicullular development. These results propose a plausible mechanism for the contribution of novel-family transcription factors and novel network motifs to the origin of multicellular organisms at transcriptional regulatory level.


Fossil evidence

The
Francevillian biota The Francevillian biota (also known as Gabon macrofossils or Gabonionta) is a group of 2.1-billion-year-old Palaeoproterozoic, macroscopic organisms known from fossils found in Gabon in the Palaeoproterozoic Francevillian B Formation, a black s ...
fossils, dated to 2.1 Ga, are the earliest known fossil organisms that are clearly multicellular. They may have had differentiated cells. Another early multicellular fossil, '' Qingshania'', dated to 1.7 Ga, appears to consist of virtually identical cells. The red algae called ''
Bangiomorpha ''Bangiomorpha pubescens'' is a red alga. It is the first known sexually reproducing organism. A multicellular fossil of ''Bangiomorpha pubescens'' was recovered from the Hunting Formation in Somerset Island, Canada that strongly resembles the ...
'', dated at 1.2 Ga, is the earliest known organism that certainly has differentiated, specialized cells, and is also the oldest known sexually reproducing organism. The 1.43 billion-year-old fossils interpreted as fungi appear to have been multicellular with differentiated cells. The "string of beads" organism '' Horodyskia'', found in rocks dated from 1.5 Ga to 900 Ma, may have been an early metazoan; however, it has also been interpreted as a colonial
foraminifera Foraminifera (; Latin for "hole bearers"; informally called "forams") are single-celled organisms, members of a phylum or class of amoeboid protists characterized by streaming granular ectoplasm for catching food and other uses; and commonly ...
n.


Emergence of animals

A family tree of the animals
Animals are multicellular eukaryotes, Myxozoa were thought to be an exception, but are now thought to be heavily modified members of the
Cnidaria Cnidaria () is a phylum under kingdom Animalia containing over 11,000 species of aquatic animals found both in freshwater and marine environments, predominantly the latter. Their distinguishing feature is cnidocytes, specialized cells that ...
.
and are distinguished from plants, algae, and fungi by lacking
cell wall A cell wall is a structural layer surrounding some types of cells, just outside the cell membrane. It can be tough, flexible, and sometimes rigid. It provides the cell with both structural support and protection, and also acts as a filtering mec ...
s. All animals are motile, if only at certain life stages. All animals except sponges have bodies differentiated into separate tissues, including
muscle Skeletal muscles (commonly referred to as muscles) are organs of the vertebrate muscular system and typically are attached by tendons to bones of a skeleton. The muscle cells of skeletal muscles are much longer than in the other types of mus ...
s, which move parts of the animal by contracting, and nerve tissue, which transmits and processes signals. In November 2019, researchers reported the discovery of '' Caveasphaera'', a
multicellular organism A multicellular organism is an organism that consists of more than one cell, in contrast to unicellular organism. All species of animals, land plants and most fungi are multicellular, as are many algae, whereas a few organisms are partially uni ...
found in 609-million-year-old rocks, that is not easily defined as an animal or non-animal, which may be related to one of the earliest instances of
animal evolution Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation ...
. Fossil studies of ''Caveasphaera'' have suggested that animal-like embryonic development arose much earlier than the oldest clearly defined animal fossils. and may be consistent with studies suggesting that animal evolution may have begun about 750 million years ago. Nonetheless, the earliest widely accepted animal fossils are the rather modern-looking
cnidaria Cnidaria () is a phylum under kingdom Animalia containing over 11,000 species of aquatic animals found both in freshwater and marine environments, predominantly the latter. Their distinguishing feature is cnidocytes, specialized cells that ...
ns (the group that includes
jellyfish Jellyfish and sea jellies are the informal common names given to the medusa-phase of certain gelatinous members of the subphylum Medusozoa, a major part of the phylum Cnidaria. Jellyfish are mainly free-swimming marine animals with umbre ...
,
sea anemone Sea anemones are a group of predatory marine invertebrates of the order Actiniaria. Because of their colourful appearance, they are named after the '' Anemone'', a terrestrial flowering plant. Sea anemones are classified in the phylum Cnidaria, ...
s and '' Hydra''), possibly from around , although fossils from the Doushantuo Formation can only be dated approximately. Their presence implies that the cnidarian and
bilateria The Bilateria or bilaterians are animals with bilateral symmetry as an embryo, i.e. having a left and a right side that are mirror images of each other. This also means they have a head and a tail (anterior-posterior axis) as well as a belly and ...
n lineages had already diverged. The Ediacara biota, which flourished for the last 40 million years before the start of the
Cambrian The Cambrian Period ( ; sometimes symbolized Ꞓ) was the first geological period of the Paleozoic Era, and of the Phanerozoic Eon. The Cambrian lasted 53.4 million years from the end of the preceding Ediacaran Period 538.8 million years ago ...
, were the first animals more than a very few centimetres long. Many were flat and had a "quilted" appearance, and seemed so strange that there was a proposal to classify them as a separate kingdom, Vendozoa. Others, however, have been interpreted as early
mollusc Mollusca is the second-largest phylum of invertebrate animals after the Arthropoda, the members of which are known as molluscs or mollusks (). Around 85,000  extant species of molluscs are recognized. The number of fossil species is est ...
s ('' Kimberella''),
echinoderm An echinoderm () is any member of the phylum Echinodermata (). The adults are recognisable by their (usually five-point) radial symmetry, and include starfish, brittle stars, sea urchins, sand dollars, and sea cucumbers, as well as the ...
s ('' Arkarua''), and
arthropod Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and cuticle made of chiti ...
s (''
Spriggina ''Spriggina'' is a genus of early bilaterian animals whose relationship to living animals is unclear. Fossils of ''Spriggina'' are known from the late Ediacaran period in what is now South Australia. ''Spriggina floundersi'' is the official fo ...
'', '' Parvancorina''). There is still debate about the classification of these specimens, mainly because the diagnostic features which allow taxonomists to classify more recent organisms, such as similarities to living organisms, are generally absent in the Ediacarans. However, there seems little doubt that ''Kimberella'' was at least a
triploblastic Triploblasty is a condition of the gastrula in which there are three primary germ layers: the ectoderm, mesoderm, and endoderm. Germ cells are set aside in the embryo at the blastula stage, which are incorporated into the gonads during organo ...
bilaterian animal, in other words, an animal significantly more complex than the cnidarians. The small shelly fauna are a very mixed collection of fossils found between the Late Ediacaran and
Middle Cambrian Middle or The Middle may refer to: * Centre (geometry), the point equally distant from the outer limits. Places * Middle (sheading), a subdivision of the Isle of Man * Middle Bay (disambiguation) * Middle Brook (disambiguation) * Middle Creek ( ...
periods. The earliest, '' Cloudina'', shows signs of successful defense against predation and may indicate the start of an evolutionary arms race. Some tiny Early Cambrian shells almost certainly belonged to molluscs, while the owners of some "armor plates," '' Halkieria'' and '' Microdictyon'', were eventually identified when more complete specimens were found in Cambrian
lagerstätte A Lagerstätte (, from '' Lager'' 'storage, lair' '' Stätte'' 'place'; plural ''Lagerstätten'') is a sedimentary deposit that exhibits extraordinary fossils with exceptional preservation—sometimes including preserved soft tissues. These f ...
n that preserved soft-bodied animals. In the 1970s there was already a debate about whether the emergence of the modern phyla was "explosive" or gradual but hidden by the shortage of
Precambrian The Precambrian (or Pre-Cambrian, sometimes abbreviated pꞒ, or Cryptozoic) is the earliest part of Earth's history, set before the current Phanerozoic Eon. The Precambrian is so named because it preceded the Cambrian, the first period of th ...
animal fossils. A re-analysis of fossils from the Burgess Shale lagerstätte increased interest in the issue when it revealed animals, such as '' Opabinia'', which did not fit into any known
phylum In biology, a phylum (; plural: phyla) is a level of classification or taxonomic rank below kingdom and above class. Traditionally, in botany the term division has been used instead of phylum, although the International Code of Nomenclature ...
. At the time these were interpreted as evidence that the modern phyla had evolved very rapidly in the Cambrian explosion and that the Burgess Shale's "weird wonders" showed that the Early Cambrian was a uniquely experimental period of animal evolution. Later discoveries of similar animals and the development of new theoretical approaches led to the conclusion that many of the "weird wonders" were evolutionary "aunts" or "cousins" of modern groups—for example that ''Opabinia'' was a member of the lobopods, a group which includes the ancestors of the arthropods, and that it may have been closely related to the modern
tardigrade Tardigrades (), known colloquially as water bears or moss piglets, are a phylum of eight-legged segmented micro-animals. They were first described by the German zoologist Johann August Ephraim Goeze in 1773, who called them Kleiner Wasserb� ...
s. Nevertheless, there is still much debate about whether the Cambrian explosion was really explosive and, if so, how and why it happened and why it appears unique in the history of animals.


Deuterostomes and the first vertebrates

Most of the animals at the heart of the Cambrian explosion debate are
protostome Protostomia () is the clade of animals once thought to be characterized by the formation of the organism's mouth before its anus during embryonic development. This nature has since been discovered to be extremely variable among Protostomia's me ...
s, one of the two main groups of complex animals. The other major group, the
deuterostome Deuterostomia (; in Greek) are animals typically characterized by their anus forming before their mouth during embryonic development. The group's sister clade is Protostomia, animals whose digestive tract development is more varied. Some ...
s, contains
invertebrate Invertebrates are a paraphyletic group of animals that neither possess nor develop a vertebral column (commonly known as a ''backbone'' or ''spine''), derived from the notochord. This is a grouping including all animals apart from the chorda ...
s such as
starfish Starfish or sea stars are star-shaped echinoderms belonging to the class Asteroidea (). Common usage frequently finds these names being also applied to ophiuroids, which are correctly referred to as brittle stars or basket stars. Starfish a ...
and
sea urchin Sea urchins () are spiny, globular echinoderms in the class Echinoidea. About 950 species of sea urchin live on the seabed of every ocean and inhabit every depth zone from the intertidal seashore down to . The spherical, hard shells (tests) o ...
s (echinoderms), as well as
chordate A chordate () is an animal of the phylum Chordata (). All chordates possess, at some point during their larval or adult stages, five synapomorphies, or primary physical characteristics, that distinguish them from all the other taxa. These fi ...
s (see below). Many echinoderms have hard
calcite Calcite is a carbonate mineral and the most stable polymorph of calcium carbonate (CaCO3). It is a very common mineral, particularly as a component of limestone. Calcite defines hardness 3 on the Mohs scale of mineral hardness, based on scratc ...
"shells," which are fairly common from the Early Cambrian small shelly fauna onwards. Other deuterostome groups are soft-bodied, and most of the significant Cambrian deuterostome fossils come from the Chengjiang fauna, a lagerstätte in
China China, officially the People's Republic of China (PRC), is a country in East Asia. It is the world's List of countries and dependencies by population, most populous country, with a Population of China, population exceeding 1.4 billion, slig ...
. The chordates are another major deuterostome group: animals with a distinct dorsal nerve cord. Chordates include soft-bodied invertebrates such as
tunicate A tunicate is a marine invertebrate animal, a member of the subphylum Tunicata (). It is part of the Chordata, a phylum which includes all animals with dorsal nerve cords and notochords (including vertebrates). The subphylum was at one t ...
s as well as vertebrates—animals with a backbone. While tunicate fossils predate the Cambrian explosion, the Chengjiang fossils ''
Haikouichthys ''Haikouichthys'' is an extinct genus of craniate (animals with notochords and distinct heads) that lived 518 million years ago, during the Cambrian explosion of multicellular life. ''Haikouichthys'' had a defined skull and other characteristi ...
'' and ''
Myllokunmingia ''Myllokunmingia'' is a genus of basal chordate from the Lower Cambrian Maotianshan shales of China 518 to 490 mya and is thought to be a vertebrate, although this is not conclusively proven. The species M. fengjiaoa is 28 mm long and 6&nbs ...
'' appear to be true vertebrates, and ''Haikouichthys'' had distinct
vertebrae The spinal column, a defining synapomorphy shared by nearly all vertebrates, Hagfish are believed to have secondarily lost their spinal column is a moderately flexible series of vertebrae (singular vertebra), each constituting a characteristi ...
, which may have been slightly mineralized. Vertebrates with jaws, such as the acanthodians, first appeared in the Late
Ordovician The Ordovician ( ) is a geologic period and system, the second of six periods of the Paleozoic Era. The Ordovician spans 41.6 million years from the end of the Cambrian Period million years ago (Mya) to the start of the Silurian Period Mya. T ...
.


Colonization of land

Adaptation to life on land is a major challenge: all land organisms need to avoid drying-out and all those above microscopic size must create special structures to withstand gravity; respiration and
gas exchange Gas exchange is the physical process by which gases move passively by diffusion across a surface. For example, this surface might be the air/water interface of a water body, the surface of a gas bubble in a liquid, a gas-permeable membrane, or a ...
systems have to change; reproductive systems cannot depend on water to carry eggs and
sperm Sperm is the male reproductive cell, or gamete, in anisogamous forms of sexual reproduction (forms in which there is a larger, female reproductive cell and a smaller, male one). Animals produce motile sperm with a tail known as a flagellum, ...
towards each other. Although the earliest good evidence of land plants and animals dates back to the Ordovician period (), and a number of microorganism lineages made it onto land much earlier, modern land
ecosystem An ecosystem (or ecological system) consists of all the organisms and the physical environment with which they interact. These biotic and abiotic components are linked together through nutrient cycles and energy flows. Energy enters the syst ...
s only appeared in the Late
Devonian The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, million years ago (Mya), to the beginning of the Carboniferous, Mya. It is named after Devon, England, wh ...
, about . In May 2017, evidence of the earliest known life on land may have been found in 3.48-billion-year-old geyserite and other related mineral deposits (often found around
hot spring A hot spring, hydrothermal spring, or geothermal spring is a spring produced by the emergence of geothermally heated groundwater onto the surface of the Earth. The groundwater is heated either by shallow bodies of magma (molten rock) or by circ ...
s and
geyser A geyser (, ) is a spring characterized by an intermittent discharge of water ejected turbulently and accompanied by steam. As a fairly rare phenomenon, the formation of geysers is due to particular hydrogeological conditions that exist only i ...
s) uncovered in the Pilbara Craton of
Western Australia Western Australia (commonly abbreviated as WA) is a state of Australia occupying the western percent of the land area of Australia excluding external territories. It is bounded by the Indian Ocean to the north and west, the Southern Ocean to t ...
. In July 2018, scientists reported that the earliest life on land may have been
bacteria Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one biological cell. They constitute a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria were am ...
living on land 3.22 billion years ago. In May 2019, scientists reported the discovery of a
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
ized
fungus A fungus ( : fungi or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. These organisms are classified as a kingdom, separately fr ...
, named ''
Ourasphaira giraldae ''Ourasphaira giraldae'' is an extinct process-bearing multicellular eukaryotic microorganism. Corentin Loron argues that it was an early fungus. It existed approximately a billion years ago during the time of the transition from the Mesoprotero ...
'', in the
Canadian Arctic Northern Canada, colloquially the North or the Territories, is the vast northernmost region of Canada variously defined by geography and politics. Politically, the term refers to the three territories of Canada: Yukon, Northwest Territories and ...
, that may have grown on land a billion years ago, well before
plant Plants are predominantly photosynthetic eukaryotes of the kingdom Plantae. Historically, the plant kingdom encompassed all living things that were not animals, and included algae and fungi; however, all current definitions of Plantae excl ...
s were living on land. "A version of this article appears in print on May 28, 2019, Section D, Page 3 of the New York edition with the headline: Finding a Clue to Life's Arrival."


Evolution of terrestrial antioxidants

Oxygen began to accumulate in Earth's atmosphere over 3 Ga, as a by-product of
photosynthesis Photosynthesis is a process used by plants and other organisms to convert light energy into chemical energy that, through cellular respiration, can later be released to fuel the organism's activities. Some of this chemical energy is stored in ...
in cyanobacteria (blue-green algae). However, oxygen produces destructive chemical oxydation which was toxic to most previous organisms. Protective endogenous antioxidant enzymes and exogenous dietary antioxidants helped to prevent oxidative damage. For example, brown algae accumulate inorganic mineral
antioxidant Antioxidants are compounds that inhibit oxidation, a chemical reaction that can produce free radicals. This can lead to polymerization and other chain reactions. They are frequently added to industrial products, such as fuels and lubrica ...
s such as
rubidium Rubidium is the chemical element with the symbol Rb and atomic number 37. It is a very soft, whitish-grey solid in the alkali metal group, similar to potassium and caesium. Rubidium is the first alkali metal in the group to have a density higher ...
,
vanadium Vanadium is a chemical element with the symbol V and atomic number 23. It is a hard, silvery-grey, malleable transition metal. The elemental metal is rarely found in nature, but once isolated artificially, the formation of an oxide layer ( pass ...
,
zinc Zinc is a chemical element with the symbol Zn and atomic number 30. Zinc is a slightly brittle metal at room temperature and has a shiny-greyish appearance when oxidation is removed. It is the first element in group 12 (IIB) of the periodi ...
, iron,
copper Copper is a chemical element with the symbol Cu (from la, cuprum) and atomic number 29. It is a soft, malleable, and ductile metal with very high thermal and electrical conductivity. A freshly exposed surface of pure copper has a pink ...
,
molybdenum Molybdenum is a chemical element with the symbol Mo and atomic number 42 which is located in period 5 and group 6. The name is from Neo-Latin ''molybdaenum'', which is based on Ancient Greek ', meaning lead, since its ores were confused with lead ...
, selenium and
iodine Iodine is a chemical element with the Symbol (chemistry), symbol I and atomic number 53. The heaviest of the stable halogens, it exists as a semi-lustrous, non-metallic solid at standard conditions that melts to form a deep violet liquid at , ...
, concentrated more than 30,000 times more than in seawater. Most marine mineral antioxidants act in the cells as essential trace elements in
redox Redox (reduction–oxidation, , ) is a type of chemical reaction in which the oxidation states of substrate change. Oxidation is the loss of electrons or an increase in the oxidation state, while reduction is the gain of electrons or ...
and antioxidant metalloenzymes. When plants and animals began to enter rivers and land about 500 Ma, environmental deficiency of these marine mineral antioxidants was a challenge to the evolution of terrestrial life. Terrestrial plants slowly optimized the production of new endogenous antioxidants such as
ascorbic acid Vitamin C (also known as ascorbic acid and ascorbate) is a water-soluble vitamin found in citrus and other fruits and vegetables, also sold as a dietary supplement and as a topical 'serum' ingredient to treat melasma (dark pigment spots) ...
,
polyphenol Polyphenols () are a large family of naturally occurring organic compounds characterized by multiples of phenol units. They are abundant in plants and structurally diverse. Polyphenols include flavonoids, tannic acid, and ellagitannin, some ...
s,
flavonoid Flavonoids (or bioflavonoids; from the Latin word ''flavus'', meaning yellow, their color in nature) are a class of polyphenolic secondary metabolites found in plants, and thus commonly consumed in the diets of humans. Chemically, flavonoids ...
s, tocopherols, etc. A few of these appeared more recently, in last 200–50 Ma, in
fruit In botany, a fruit is the seed-bearing structure in flowering plants that is formed from the ovary after flowering. Fruits are the means by which flowering plants (also known as angiosperms) disseminate their seeds. Edible fruits in partic ...
s and
flower A flower, sometimes known as a bloom or blossom, is the reproductive structure found in flowering plants (plants of the division Angiospermae). The biological function of a flower is to facilitate reproduction, usually by providing a mechanis ...
s of
angiosperm Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants ...
plants. In fact, angiosperms (the dominant type of plant today) and most of their antioxidant pigments evolved during the Late Jurassic period. Plants employ antioxidants to defend their structures against reactive oxygen species produced during photosynthesis. Animals are exposed to the same oxidants, and they have evolved endogenous enzymatic antioxidant systems.
Iodine Iodine is a chemical element with the Symbol (chemistry), symbol I and atomic number 53. The heaviest of the stable halogens, it exists as a semi-lustrous, non-metallic solid at standard conditions that melts to form a deep violet liquid at , ...
in the form of the iodide ion I- is the most primitive and abundant electron-rich essential element in the diet of marine and terrestrial organisms; it acts as an electron donor and has this ancestral antioxidant function in all iodide-concentrating cells, from primitive marine algae to terrestrial vertebrates.


Evolution of soil

Before the colonization of land there was no
soil Soil, also commonly referred to as earth or dirt, is a mixture of organic matter, minerals, gases, liquids, and organisms that together support life. Some scientific definitions distinguish ''dirt'' from ''soil'' by restricting the former ...
, a combination of mineral particles and decomposed
organic matter Organic matter, organic material, or natural organic matter refers to the large source of carbon-based compounds found within natural and engineered, terrestrial, and aquatic environments. It is matter composed of organic compounds that have c ...
. Land surfaces were either bare rock or shifting sand produced by
weathering Weathering is the deterioration of rocks, soils and minerals as well as wood and artificial materials through contact with water, atmospheric gases, and biological organisms. Weathering occurs '' in situ'' (on site, with little or no movement ...
. Water and dissolved nutrients would have drained away very quickly. In the
Sub-Cambrian peneplain The sub-Cambrian peneplain is an ancient, extremely flat, erosion surface ( peneplain) that has been exhumed and exposed by erosion from under Cambrian strata over large swathes of Fennoscandia. Eastward, where this peneplain dips below Cambria ...
in Sweden, for example, maximum depth of
kaolin Kaolinite ( ) is a clay mineral, with the chemical composition Al2 Si2 O5( OH)4. It is an important industrial mineral. It is a layered silicate mineral, with one tetrahedral sheet of silica () linked through oxygen atoms to one octahedra ...
itization by Neoproterozoic
weathering Weathering is the deterioration of rocks, soils and minerals as well as wood and artificial materials through contact with water, atmospheric gases, and biological organisms. Weathering occurs '' in situ'' (on site, with little or no movement ...
is about 5 m, while nearby kaolin deposits developed in the
Mesozoic The Mesozoic Era ( ), also called the Age of Reptiles, the Age of Conifers, and colloquially as the Age of the Dinosaurs is the second-to-last era of Earth's geological history, lasting from about , comprising the Triassic, Jurassic and Cretace ...
are much thicker. It has been argued that in the late Neoproterozoic
sheet wash Sheet erosion or sheet wash is the even erosion of substrate along a wide area. It occurs in a wide range of settings such as coastal plains, hillslopes, floodplains, beaches, savanna plains and semi-arid plains. Water moving fairly uniformly with ...
was a dominant process of erosion of surface material due to the lack of plants on land. Films of cyanobacteria, which are not plants but use the same photosynthesis mechanisms, have been found in modern deserts in areas unsuitable for
vascular plant Vascular plants (), also called tracheophytes () or collectively Tracheophyta (), form a large group of land plants ( accepted known species) that have lignified tissues (the xylem) for conducting water and minerals throughout the plant. They ...
s. This suggests that microbial mats may have been the first organisms to colonize dry land, possibly in the Precambrian. Mat-forming cyanobacteria could have gradually evolved resistance to desiccation as they spread from the seas to
intertidal zone The intertidal zone, also known as the foreshore, is the area above water level at low tide and underwater at high tide (in other words, the area within the tidal range). This area can include several types of habitats with various species ...
s and then to land. Shear 2000, "The Early Development of Terrestrial Ecosystems," pp
169–184
/ref>
Lichen A lichen ( , ) is a composite organism that arises from algae or cyanobacteria living among filaments of multiple fungi species in a mutualistic relationship.symbiotic Symbiosis (from Greek , , "living together", from , , "together", and , bíōsis, "living") is any type of a close and long-term biological interaction between two different biological organisms, be it mutualistic, commensalistic, or para ...
combinations of a fungus (almost always an
ascomycete Ascomycota is a phylum of the kingdom Fungi that, together with the Basidiomycota, forms the subkingdom Dikarya. Its members are commonly known as the sac fungi or ascomycetes. It is the largest phylum of Fungi, with over 64,000 species. The defi ...
) and one or more photosynthesizers (green algae or cyanobacteria), are also important colonizers of lifeless environments, and their ability to break down rocks contributes to soil formation where plants cannot survive. The earliest known ascomycete fossils date from in the
Silurian The Silurian ( ) is a geologic period and system spanning 24.6 million years from the end of the Ordovician Period, at million years ago ( Mya), to the beginning of the Devonian Period, Mya. The Silurian is the shortest period of the Paleoz ...
. Soil formation would have been very slow until the appearance of burrowing animals, which mix the mineral and organic components of soil and whose
feces Feces ( or faeces), known colloquially and in slang as poo and poop, are the solid or semi-solid remains of food that was not digested in the small intestine, and has been broken down by bacteria in the large intestine. Feces contain a rela ...
are a major source of organic components. Burrows have been found in Ordovician sediments, and are attributed to
annelid The annelids (Annelida , from Latin ', "little ring"), also known as the segmented worms, are a large phylum, with over 22,000 extant species including ragworms, earthworms, and leeches. The species exist in and have adapted to various ecol ...
s (worms) or arthropods.


Plants and the Late Devonian wood crisis

In aquatic algae, almost all cells are capable of photosynthesis and are nearly independent. Life on land required plants to become internally more complex and specialized: photosynthesis is most efficient at the top; roots extract water and nutrients from the ground; and the intermediate parts support and transport. Spores of land plants resembling
liverworts The Marchantiophyta () are a division of non-vascular land plants commonly referred to as hepatics or liverworts. Like mosses and hornworts, they have a gametophyte-dominant life cycle, in which cells of the plant carry only a single set of ...
have been found in Middle Ordovician rocks from . Middle Silurian rocks from contain fossils of true plants, including clubmosses such as ''
Baragwanathia ''Baragwanathia'' is a genus of extinct lycopsid plants of Late Silurian to Early Devonian age (), fossils of which have been found in Australia, Canada, China and Czechia. The name derives from William Baragwanath who discovered the first specim ...
''; most were under high, and some appear closely related to
vascular plant Vascular plants (), also called tracheophytes () or collectively Tracheophyta (), form a large group of land plants ( accepted known species) that have lignified tissues (the xylem) for conducting water and minerals throughout the plant. They ...
s, the group that includes
tree In botany, a tree is a perennial plant with an elongated stem, or trunk, usually supporting branches and leaves. In some usages, the definition of a tree may be narrower, including only woody plants with secondary growth, plants that are ...
s. By the Late Devonian , abundant trees such as ''
Archaeopteris ''Archaeopteris'' is an extinct genus of progymnosperm tree with fern-like leaves. A useful index fossil, this tree is found in strata dating from the Upper Devonian to Lower Carboniferous (), the oldest fossils being 385 million years old, ...
'' bound the soil so firmly that they changed river systems from mostly
braided Braided is a musical group consisting of Casey LeBlanc, Ashley Leitão, and Amber Fleury, who all competed on the third season of ''Canadian Idol'' in 2005. They are the third music group to come from an Idol show in the world, after Young Divas ...
to mostly
meander A meander is one of a series of regular sinuous curves in the channel of a river or other watercourse. It is produced as a watercourse erodes the sediments of an outer, concave bank ( cut bank) and deposits sediments on an inner, convex ban ...
ing. This caused the "Late Devonian wood crisis" because: * They removed more carbon dioxide from the atmosphere, reducing the
greenhouse effect The greenhouse effect is a process that occurs when energy from a planet's host star goes through the planet's atmosphere and heats the planet's surface, but greenhouse gases in the atmosphere prevent some of the heat from returning directly ...
and thus causing an
ice age An ice age is a long period of reduction in the temperature of Earth's surface and atmosphere, resulting in the presence or expansion of continental and polar ice sheets and alpine glaciers. Earth's climate alternates between ice ages and gre ...
in the
Carboniferous The Carboniferous ( ) is a geologic period and system of the Paleozoic that spans 60 million years from the end of the Devonian Period million years ago ( Mya), to the beginning of the Permian Period, million years ago. The name ''Carboniferou ...
period. This did not repeat in later ecosystems, since the carbon dioxide "locked up" in wood was returned to the atmosphere by decomposition of dead wood, but the earliest fossil evidence of fungi that can decompose wood also comes from the Late Devonian. * The increasing depth of plants' roots led to more washing of nutrients into rivers and seas by rain. This caused algal blooms whose high consumption of oxygen caused anoxic events in deeper waters, increasing the extinction rate among deep-water animals.


Land invertebrates (tetrapods)

Animals had to change their feeding and excretory systems, and most land animals developed internal fertilization of their eggs. The difference in
refractive index In optics, the refractive index (or refraction index) of an optical medium is a dimensionless number that gives the indication of the light bending ability of that medium. The refractive index determines how much the path of light is bent, ...
between water and air required changes in their eyes. On the other hand, in some ways movement and breathing became easier, and the better transmission of high-frequency sounds in air encouraged the development of hearing. The oldest known air-breathing animal is '' Pneumodesmus'', an
archipolypoda Archipolypoda is an extinct group of millipedes known from fossils in Europe and North America and containing the earliest known land animals. The Archipolypoda was erected by Scudder (1882) but redefined in 2005 with the description of sever ...
n millipede from the Middle Silurian, about . Its air-breathing, terrestrial nature is evidenced by the presence of spiracles, the openings to tracheal systems. However, some earlier trace fossils from the Cambrian-Ordovician boundary about are interpreted as the tracks of large
amphibious Amphibious means able to use either land or water. In particular it may refer to: Animals * Amphibian, a vertebrate animal of the class Amphibia (many of which live on land and breed in water) * Amphibious caterpillar * Amphibious fish, a fish ...
arthropods on coastal
sand dune A dune is a landform composed of wind- or water-driven sand. It typically takes the form of a mound, ridge, or hill. An area with dunes is called a dune system or a dune complex. A large dune complex is called a dune field, while broad, f ...
s, and may have been made by
euthycarcinoid Euthycarcinoidea are an enigmatic group of extinct possibly amphibious arthropods that ranged from Cambrian to Triassic times. Fossils are known from Europe, North America, Argentina, Australia and Antarctica. Description The euthycarcinoid body ...
s, which are thought to be evolutionary "aunts" of myriapods. Other trace fossils from the Late Ordovician a little over probably represent land invertebrates, and there is clear evidence of numerous arthropods on coasts and
alluvial plain An alluvial plain is a largely flat landform created by the deposition of sediment over a long period of time by one or more rivers coming from highland regions, from which alluvial soil forms. A floodplain is part of the process, being the smal ...
s shortly before the Silurian-Devonian boundary, about , including signs that some arthropods ate plants. Arthropods were well
pre-adapted Exaptation and the related term co-option describe a shift in the function of a trait during evolution. For example, a trait can evolve because it served one particular function, but subsequently it may come to serve another. Exaptations are common ...
to colonise land, because their existing jointed exoskeletons provided protection against desiccation, support against gravity and a means of locomotion that was not dependent on water. The fossil record of other major invertebrate groups on land is poor: none at all for non-
parasitic Parasitism is a close relationship between species, where one organism, the parasite, lives on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The entomologist E. O. Wilson ha ...
flatworm The flatworms, flat worms, Platyhelminthes, or platyhelminths (from the Greek πλατύ, ''platy'', meaning "flat" and ἕλμινς (root: ἑλμινθ-), ''helminth-'', meaning "worm") are a phylum of relatively simple bilaterian, unsegmen ...
s,
nematode The nematodes ( or grc-gre, Νηματώδη; la, Nematoda) or roundworms constitute the phylum Nematoda (also called Nemathelminthes), with plant- parasitic nematodes also known as eelworms. They are a diverse animal phylum inhabiting a bro ...
s or nemerteans; some parasitic nematodes have been fossilized in
amber Amber is fossilized tree resin that has been appreciated for its color and natural beauty since Neolithic times. Much valued from antiquity to the present as a gemstone, amber is made into a variety of decorative objects."Amber" (2004). In M ...
; annelid worm fossils are known from the Carboniferous, but they may still have been aquatic animals; the earliest fossils of
gastropod The gastropods (), commonly known as snails and slugs, belong to a large taxonomic class of invertebrates within the phylum Mollusca called Gastropoda (). This class comprises snails and slugs from saltwater, from freshwater, and from land. T ...
s on land date from the Late Carboniferous, and this group may have had to wait until
leaf litter Plant litter (also leaf litter, tree litter, soil litter, litterfall or duff) is dead plant material (such as leaves, bark, needles, twigs, and cladodes) that have fallen to the ground. This detritus or dead organic material and its constituent ...
became abundant enough to provide the moist conditions they need. Selden 2001, "Terrestrialization of Animals," pp. 71–74 The earliest confirmed fossils of flying
insect Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body ( head, thorax and abdomen), three pa ...
s date from the Late Carboniferous, but it is thought that insects developed the ability to fly in the Early Carboniferous or even Late Devonian. This gave them a wider range of
ecological niche In ecology, a niche is the match of a species to a specific environmental condition. Three variants of ecological niche are described by It describes how an organism or population responds to the distribution of resources and competitors (for ...
s for feeding and breeding, and a means of escape from predators and from unfavorable changes in the environment. About 99% of modern insect species fly or are descendants of flying species.


Amphibians (Early land vertebrates)

Family tree of
tetrapod Tetrapods (; ) are four-limbed vertebrate animals constituting the superclass Tetrapoda (). It includes extant and extinct amphibians, sauropsids ( reptiles, including dinosaurs and therefore birds) and synapsids ( pelycosaurs, extinct t ...
s
Tetrapod Tetrapods (; ) are four-limbed vertebrate animals constituting the superclass Tetrapoda (). It includes extant and extinct amphibians, sauropsids ( reptiles, including dinosaurs and therefore birds) and synapsids ( pelycosaurs, extinct t ...
s, vertebrates with four limbs, evolved from other rhipidistian fish over a relatively short timespan during the Late Devonian (). The early groups are grouped together as
Labyrinthodontia "Labyrinthodontia" ( Greek, 'maze-toothed') is an informal grouping of extinct predatory amphibians which were major components of ecosystems in the late Paleozoic and early Mesozoic eras (about 390 to 150 million years ago). Traditionally con ...
. They retained aquatic, fry-like
tadpole A tadpole is the larval stage in the biological life cycle of an amphibian. Most tadpoles are fully aquatic, though some species of amphibians have tadpoles that are terrestrial. Tadpoles have some fish-like features that may not be found ...
s, a system still seen in modern amphibians. Iodine and T4/T3 stimulate the amphibian metamorphosis and the
evolution of nervous systems The evolution of nervous systems dates back to the first development of nervous systems in animals (or metazoans). Neurons developed as specialized electrical signaling cells in multicellular animals, adapting the mechanism of action potentials ...
transforming the aquatic, vegetarian tadpole into a "more evolved" terrestrial, carnivorous frog with better neurological, visuospatial, olfactory and cognitive abilities for hunting. The new hormonal action of T3 was made possible by the formation of T3-receptors in the cells of vertebrates. Firstly, about 600-500 million years ago, in primitive Chordata appeared the alpha T3-receptors with a metamorphosing action and then, about 250-150 million years ago, in the Birds and Mammalia appeared the beta T3-receptors with metabolic and thermogenetic actions. From the 1950s to the early 1980s it was thought that tetrapods evolved from fish that had already acquired the ability to crawl on land, possibly in order to go from a pool that was drying out to one that was deeper. However, in 1987, nearly complete fossils of '' Acanthostega'' from about showed that this Late Devonian transitional animal had legs and both
lung The lungs are the primary organs of the respiratory system in humans and most other animals, including some snails and a small number of fish. In mammals and most other vertebrates, two lungs are located near the backbone on either side of ...
s and
gill A gill () is a respiratory organ that many aquatic organisms use to extract dissolved oxygen from water and to excrete carbon dioxide. The gills of some species, such as hermit crabs, have adapted to allow respiration on land provided they ar ...
s, but could never have survived on land: its limbs and its wrist and ankle joints were too weak to bear its weight; its ribs were too short to prevent its lungs from being squeezed flat by its weight; its fish-like tail fin would have been damaged by dragging on the ground. The current hypothesis is that ''Acanthostega'', which was about long, was a wholly aquatic predator that hunted in shallow water. Its skeleton differed from that of most fish, in ways that enabled it to raise its head to breathe air while its body remained submerged, including: its jaws show modifications that would have enabled it to gulp air; the bones at the back of its skull are locked together, providing strong attachment points for muscles that raised its head; the head is not joined to the
shoulder girdle The shoulder girdle or pectoral girdle is the set of bones in the appendicular skeleton which connects to the arm on each side. In humans it consists of the clavicle and scapula; in those species with three bones in the shoulder, it consists ...
and it has a distinct neck. The Devonian proliferation of land plants may help to explain why air breathing would have been an advantage: leaves falling into streams and rivers would have encouraged the growth of aquatic vegetation; this would have attracted grazing invertebrates and small fish that preyed on them; they would have been attractive prey but the environment was unsuitable for the big marine predatory fish; air-breathing would have been necessary because these waters would have been short of oxygen, since warm water holds less dissolved oxygen than cooler marine water and since the decomposition of vegetation would have used some of the oxygen. Later discoveries revealed earlier transitional forms between ''Acanthostega'' and completely fish-like animals. Unfortunately, there is then a gap (
Romer's gap Romer's gap is an example of an apparent gap in the tetrapod fossil record used in the study of evolutionary biology. Such gaps represent periods from which excavators have not yet found relevant fossils. Romer's gap is named after paleontologist ...
) of about 30 Ma between the fossils of ancestral tetrapods and Middle Carboniferous fossils of vertebrates that look well-adapted for life on land, during which only some fossils are found, which had five digits in the terminating point of the four limbs, showing true or crown
tetrapods Tetrapods (; ) are four-limbed vertebrate animals constituting the superclass Tetrapoda (). It includes extant and extinct amphibians, sauropsids (reptiles, including dinosaurs and therefore birds) and synapsids ( pelycosaurs, extinct therapsi ...
appeared in the gap around 350 Ma. Some of the fossils after this gap look as if the animals which they belonged to were early relatives of modern amphibians, all of which need to keep their skins moist and to lay their eggs in water, while others are accepted as early relatives of the
amniote Amniotes are a clade of tetrapod vertebrates that comprises sauropsids (including all reptiles and birds, and extinct parareptiles and non-avian dinosaurs) and synapsids (including pelycosaurs and therapsids such as mammals). They are dis ...
s, whose waterproof skin and egg membranes enable them to live and breed far from water. The Carboniferous Rainforest Collapse may have paved the way for amniotes to become dominant over amphibians.


Reptiles (inlcuding dinosaurs)

Possible family tree of
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the evolution of dinosaurs is t ...
s,
bird Birds are a group of warm-blooded vertebrates constituting the class Aves (), characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, and a strong yet lightweig ...
s and
mammal Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fur ...
s
Amniotes, whose eggs can survive in dry environments, probably evolved in the Late Carboniferous period (). The earliest fossils of the two surviving amniote groups, synapsids and
sauropsid Sauropsida ("lizard faces") is a clade of amniotes, broadly equivalent to the class Reptilia. Sauropsida is the sister taxon to Synapsida, the other clade of amniotes which includes mammals as its only modern representatives. Although early syn ...
s, date from around . The synapsid pelycosaurs and their descendants the
therapsid Therapsida is a major group of eupelycosaurian synapsids that includes mammals, their ancestors and relatives. Many of the traits today seen as unique to mammals had their origin within early therapsids, including limbs that were oriented more ...
s are the most common land vertebrates in the best-known Permian () fossil beds. However, at the time these were all in
temperate In geography, the temperate climates of Earth occur in the middle latitudes (23.5° to 66.5° N/S of Equator), which span between the tropics and the polar regions of Earth. These zones generally have wider temperature ranges throughout ...
zones at middle
latitude In geography, latitude is a coordinate that specifies the north– south position of a point on the surface of the Earth or another celestial body. Latitude is given as an angle that ranges from –90° at the south pole to 90° at the north ...
s, and there is evidence that hotter, drier environments nearer the Equator were dominated by sauropsids and amphibians. The
Permian–Triassic extinction event The Permian–Triassic (P–T, P–Tr) extinction event, also known as the Latest Permian extinction event, the End-Permian Extinction and colloquially as the Great Dying, formed the boundary between the Permian and Triassic geologic periods, ...
wiped out almost all land vertebrates, as well as the great majority of other life. During the slow recovery from this catastrophe, estimated to have taken 30 million years, a previously obscure sauropsid group became the most abundant and diverse terrestrial vertebrates: a few fossils of archosauriformes ("ruling lizard forms") have been found in Late Permian rocks, but, by the
Middle Triassic In the geologic timescale, the Middle Triassic is the second of three epochs of the Triassic period or the middle of three series in which the Triassic system is divided in chronostratigraphy. The Middle Triassic spans the time between Ma an ...
, archosaurs were the dominant land vertebrates. Dinosaurs distinguished themselves from other archosaurs in the Late Triassic, and became the dominant land vertebrates of the Jurassic and Cretaceous periods ().


Birds

During the Late Jurassic, birds evolved from small, predatory theropod dinosaurs. The first birds inherited teeth and long, bony tails from their dinosaur ancestors, but some had developed horny, toothless
beak The beak, bill, or rostrum is an external anatomical structure found mostly in birds, but also in turtles, non-avian dinosaurs and a few mammals. A beak is used for eating, preening, manipulating objects, killing prey, fighting, probing for fo ...
s by the very Late Jurassic and short pygostyle tails by the
Early Cretaceous The Early Cretaceous (geochronology, geochronological name) or the Lower Cretaceous (chronostratigraphy, chronostratigraphic name), is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 145& ...
.


Mammals

While the archosaurs and dinosaurs were becoming more dominant in the Triassic, the mammaliaform successors of the therapsids evolved into small, mainly nocturnal
insectivore A robber fly eating a hoverfly An insectivore is a carnivorous animal or plant that eats insects. An alternative term is entomophage, which can also refer to the human practice of eating insects. The first vertebrate insectivores were ...
s. This ecological role may have promoted the
evolution of mammals The evolution of mammals has passed through many stages since the first appearance of their synapsid ancestors in the Pennsylvanian sub-period of the late Carboniferous period. By the mid-Triassic, there were many synapsid species that looked l ...
, for example nocturnal life may have accelerated the development of
endotherm An endotherm (from Greek ἔνδον ''endon'' "within" and θέρμη ''thermē'' "heat") is an organism that maintains its body at a metabolically favorable temperature, largely by the use of heat released by its internal bodily functions inst ...
y ("warm-bloodedness") and hair or fur. By in the
Early Jurassic The Early Jurassic Epoch (geology), Epoch (in chronostratigraphy corresponding to the Lower Jurassic series (stratigraphy), Series) is the earliest of three epochs of the Jurassic Period. The Early Jurassic starts immediately after the Triassic-J ...
there were animals that were very like today's mammals in a number of respects. Unfortunately, there is a gap in the fossil record throughout the Middle Jurassic. However, fossil teeth discovered in
Madagascar Madagascar (; mg, Madagasikara, ), officially the Republic of Madagascar ( mg, Repoblikan'i Madagasikara, links=no, ; french: République de Madagascar), is an island country in the Indian Ocean, approximately off the coast of East Afric ...
indicate that the split between the lineage leading to
monotreme Monotremes () are prototherian mammals of the order Monotremata. They are one of the three groups of living mammals, along with placentals ( Eutheria), and marsupials (Metatheria). Monotremes are typified by structural differences in their brai ...
s and the one leading to other living mammals had occurred by . After dominating land vertebrate niches for about 150 Ma, the non-avian dinosaurs perished in the Cretaceous–Paleogene extinction event () along with many other groups of organisms. Mammals throughout the time of the dinosaurs had been restricted to a narrow range of taxa, sizes and shapes, but increased rapidly in size and diversity after the extinction, with bats taking to the air within 13 million years, and
cetacea Cetacea (; , ) is an infraorder of aquatic mammals that includes whales, dolphins, and porpoises. Key characteristics are their fully aquatic lifestyle, streamlined body shape, often large size and exclusively carnivorous diet. They propel th ...
ns to the sea within 15 million years.


Flowering plants

The first flowering plants appeared around 130 Ma. The 250,000 to 400,000 species of flowering plants outnumber all other ground plants combined, and are the dominant vegetation in most terrestrial ecosystems. There is fossil evidence that flowering plants diversified rapidly in the Early Cretaceous, from , Crane, Friis and Pedersen 2000, "The Origin and Early Diversification of Angiosperms," pp
233–250
/ref> and that their rise was associated with that of
pollinating Pollination is the transfer of pollen from an anther of a plant to the stigma of a plant, later enabling fertilisation and the production of seeds, most often by an animal or by wind. Pollinating agents can be animals such as insects, birds, a ...
insects. Among modern flowering plants ''
Magnolia ''Magnolia'' is a large genus of about 210 to 340The number of species in the genus ''Magnolia'' depends on the taxonomic view that one takes up. Recent molecular and morphological research shows that former genera ''Talauma'', ''Dugandiodendr ...
'' are thought to be close to the common ancestor of the group. However, paleontologists have not succeeded in identifying the earliest stages in the evolution of flowering plants.


Social insects

The social insects are remarkable because the great majority of individuals in each colony are sterile. This appears contrary to basic concepts of evolution such as natural selection and the selfish gene. In fact, there are very few eusocial insect species: only 15 out of approximately 2,600 living families of insects contain eusocial species, and it seems that eusociality has evolved independently only 12 times among arthropods, although some eusocial lineages have diversified into several families. Nevertheless, social insects have been spectacularly successful; for example although ants and
termite Termites are small insects that live in colonies and have distinct castes ( eusocial) and feed on wood or other dead plant matter. Termites comprise the infraorder Isoptera, or alternatively the epifamily Termitoidae, within the order Blat ...
s account for only about 2% of known insect species, they form over 50% of the total mass of insects. Their ability to control a territory appears to be the foundation of their success. The sacrifice of breeding opportunities by most individuals has long been explained as a consequence of these species' unusual haplodiploid method of sex determination, which has the paradoxical consequence that two sterile worker daughters of the same queen share more genes with each other than they would with their offspring if they could breed. However, E. O. Wilson and Bert Hölldobler argue that this explanation is faulty: for example, it is based on
kin selection Kin selection is the evolutionary strategy that favours the reproductive success of an organism's relatives, even when at a cost to the organism's own survival and reproduction. Kin altruism can look like altruistic behaviour whose evolution ...
, but there is no evidence of
nepotism Nepotism is an advantage, privilege, or position that is granted to relatives and friends in an occupation or field. These fields may include but are not limited to, business, politics, academia, entertainment, sports, fitness, religion, an ...
in colonies that have multiple queens. Instead, they write, eusociality evolves only in species that are under strong pressure from predators and competitors, but in environments where it is possible to build "fortresses"; after colonies have established this security, they gain other advantages through co-operative
foraging Foraging is searching for wild food resources. It affects an animal's fitness because it plays an important role in an animal's ability to survive and reproduce. Foraging theory is a branch of behavioral ecology that studies the foraging behavi ...
. In support of this explanation they cite the appearance of eusociality in bathyergid mole rats, which are not haplodiploid. The earliest fossils of insects have been found in Early Devonian rocks from about , which preserve only a few varieties of flightless insect. The Mazon Creek lagerstätten from the Late Carboniferous, about , include about 200 species, some gigantic by modern standards, and indicate that insects had occupied their main modern ecological niches as
herbivore A herbivore is an animal anatomically and physiologically adapted to eating plant material, for example foliage or marine algae, for the main component of its diet. As a result of their plant diet, herbivorous animals typically have mouthpar ...
s,
detritivore Detritivores (also known as detrivores, detritophages, detritus feeders, or detritus eaters) are heterotrophs that obtain nutrients by consuming detritus (decomposing plant and animal parts as well as feces). There are many kinds of invertebrate ...
s and insectivores. Social termites and ants first appear in the Early Cretaceous, and advanced social bees have been found in Late Cretaceous rocks but did not become abundant until the Middle
Cenozoic The Cenozoic ( ; ) is Earth's current geological era, representing the last 66million years of Earth's history. It is characterised by the dominance of mammals, birds and flowering plants, a cooling and drying climate, and the current configu ...
.


Humans

The idea that, along with other life forms, modern-day humans evolved from an ancient, common ancestor was proposed by Robert Chambers in 1844 and taken up by
Charles Darwin Charles Robert Darwin ( ; 12 February 1809 – 19 April 1882) was an English naturalist, geologist, and biologist, widely known for his contributions to evolutionary biology. His proposition that all species of life have descended ...
in 1871. Modern humans evolved from a lineage of upright-walking apes that has been traced back over to '' Sahelanthropus''. The first known
stone tool A stone tool is, in the most general sense, any tool made either partially or entirely out of stone. Although stone tool-dependent societies and cultures still exist today, most stone tools are associated with prehistoric (particularly Stone A ...
s were made about , apparently by '' Australopithecus garhi'', and were found near animal bones that bear scratches made by these tools. The earliest
hominine Homininae (), also called "African hominids" or "African apes", is a subfamily of Hominidae. It includes two tribes, with their extant as well as extinct species: 1) the tribe Hominini (with the genus ''Homo'' including modern humans and numerou ...
s had
chimpanzee The chimpanzee (''Pan troglodytes''), also known as simply the chimp, is a species of great ape native to the forest and savannah of tropical Africa. It has four confirmed subspecies and a fifth proposed subspecies. When its close relative t ...
-sized brains, but there has been a fourfold increase in the last 3 Ma; a statistical analysis suggests that hominine brain sizes depend almost completely on the date of the fossils, while the species to which they are assigned has only slight influence. There is a long-running debate about whether modern humans evolved all over the world simultaneously from existing advanced hominines or are descendants of a single small population in Africa, which then migrated all over the world less than 200,000 years ago and replaced previous hominine species. There is also debate about whether anatomically modern humans had an intellectual, cultural and technological "Great Leap Forward" under 100,000 years ago and, if so, whether this was due to neurological changes that are not visible in fossils.


Mass extinctions

Life on Earth has suffered occasional mass extinctions at least since . Although they were disasters at the time, mass extinctions have sometimes accelerated the evolution of life on Earth. When dominance of particular ecological niches passes from one group of organisms to another, it is rarely because the new dominant group is "superior" to the old and usually because an extinction event eliminates the old dominant group and makes way for the new one. The fossil record appears to show that the gaps between mass extinctions are becoming longer and the average and background rates of extinction are decreasing. Both of these phenomena could be explained in one or more ways: * The oceans may have become more hospitable to life over the last 500 Ma and less vulnerable to mass extinctions: dissolved oxygen became more widespread and penetrated to greater depths; the development of life on land reduced the run-off of nutrients and hence the risk of
eutrophication Eutrophication is the process by which an entire body of water, or parts of it, becomes progressively enriched with minerals and nutrients, particularly nitrogen and phosphorus. It has also been defined as "nutrient-induced increase in phyt ...
and anoxic events; and marine ecosystems became more diversified so that food chains were less likely to be disrupted. * Reasonably complete fossils are very rare, most extinct organisms are represented only by partial fossils, and complete fossils are rarest in the oldest rocks. So paleontologists have mistakenly assigned parts of the same organism to different genera, which were often defined solely to accommodate these finds—the story of ''
Anomalocaris ''Anomalocaris'' ("unlike other shrimp", or "abnormal shrimp") is an extinct genus of radiodont, an order of early-diverging stem-group arthropods. The first fossils of ''Anomalocaris'' were discovered in the ''Ogygopsis'' Shale of the Stephen F ...
'' is an example of this. The risk of this mistake is higher for older fossils because these are often both unlike parts of any living organism and poorly conserved. Many of the "superfluous" genera are represented by fragments which are not found again and the "superfluous" genera appear to become extinct very quickly.
Biodiversity Biodiversity or biological diversity is the variety and variability of life on Earth. Biodiversity is a measure of variation at the genetic ('' genetic variability''), species ('' species diversity''), and ecosystem ('' ecosystem diversity'') ...
in the fossil record, which is "...the number of distinct genera alive at any given time; that is, those whose first occurrence predates and whose last occurrence postdates that time" shows a different trend: a fairly swift rise from ; a slight decline from , in which the devastating Permian–Triassic extinction event is an important factor; and a swift rise from to the present.


See also

* History of evolutionary thought * ''
On the Origin of Species ''On the Origin of Species'' (or, more completely, ''On the Origin of Species by Means of Natural Selection, or the Preservation of Favoured Races in the Struggle for Life''),The book's full original title was ''On the Origin of Species by Me ...
'' *
Speciation Speciation is the evolutionary process by which populations evolve to become distinct species. The biologist Orator F. Cook coined the term in 1906 for cladogenesis, the splitting of lineages, as opposed to anagenesis, phyletic evolution withi ...
*
Taxonomy of commonly fossilised invertebrates Although the phylogenetic classification of non-vertebrate animals (both extinct and extant) remains a work-in-progress, the following taxonomy attempts to be useful by combining both traditional (old) ''and'' new (21st-century) paleozoological ...
* ''
Treatise on Invertebrate Paleontology The ''Treatise on Invertebrate Paleontology'' (or ''TIP'') published by the Geological Society of America and the University of Kansas Press, is a definitive multi-authored work of some 50 volumes, written by more than 300 paleontologists, and ...
'' * Viral evolution


Footnotes


References


Bibliography

* * * * * * * * * * * * * * * * * * "Proceedings from Nobel Symposium 70 held at
Alfred Nobel Alfred Bernhard Nobel ( , ; 21 October 1833 – 10 December 1896) was a Swedish chemist, engineer, inventor, businessman, and philanthropist. He is best known for having bequeathed his fortune to establish the Nobel Prize, though he al ...
's Björkborn, Karlskoga, Sweden, August 29 – September 2, 1988" * * * * * * * "Preview booklet for 'Gondwana alive : biodiversity and the evolving terrestrial biosphere', book planned for September 2000, and associated projects." * * * * * * * * * * * *


Further reading

* * * *


External links

* * * General information on evolution compiled by Roger Perkins. * * * Tree of life diagram by Neal Olander. * * * Tutorial created by Dennis O'Neil. {{DEFAULTSORT:History Of Life Evolution-related timelines History by topic Paleontology Evolutionary biology Earth sciences Geochronology