History of life
The history of life on Earth begins with a planet that formed about 4.54 billion years ago, give or take 50 million. Scientists abbreviate that span as Ga, for gigaannum, and they suspect life appeared before 3.7 Ga. Every species alive today, from a microbe to a whale, traces back to a common ancestor through evolution. The proof is in the resemblance. All known organisms share the same basic biochemical machinery. That shared inheritance raises hard questions. How did life cross the line from non-living chemistry to a thing that copies itself? Why did oxygen, a waste product, end up reshaping the whole planet? What made complex cells possible, and why does almost everything reproduce sexually when an asexual rival could out-breed it in fifty generations? Consider one striking estimate. Earth may hold a trillion species, yet only 1.75 to 1.8 million have been named. The currently living species represent less than one percent of all species that have ever lived. The rest are gone, and much of what follows is the story of who replaced whom.
Biogenic carbon signatures and stromatolite fossils sit inside 3.7 billion-year-old metasedimentary rocks from western Greenland, the earliest clear evidence of life. Older claims push further back and grow shakier. In 2015, possible remains of biotic life turned up in 4.1 billion-year-old rocks in Western Australia. Fossilized microorganisms in hydrothermal vent precipitates from the Nuvvuagittuq Belt in Quebec, Canada, may have lived as early as 4.28 billion years ago, though the evidence is disputed as inconclusive. The oceans had formed by 4.4 billion years ago, so life, if real, arrived soon after there was water to hold it. The earliest organisms make detective work brutal. They were minute and relatively featureless, and their fossils look like small rods nearly impossible to separate from structures abiotic physics can produce. The oldest undisputed evidence, read as fossilized bacteria, dates to 3 Ga. Older finds at roughly 3.5 Ga and geochemical hints at 3.8 Ga drew close scrutiny, and researchers found non-biological processes that could mimic every reported signature of life. That does not prove the structures were lifeless. It means they cannot stand as clear proof. Stromatolites add their own quarrel. Critics argue that alleged examples from before 3 Ga could be non-biological, though in 2006 a fresh find emerged from Australia in rocks dated to 3.5 Ga.
Carbon and water are the foundation of life on Earth, and no substitute comes close. Carbon builds stable frameworks for complex chemicals and pulls easily from carbon dioxide. Silicon sits directly below carbon on the periodic table, yet it forms few complex stable molecules, and silicon dioxide is a hard abrasive solid rather than a convenient gas. Water adds two quiet gifts. Ice floats, so aquatic organisms survive beneath it in winter, and water's electrically charged ends let it form a wider range of compounds than rivals like ammonia. Researchers chasing the leap from chemistry to biology focus on three starting points. There is self-replication, the ability to make near-copies of oneself. There is metabolism, the ability to feed and repair. And there are external cell membranes that admit food, expel waste, and bar unwanted substances. From these threads come competing origin stories. The replication-first view imagines an RNA world, because some RNA molecules can catalyze both their own replication and the building of proteins. Such an RNA world would have had individuals but no species, with mutations and horizontal gene transfer making offspring genetically unlike their parents. Evidence suggests the first RNA molecules formed prior to 4.17 Ga. Rival accounts put membranes first, with lipid bubbles called liposomes that reproduce themselves, or hand the early work to clays. Montmorillonite clay grows by self-replicating its crystal pattern and can catalyze the formation of RNA molecules. A metabolism-first camp points to iron-sulfur and nickel-sulfide catalysts that, in experiments from 1997, began building proteins from carbon monoxide and hydrogen sulfide near hydrothermal vents.
Wet-dry cycles at geothermal springs solve a stubborn problem, hydrolysis, and they push biopolymers to polymerize and pack into vesicles. Analysis of hydrothermal veins at a 3.5 Gya geothermal spring setting found the elements an origin needs, including potassium, boron, hydrogen, sulfur, phosphorus, zinc, nitrogen, and oxygen. Mulkidjanian and colleagues note that such freshwater settings carry ionic concentrations identical to the cytoplasm of modern cells. Deep sea hydrothermal vents offer a competing cradle. In simulated white-smoker conditions, scientists oligomerized RNA four units long, and in alkaline vent pores RNA molecules of 22 bases can be polymerized. A genomic analysis found 355 genes likely tracing to LUCA, the last universal common ancestor, among 6.1 million sequenced prokaryotic genes, and reconstructed LUCA as a thermophilic anaerobe with a Wood-Ljungdahl pathway. Carbonate-rich lakes answer a different shortage, phosphate, which natural environments deplete through microbial uptake and binding to calcium. In carbonate water, calcium locks into calcium carbonate instead of apatite, freeing phosphate to rise. Models put early lake phosphate at roughly 100 times today's levels. Some accounts skip Earth entirely. The panspermia hypothesis, the idea that life was seeded from elsewhere, dates back at least to the Greek philosopher Anaximander in the sixth century BCE, and was later proposed by Svante Arrhenius, by Fred Hoyle and Chandra Wickramasinghe, and by Francis Crick and Leslie Orgel. In low Earth orbit experiments such as EXOSTACK, some microorganism spores survived outer space radiation for at least 5.7 years.
Microbial mats are multi-layered, multi-species colonies only a few millimeters thick, yet each holds a range of chemical environments. The by-products of one group of microorganisms feed the next, so each mat runs its own small food chain. These mats of coexisting bacteria and archaea dominated the early Archean, and many major steps in early evolution likely happened inside them. Stromatolites grew from this world as stubby pillars, built when microorganisms migrated upward to escape sediment settling on them. Cyanobacteria invented photosynthesis around 3.5 Ga, and their waste product would remake the planet. Oxygenic photosynthesis by mat bacteria raised biological productivity by a factor between 100 and 1,000, drawing hydrogen from water rather than scarce geological reducing agents. From that point on, life produced more of its own resources than geochemistry did. Free oxygen first saturated every available reductant on Earth's surface, then spilled into the atmosphere, driving the Great Oxygenation Event around 2.4 Ga. Oxygen is toxic to organisms not adapted to it, yet it sharply boosts the efficiency of those that are. Inside the mats, the boundary between oxygen-rich and oxygen-free layers rose at night when photosynthesis stopped and sank again by day. That daily shift pressured organisms in the middle zone to tolerate oxygen and then to use it, possibly through endosymbiosis, where one organism lives inside another to mutual benefit.
Mitochondria fuel the production of ATP, the internal energy supply of all known cells, and most modern eukaryotes cannot live without the oxygen those mitochondria burn. In the 1970s a vigorous debate settled on an answer for where complex cells came from. Eukaryotes arose through a sequence of endosymbiosis between prokaryotes. A predatory microorganism invaded a large prokaryote, probably an archaean, and instead of killing its prey it took up residence and became mitochondria. The earliest evidence of eukaryotes dates from 1.85 Ga, drawn from fossils of the alga Grypania in rocks of that age. Their diversification sped up once aerobic respiration by mitochondria delivered a richer supply of biological energy. Plastids tell the second chapter of this bargain. Around 1.5 Ga, a eukaryote tried to swallow a photosynthesizing cyanobacterium, the victim survived inside, and the pairing became the ancestor of plants. From that single event grew three lineages, chloroplasts in green algae and plants, rhodoplasts in red algae, and cyanelles in glaucophytes. Around 1.6 Ga, eukaryotes that gained photosynthesis gave rise to algae that eventually overtook cyanobacteria as the dominant primary producers. Dating remains contested. Steranes in Australian shales were once read as eukaryotes at 2.7 Ga, but a 2008 analysis concluded the chemicals infiltrated the rocks less than 2.2 Ga and prove nothing. The earliest known fossils of fungi date from 1.43 Ga.
Meiosis and fertilization define sexual reproduction in eukaryotes, giving offspring 50 percent of their genes from each parent through genetic recombination. The puzzle is why it persists, since an asexual population can out-breed and displace an otherwise equal sexual one in as little as 50 generations. Males, after all, do not directly add to the next generation's numbers. Several hypotheses circle the problem without closing it. The Red Queen hypothesis argues that sex defends against parasites by presenting a moving target, yet Kathryn A. Hanley and colleagues found mites significantly more prevalent in sexual geckos than in asexual ones sharing the same habitat. Alexey Kondrashov's deterministic mutation hypothesis assumes harmful mutations combine to do more damage than their sum, so recombination can purge them, but many species average less than one harmful mutation per individual. John A. Birdsell and Christopher Wills reviewed the competing models, and one alternative holds that sex arose to repair DNA damage, with genetic variety as a byproduct. Sexual reproduction also opened the door to complex bodies. Multicellularity evolved independently in sponges and other animals, fungi, plants, brown algae, cyanobacteria, slime molds, and myxobacteria. Sexual reproduction eliminates rogue cells that would otherwise reduce an asexual organism to undifferentiated mush, so it appears to be a prerequisite for complex multicellularity. The red alga Bangiomorpha, dated at 1.2 Ga, is the earliest known organism with truly differentiated, specialized cells, and also the oldest known sexually reproducing organism.
Bilateria, animals with mirror-image left and right sides, appeared by 555 Ma, and most modern phyla originated during the Cambrian explosion. In November 2019 researchers reported Caveasphaera from 609-million-year-old rocks, a multicellular organism not easily called animal or non-animal, hinting animal evolution may have begun about 750 million years ago. The Ediacara biota flourished for the last 40 million years before the Cambrian, the first animals more than a very few centimeters long. The Chengjiang fossils Haikouichthys and Myllokunmingia appear to be true vertebrates, and Haikouichthys had distinct vertebrae. Moving onto land demanded near-total redesign. Organisms had to avoid drying out, withstand gravity, rework respiration, and free reproduction from water. Modern land ecosystems only appeared in the Late Devonian. Trees such as Archaeopteris bound the soil so firmly that rivers shifted from braided to meandering, triggering the Late Devonian wood crisis. By drawing down carbon dioxide they reduced the greenhouse effect and helped cause a Carboniferous ice age, while their deepening roots washed nutrients into water and fed anoxic algal blooms. Acanthostega, about 1 meter long, was a Late Devonian animal with legs, lungs, and gills, yet too weak to survive on land, a wholly aquatic predator of shallow water. The Permian-Triassic extinction event wiped out almost all land vertebrates, and recovery took an estimated 30 million years. From it rose the archosaurs, and one group, the dinosaurs, ruled the Jurassic and Cretaceous. After the Cretaceous-Paleogene extinction killed the non-avian dinosaurs, mammals increased rapidly in size and diversity, with bats taking to the air within 13 million years and cetaceans to the sea within 15 million years. Modern humans evolved from upright-walking apes traced back to Sahelanthropus, and the first known stone tools were made apparently by Australopithecus garhi. Mass extinctions rarely promote a superior group. They simply clear the old dominant one and make room, and the fossil record suggests the gaps between such catastrophes have been growing longer.
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Common questions
When did life first appear in the history of life on Earth?
Evidence suggests life emerged before 3.7 Ga, the age of biogenic carbon signatures and stromatolite fossils found in metasedimentary rocks from western Greenland. Earth itself formed about 4.54 billion years ago, give or take 50 million, and the oceans had formed by 4.4 billion years ago. Disputed evidence from the Nuvvuagittuq Belt in Quebec, Canada, suggests microorganisms may have lived as early as 4.28 billion years ago.
What was the Great Oxygenation Event in the history of life on Earth?
The Great Oxygenation Event was the buildup of free oxygen in Earth's atmosphere around 2.4 Ga. It happened after cyanobacteria evolved photosynthesis around 3.5 Ga and free oxygen saturated all available reductant substances on Earth's surface. Oxygenic photosynthesis raised biological productivity by a factor between 100 and 1,000.
How did eukaryotic cells evolve in the history of life on Earth?
Eukaryotes arose through a sequence of endosymbiosis between prokaryotes, a conclusion reached in a vigorous 1970s debate. A predatory microorganism invaded a large prokaryote, probably an archaean, and evolved into mitochondria, while a later swallowed cyanobacterium became the ancestor of plants. The earliest evidence of eukaryotes dates from 1.85 Ga.
What are the main hypotheses for the origin of life on Earth?
The leading hypotheses include the RNA world, where replication came first because some RNA molecules catalyze their own replication, a lipid world where membranes formed first, a clay hypothesis centered on montmorillonite, and an iron-sulfur world where metabolism came first. Research focuses on three starting points: self-replication, metabolism, and external cell membranes. Evidence suggests the first RNA molecules formed prior to 4.17 Ga.
Why is sexual reproduction a puzzle in the history of life on Earth?
Sexual reproduction is a puzzle because an asexual population can out-breed and displace an otherwise equal sexual population in as little as 50 generations. Competing explanations include the Red Queen hypothesis about parasites and Alexey Kondrashov's deterministic mutation hypothesis, but each faces contrary evidence. The red alga Bangiomorpha, dated at 1.2 Ga, is the oldest known sexually reproducing organism.
How did mass extinctions shape the history of life on Earth?
Mass extinctions accelerated evolution by eliminating dominant groups and making way for new ones, rather than because the newcomers were superior. The Permian-Triassic extinction event wiped out almost all land vertebrates, and during a recovery estimated at 30 million years the archosaurs and then dinosaurs rose. After the Cretaceous-Paleogene extinction killed the non-avian dinosaurs, mammals increased rapidly in size and diversity.
All sources
272 references cited across the entry
- 1JournalConstraining the Time Interval for the Origin of Life on EarthBen K. D. Pearce et al. — March 1, 2018
- 2Journal13C-Depleted Carbon Microparticles in >3700-Ma Sea-Floor Sedimentary Rocks from West GreenlandMinik T. Rosing — January 29, 1999
- 3JournalEvidence for biogenic graphite in early Archaean Isua metasedimentary rocksYoko Ohtomo et al. — January 2014
- 4Futuyma (2005)Futuyma — 2005
- 5JournalRapid emergence of life shown by discovery of 3,700-million-year-old microbial structuresAllen P. Nutman et al. — September 22, 2016
- 6NewsHints of life on what was thought to be desolate early EarthSeth Borenstein — October 19, 2015
- 7JournalPotentially biogenic carbon preserved in a 4.1 billion-year-old zirconElizabeth A. Bell et al. — November 24, 2015
- 8NewsScientists Say Canadian Bacteria Fossils May Be Earth's OldestCarl Zimmer — March 1, 2017
- 9JournalYoung poorly crystalline graphite in the >3.8-Gyr-old Nuvvuagittuq banded iron formationDominic Papineau et al. — June 2011
- 10JournalA light carbon reservoir recorded in zircon-hosted diamond from the Jack HillsAlexander A. Nemchin et al. — July 3, 2008
- 11JournalArchaean metabolic evolution of microbial matsEuan G. Nisbet et al. — December 7, 1999
- 12JournalA Whiff of Oxygen Before the Great Oxidation Event?Ariel D. Anbar et al. — September 28, 2007
- 13JournalEukaryotic organisms in Proterozoic oceansAndrew H. Knoll et al. — June 29, 2006
- 14JournalResolving the paradox of sex and recombinationSarah P. Otto et al. — April 1, 2002
- 15iTOL: Interactive Tree of LifeIvica Letunic et al. — European Molecular Biology Laboratory
- 16JournalEarth's earliest non-marine eukaryotesPaul K. Strother et al. — May 26, 2011
- 17JournalEarly life on land and the first terrestrial ecosystemsHugo Beraldi-Campesi — February 23, 2013
- 18JournalNew data on Kimberella, the Vendian mollusc-like organism (White Sea region, Russia): palaeoecological and evolutionary implicationsMikhail A. Fedonkin et al. — January 1, 2007
- 19JournalPrecambrian animal diversity: Putative phosphatized embryos from the Doushantuo Formation of ChinaChen Jun-Yuan et al. — April 25, 2000
- 20JournalLower Cambrian vertebrates from south ChinaShu D-G. et al. — November 4, 1999
- 21Synapsid ReptilesDonald F. Hoyt — California State Polytechnic University, Pomona — February 17, 1997
- 22The Great DyingPatrick L. Barry — Marshall Space Flight Center — January 28, 2002
- 23JournalAssessing the record and causes of Late Triassic extinctionsLawrence H. Tanner et al. — March 2004
- 24Benton (1997)Benton — 1997
- 25JournalThe Extinction of the Dinosaurs in North AmericaDavid E. Fastovsky et al. — March 2005
- 26NewsDinosaur Extinction Spurred Rise of Modern MammalsJohn Roach — National Geographic Society — June 20, 2007
- 27JournalMajor Patterns in the History of Carnivorous MammalsBlaire Van Valkenburgh — May 1, 1999
- 28Press releaseResearchers find that Earth may be home to 1 trillion species: Largest analysis of microbial data reveals that 99.999 percent of all species remain undiscoveredCheryl Dybas et al. — National Science Foundation — May 2, 2016
- 29JournalScaling laws predict global microbial diversityKenneth J. Locey et al. — May 24, 2016
- 30NewsPrehistory's Brilliant FutureMichael J. Novacek — November 8, 2014
- 31Catalogue of Life: 2019 Annual ChecklistSpecies 2000; Integrated Taxonomic Information System — 2019
- 32JournalEarly metazoan life: divergence, environment and ecologyDouglas H. Erwin — December 19, 2015
- 33JournalLarge colonial organisms with coordinated growth in oxygenated environments 2.1 Gyr agoAbderrazak El Albani et al. — July 1, 2010
- 35JournalBangiomorpha pubescensn. gen., n. sp.: implications for the evolution of sex, multicellularity, and the Mesoproterozoic/Neoproterozoic radiation of eukaryotesNicholas J. Butterfield — September 2000
- 36Dalrymple (1991)Dalrymple — 1991
- 37BookOrigins of the Earth, Moon, and Life: An Interdisciplinary ApproachAkio Makishima — Elsevier — 2017-01-27
- 38JournalWhere Did the Moon Come From?Edward Belbruno et al. — March 2005
- 39JournalHafnium–tungsten chronometry and the timing of terrestrial core formationDer-Chuen Lee et al. — December 1995
- 40JournalOrigin of the Earth—Moon systemErik M. Galimov et al. — December 2005
- 41NewsOldest Rocks on Earth FoundAndrea Thompson — Imaginova — September 25, 2008
- 42JournalSupport for the Lunar Cataclysm Hypothesis from Lunar Meteorite Impact Melt AgesBarbara A. Cohen et al. — December 1, 2000
- 43Press releaseEarly Earth Likely Had Continents And Was HabitableUniversity of Colorado — November 17, 2005
- 44JournalMagmatic δ18O in 4400–3900 Ma detrital zircons: A record of the alteration and recycling of crust in the Early ArcheanAaron J. Cavosie et al. — July 15, 2005
- 45JournalPatterns In Palaeontology: The first 3 billion years of evolutionRussell J. Garwood — 2012
- 46NewsEvidence for Ancient Bombardment of EarthRobert Roy Britt — Imaginova — July 24, 2002
- 47JournalA cool early EarthJohn W. Valley et al. — April 1, 2002
- 48JournalThe Late Asteroidal and Cometary Bombardment of Earth as Recorded in Water Deuterium to Protium RatioNicolas Dauphas et al. — December 2000
- 49Microbial Habitability During the Late Heavy BombardmentDaniella Scalice — NASA Astrobiology Program — May 20, 2009
- 50JournalMicrofossils of the Early Archean Apex Chert: New Evidence of the Antiquity of LifeJ. William Schopf — April 30, 1993
- 51JournalEvidence for life on Earth before 3,800 million years agoStephen J. Mojzsis et al. — November 7, 1996
- 52JournalAn abiotic model for stromatolite morphogenesisJohn P. Grotzinger et al. — October 3, 1996
- 53JournalMetasomatic Origin of Quartz-Pyroxene Rock, Akilia, Greenland, and Implications for Earth's Earliest LifeChristopher M. Fedo et al. — May 24, 2002
- 54JournalA fresh look at the fossil evidence for early Archaean cellular lifeMartin Brasier et al. — June 2006
- 55JournalFossil evidence of Archaean lifeJ. William Schopf — June 29, 2006
- 56JournalEvidence for early life in Earth's oldest hydrothermal vent precipitatesMatthew S. Dodd et al. — March 2, 2017
- 57NewsThis May Be the Oldest Known Sign of Life on EarthNadia Drake — National Geographic Society — March 1, 2017
- 58JournalToward Automatic Reconstruction of a Highly Resolved Tree of LifeFrancesca D. Ciccarelli et al. — March 3, 2006
- 59JournalOrigins of biomolecular handednessStephen F. Mason — September 6, 1984
- 60MagazineThe Origin of Life on the EarthLeslie E. Orgel — October 1994
- 61BookEarly Life on Earth. Nobel Symposium 84Otto Kandler — Columbia University Press — 1994
- 62JournalCell Wall Biochemistry in Archaea and its Phylogenetic ImplicationsOtto Kandler — 1995
- 63BookThermophiles: The keys to molecular evolution and the origin of life?Otto Kandler — Taylor and Francis Ltd. — 1998
- 64JournalTowards a natural system of organisms: proposal for the domains Archaea, Bacteria, and EucaryaCarl R. Woese et al. — 1990
- 65Bennett (2008) p. 82–85Bennett — 2008
- 66JournalThe prospect of alien life in exotic forms on other worldsDirk Schulze-Makuch et al. — April 2006
- 67JournalControversies on the origin of lifeJuli Peretó — March 2005
- 68JournalIn search of the simplest cellEörs Szathmáry — February 3, 2005
- 69JournalApproaches to semi-synthetic minimal cells: a reviewPier Luigi Luisi et al. — January 2006
- 70JournalThe antiquity of RNA-based evolutionGerald F. Joyce — July 11, 2002
- 71JournalEvolution without speciation but with selection: LUCA, the Last Universal Common Ancestor in Gilbert's RNA worldHugo Hoenigsberg — December 30, 2003
- 72JournalChance and necessity do not explain the origin of lifeJack T. Trevors et al. — November 2004
- 73JournalThe nature of the last universal ancestor and the root of the tree of life, still open questionsPatrick Forterre et al. — 1992
- 74JournalThe Ribosome Is a RibozymeThomas R. Cech — August 11, 2000
- 75JournalOrigin of the RNA world: The fate of nucleobases in warm little pondsBen K. D. Pearce et al. — October 24, 2017
- 76JournalRNA-Catalyzed RNA Polymerization: Accurate and General RNA-Templated Primer ExtensionWendy K. Johnston et al. — May 18, 2001
- 77JournalThe stability of the RNA bases: Implications for the origin of lifeMatthew Levy et al. — July 7, 1998
- 78JournalA Simpler Nucleic AcidLeslie E. Orgel — November 17, 2000
- 79JournalPeptide nucleic acids rather than RNA may have been the first genetic moleculeKevin E. Nelson et al. — April 11, 2000
- 80JournalOn the origins of cells: a hypothesis for the evolutionary transitions from abiotic geochemistry to chemoautotrophic prokaryotes, and from prokaryotes to nucleated cellsWilliam Martin et al. — January 29, 2003
- 81JournalFrom self-assembly of life to present-day bacteria: a possible role for nanocellsJack T. Trevors et al. — December 2001
- 82JournalThe Lipid WorldDaniel Segré et al. — February 2001
- 83Cairns-Smith (1968) p. 57–66Cairns-Smith — 1968
- 84JournalPrebiotic Synthesis on Minerals: Bridging the Prebiotic and RNA WorldsJames P. Ferris — June 1999
- 85JournalExperimental Models of Primitive Cellular Compartments: Encapsulation, Growth, and DivisionMartin M. Hanczyc et al. — October 24, 2003
- 86JournalPhotosynthesis and the Origin of LifeHyman Hartman — October 1998
- 87JournalLife as We Don't Know ItGünter Wächtershäuser — August 25, 2000
- 88JournalWhere Did Life Begin? Testing Ideas in Prebiotic Analogue ConditionsDavid Deamer — February 10, 2021
- 89JournalThe Hot Spring Hypothesis for an Origin of LifeBruce Damer et al. — 2020-04-01
- 90JournalTerrestrial Hot Spring Systems: IntroductionDavid J. Des Marais et al. — 2019-12-01
- 91JournalOrigin of first cells at terrestrial, anoxic geothermal fieldsArmen Y. Mulkidjanian et al. — 2012-04-03
- 92JournalCommon origins of RNA, protein and lipid precursors in a cyanosulfidic protometabolismBhavesh H. Patel et al. — March 16, 2015
- 93JournalElements for the Origin of Life on Land: A Deep-Time Perspective from the Pilbara Craton of Western AustraliaMartin J. Van Kranendonk et al. — 2021-01-01
- 94JournalAmphiphilic Compounds Assemble into Membranous Vesicles in Hydrothermal Hot Spring Water but Not in SeawaterDaniel Milshteyn et al. — 2018-05-10
- 95JournalA carbonate-rich lake solution to the phosphate problem of the origin of lifeJonathan D. Toner et al. — 2020-01-14
- 97JournalRNA Oligomerization in Laboratory Analogues of Alkaline Hydrothermal Vent SystemsBradley T. Burcar et al. — 2015-07-01
- 98JournalA biophysical basis for the emergence of the genetic code in protocellsStuart A. Harrison et al. — 2022-11-01
- 99JournalPromotion of protocell self-assembly from mixed amphiphiles at the origin of lifeSean F. Jordan et al. — November 4, 2019
- 100Hydrothermal vents and prebiotic chemistry: a reviewMaría Colín-García et al. — 2016
- 101JournalHow did LUCA make a living? Chemiosmosis in the origin of lifeNick Lane et al. — 2010-01-27
- 102JournalA Hydrothermal-Sedimentary Context for the Origin of LifeF. Westall et al. — 2018-03-01
- 103JournalThe Origin of Membrane BioenergeticsNick Lane et al. — 2012-12-21
- 104JournalExtreme accumulation of nucleotides in simulated hydrothermal pore systemsPhilipp Baaske et al. — 2007-05-29
- 105JournalThe physiology and habitat of the last universal common ancestorMadeline C. Weiss et al. — 2016-07-25
- 106JournalThe last universal common ancestor between ancient Earth chemistry and the onset of geneticsMadeline C. Weiss et al. — 2018-08-16
- 107JournalLife as a guide to prebiotic nucleotide synthesisStuart A. Harrison et al. — 2018-12-12
- 108JournalPhosphorus in prebiotic chemistryAlan W. Schwartz — 2006-09-07
- 109JournalStability relations of apatites and calcium carbonatesYaacov Nathan et al. — November 1981
- 110JournalPhysical and chemical factors in the formation of marine apatiteR. A. Gulbrandsen — 1969-06-01
- 111JournalEarth's Earliest AtmospheresK. Zahnle et al. — 2010-06-23
- 112JournalDry/Wet Cycling and the Thermodynamics and Kinetics of Prebiotic Polymer SynthesisDavid Ross et al. — 2016-07-26
- 113O'Leary (2008)O'Leary — 2008
- 114Arrhenius (1980) p. 32Arrhenius — 1980
- 115JournalOn the Nature of Interstellar GrainsFred Hoyle et al. — November 1979
- 116JournalDirected PanspermiaFrancis H. Crick et al. — July 1973
- 117MagazineDid Life Come From Another World?David Warmflash et al. — November 2005
- 118JournalOn the possibility of microbiota transfer from Venus to EarthNalin C. Wickramasinghe et al. — September 2008
- 119Clancy, Brack, Horneck (2005)Clancy, Brack, Horneck — 2005
- 120JournalSpace MicrobiologyGerda Horneck et al. — March 2010
- 121JournalSearch for Past Life on Mars: Possible Relic Biogenic Activity in Martian Meteorite ALH84001David S. McKay et al. — 1996-08-16
- 122NewsClaim of Martian Life Called 'Bogus'Ker Than — Imaginova — August 23, 2007
- 123Krumbein, Brehm, Gerdes (2003) p. 1–28Krumbein, Brehm, Gerdes — 2003
- 124JournalCommunity structure of a microbial mat: The phylogenetic dimensionJ. Bruno Risatti et al. — October 11, 1994
- 125JournalBiodiversity rocksJune 8, 2006
- 126JournalStromatolite reef from the Early Archaean era of AustraliaAbigail C. Allwood et al. — June 8, 2006
- 127JournalToxic oxygen: The radical life-giverDoris Abele — November 7, 2002
- 128Introduction to Aerobic RespirationBecky B. Westerdahl — University of California, Davis — 2007
- 129JournalMolecular evidence for the evolution of photosynthesisRobert E. Blankenship — January 1, 2001
- 130JournalThe role of microbial mats in the production of reduced gases on the early EarthTori M. Hoehler et al. — July 19, 2001
- 131JournalThe Great Oxidation at ~2.4 Ga as a bistability in atmospheric oxygen due to UV shielding by ozoneColin Goldblatt et al. — 2006
- 132JournalThe Last Universal Common Ancestor: emergence, constitution and genetic legacy of an elusive forerunnerNicolas Glansdorff et al. — July 9, 2008
- 133JournalArchean Molecular Fossils and the Early Rise of EukaryotesJochen J. Brocks et al. — August 13, 1999
- 134JournalA molecular timescale of eukaryote evolution and the rise of complex multicellular lifeS. Blair Hedges et al. — January 28, 2004
- 135Margulis (1981)Margulis — 1981
- 136JournalThe origin of eukaryotes: the difference between prokaryotic and eukaryotic cellsTibor Vellai et al. — August 7, 1999
- 137JournalReducing the genome size of organelles favours gene transfer to the nucleusMarc-André Selosse et al. — March 1, 2001
- 138JournalSupertrees Disentangle the Chimerical Origin of Eukaryotic GenomesDavide Pisani et al. — August 2007
- 139JournalMitochondrial EvolutionMichael W. Gray et al. — March 5, 1999
- 140JournalReassessing the first appearance of eukaryotes and cyanobacteriaBirger Rasmussen et al. — October 23, 2008
- 141JournalMegascopic eukaryotic algae from the 2.1-billion-year-old negaunee iron-formation, MichiganHan Tsu-Ming et al. — July 10, 1992
- 142JournalTEM evidence for eukaryotic diversity in mid-Proterozoic oceansEmmanuelle J. Javaux et al. — July 2004
- 143JournalProbable Proterozoic fungiNicholas J. Butterfield — Winter 2005
- 144BookAdvances in Photosynthesis and RespirationRobert R. Wise — Springer — 2006
- 145Holmes, Jobling (1996)Holmes, Jobling — 1996
- 146JournalType IV secretion: intercellular transfer of macromolecules by systems ancestrally related to conjugation machinesPeter J. Christie — April 2001
- 147JournalAdaptive value of sex in microbial pathogensRichard E. Michod et al. — May 2008
- 148JournalEvolutionary Origin of Recombination during MeiosisHarris Bernstein et al. — July 2010
- 149JournalNatural genetic transformation: prevalence, mechanisms and functionOla Johnsborg et al. — December 2007
- 150Bernstein, Bernstein, Michod (2012) p. 1–50Bernstein, Bernstein, Michod — 2012
- 151Maurine Neiman et al.John Wiley & Sons — 2010
- 152JournalA Phylogenomic Inventory of Meiotic Genes: Evidence for Sex in Giardia and an Early Eukaryotic Origin of MeiosisMarilee A. Ramesh et al. — January 26, 2005
- 153JournalWhy have sex? The population genetics of sex and recombinationSarah P. Otto et al. — August 2006
- 154JournalLower Mite Infestations in an Asexual Gecko Compared With Its Sexual AncestorsKathryn A. Hanley et al. — June 1995
- 155JournalPathogens and sex in plantsMatthew A. Parker — September 1994
- 156Birdsell, Wills (2003) p. 27–137Birdsell, Wills — 2003
- 157Bernstein, Hopf, Michod (1987) p. 323–370Bernstein, Hopf, Michod — 1987
- 158JournalSize and complexity among multicellular organismsGraham Bell et al. — March 1997
- 159JournalThe origin of the Metazoa in the light of the Proterozoic fossil recordMikhail A. Fedonkin — March 31, 2003
- 160JournalBuilding a Multicellular OrganismDale Kaiser — December 2001
- 161JournalThe origins of multicellularityJohn Tyler Bonner — January 7, 1998
- 162JournalSucrose Utilization in Budding Yeast as a Model for the Origin of Undifferentiated MulticellularityJohn H. Koschwanez et al. — August 9, 2011
- 163JournalMaze-solving by an amoeboid organismToshiyuki Nakagaki et al. — September 28, 2000
- 164JournalAn Arabidopsis transcriptional regulatory map reveals distinct functional and evolutionary features of novel transcription factorsJin Jinpu et al. — July 2015
- 165JournalFungus-like mycelial fossils in 2.4-billion-year-old vesicular basaltStefan Bengtson et al. — 24 April 2017
- 166MagazineEvidence for earlier multicellular lifeGwyneth Dickey — July 31, 2010
- 167JournalBangiomorpha pubescens n. gen., n. sp.: implications for the evolution of sex, multicellularity, and the Mesoproterozoic/Neoproterozoic radiation of eukaryotesNicholas J. Butterfield — Summer 2000
- 168JournalSilicified Horodyskia and Palaeopascichnus from upper Ediacaran cherts in South China: tentative phylogenetic interpretation and implications for evolutionary stasisLin Dong et al. — January 1, 2008
- 169BookThe Invertebrate Tree of LifeGonzalo Giribet et al. — Princeton University Press — 3 March 2020
- 170JournalBuddenbrockia Is a Cnidarian WormEva Jímenez-Guri et al. — July 6, 2007
- 171Animal Cell StructureMichael W. Davidson — Florida State University — May 26, 2005
- 172Concepts of BiologyStephen G. Saupe — College of Saint Benedict and Saint John's University — January 3, 2004
- 173Hinde (2001) p. 28–57Hinde — 2001
- 174Press releaseResearchers say animal-like embryos preceded animal appearanceXiaozheng Chen — American Association for the Advancement of Science — November 27, 2019
- 176JournalThe origin of animals: Can molecular clocks and the fossil record be reconciled?John A. Cunningham et al. — January 2017
- 177JournalPrecambrian Animal Life: Probable Developmental and Adult Cnidarian Forms from Southwest ChinaChen Jun-Yuan et al. — August 1, 2002
- 178JournalPatterns of distribution in the Ediacaran biotas: facies versus biogeography and evolutionDima Grazhdankin — June 2004
- 179JournalVendobionta and Psammocorallia: lost constructions of Precambrian evolutionAdolf Seilacher — August 1992
- 180JournalAge of Neoproterozoic Bilaterian Body and Trace Fossils, White Sea, Russia: Implications for Metazoan EvolutionMark W. Martin et al. — May 5, 2000
- 181JournalThe late Precambrian fossil Kimberella is a mollusc-like bilaterian organismMikhail A. Fedonkin et al. — August 28, 1997
- 182JournalEvolution Within a Bizarre Phylum: Homologies of the First EchinodermsRich Mooi et al. — December 1998
- 183Spriggina is a trilobitoid ecdysozoanMark A. S. McMenamin — Geological Society of America — September 2003
- 184JournalA Parvancorina-like arthropod from the Cambrian of South ChinaLin Jih-Pai et al. — 2006
- 185JournalHooking some stem-group 'worms': fossil lophotrochozoans in the Burgess ShaleNicholas J. Butterfield — December 2006
- 186Bengtson (2004) p. 67–78Bengtson — 2004
- 187Gould (1989) p. 124–136Gould — 1989
- 188Gould (1989)Gould — 1989
- 189JournalThe Cambrian Fossil Record and the Origin of the PhylaGraham E. Budd — February 2003
- 190JournalThe morphology of Opabinia regalis and the reconstruction of the arthropod stem-groupGraham E. Budd — March 1996
- 191JournalExplaining the Cambrian 'Explosion' of AnimalsCharles R. Marshall — May 30, 2006
- 192Philippe JanvierJohn Wiley & Sons — 2001
- 193MagazineOnce we were wormsSimon Conway Morris — August 2, 2003
- 194JournalThe first tunicate from the Early Cambrian of South ChinaJun-Yuan Chen et al. — July 8, 2003
- 195JournalHead and backbone of the Early Cambrian vertebrate HaikouichthysD.-G. Shu — January 30, 2003
- 196Sansom, Smith, Smith (2001) p. 156–171Sansom, Smith, Smith — 2001
- 197Cowen (2000) p. 120–122Cowen — 2000
- 198JournalEarly Terrestrial Animals, Evolution, and UncertaintyRussell J. Garwood et al. — September 2011
- 199JournalA genomic timescale of prokaryote evolution: insights into the origin of methanogenesis, phototrophy, and the colonization of landFabia U. Battistuzzi et al. — November 9, 2004
- 200Weber, Büdel, Belnap (2016) p. 37–54Weber, Büdel, Belnap — 2016
- 201Press releaseOldest evidence of life on land found in 3.48 billion-year-old Australian rocksDeborah Smith — UNSW Media — May 10, 2017
- 202JournalEarliest signs of life on land preserved in ca. 3.5 Ga hot spring depositsTara Djokic et al. — May 9, 2017
- 203JournalMicrobial life and biogeochemical cycling on land 3,220 million years agoMartin Homann et al. — September 2018
- 204NewsA Billion-Year-Old Fungus May Hold Clues to Life's Arrival on LandCarl Zimmer — May 22, 2019
- 205JournalEarly fungi from the Proterozoic era in Arctic CanadaCorentin C. Loron et al. — June 13, 2019
- 206Billion-year-old fossils may be early fungusJohn Timmer — May 22, 2019
- 207JournalEvolutionary Significance of IodineSebastiano Venturi — September 2011
- 208JournalEvolutionary roots of iodine and thyroid hormones in cell-cell signalingSusan J. Crockford — August 2009
- 209JournalEnvironmental Iodine Deficiency: A Challenge to the Evolution of Terrestrial Life?Sebastiano Venturi et al. — August 2000
- 210JournalIodide accumulation provides kelp with an inorganic antioxidant impacting atmospheric chemistryFrithjof C. Küpper et al. — May 13, 2008
- 211JournalStratigraphic Landscape Analysis and geomorphological paradigms: Scandinavia as an example of Phanerozoic uplift and subsidenceKarna Lidmar-Bergström et al. — January 2013
- 212JournalDenudation surfaces and tectonics in the southernmost part of the Baltic ShieldKarna Lidmar-Bergström — December 1993
- 214David L. HawksworthJohn Wiley & Sons — 2002
- 215JournalTrace Fossil Evidence for Late Ordovician Animals on LandGregory J. Retallack et al. — January 2, 1987
- 216JournalThe origin and early evolution of plants on landPaul Kenrick et al. — September 4, 1997
- 218JournalTerrestrial-marine teleconnections in the Devonian: links between the evolution of land plants, weathering processes, and marine anoxic eventsThomas J. Algeo et al. — January 29, 1998
- 219JournalThe importance of fungi in shaping the paleoecosystemThomas N. Taylor et al. — February 1996
- 220JournalMyriapod divergence times differ between molecular clock and fossil evidence: U/Pb zircon ages of the earliest fossil millipede-bearing sediments and their significanceM. E. Brookfield et al. — 2021-10-03
- 221JournalThe geological record and phylogeny of the MyriapodaWilliam A. Shear et al. — March–May 2010
- 222JournalFirst steps on land: Arthropod trackways in Cambrian-Ordovician eolian sandstone, southeastern Ontario, CanadaRobert B. MacNaughton et al. — May 2002
- 223JournalCambrian origins and affinities of an enigmatic fossil group of arthropodsN. Emilio Vaccari et al. — July 29, 2004
- 224JournalThe Ichnologic Record of the Continental Invertebrate Invasion: Evolutionary Trends in Environmental Expansion, Ecospace Utilization, and Behavioral ComplexityLuis A. Buatois et al. — June 1998
- 225Cowen (2000) p. 126Cowen — 2000
- 226Grimaldi, Engel (2005) p. 155–160Grimaldi, Engel — 2005
- 227Grimaldi, Engel (2005) p. 12Grimaldi, Engel — 2005
- 228JournalIntroduction to the Special Collection: Revisiting the Vertebrate Invasion of the LandMalcolm S. Gordon et al. — September–October 2004
- 230MagazineGetting a Leg Up on LandJennifer A. Clack — December 2005
- 231JournalA Devonian tetrapod-like fish and the evolution of the tetrapod body planEdward B. Daeschler et al. — April 6, 2006
- 232JournalThe origin and early diversification of tetrapodsPer E. Ahlberg et al. — April 7, 1994
- 233JournalReptile phylogeny and the interrelationships of turtlesMichael deBraga et al. — July 1997
- 234JournalPaleontological Evidence to Date the Tree of LifeMichael J. Benton et al. — January 2007
- 235JournalPhylogeny of the major tetrapod groups: Morphological data and divergence datesMichael J. Benton — May 1990
- 236JournalPermian tetrapods from the Sahara show climate-controlled endemism in PangaeaChristian A. Sidor et al. — April 14, 2005
- 237JournalThe recovery of terrestrial vertebrate diversity in the South African Karoo Basin after the end-Permian extinctionRoger Smith et al. — September–October 2005
- 238Benton (2005)Benton — 2005
- 239JournalRecovery from the most profound mass extinction of all timeSarda Sahney et al. — April 7, 2008
- 240Gauthier, Cannatella, de Queiroz (1989) p. 345Gauthier, Cannatella, de Queiroz — 1989
- 241JournalDinosaur Success in the Triassic: A Noncompetitive Ecological ModelMichael J. Benton — March 1983
- 242Padian (2004) p. 210–231Padian — 2004
- 243JournalA beaked bird from the Jurassic of ChinaLian-hai Hou et al. — October 19, 1995
- 244JournalInsight into the evolution of avian flight from a new clade of Early Cretaceous ornithurines from China and the morphology of Yixianornis grabauiJulia A. Clarke et al. — March 2006
- 245JournalSelective Factors Associated with the Origin of Fur and FeathersJohn A. Ruben et al. — August 2000
- 246JournalA New Mammaliaform from the Early Jurassic and Evolution of Mammalian CharacteristicsLuo Zhe-Xi et al. — May 25, 2001
- 247JournalEarly mammalian radiationsRichard L. Cifelli — November 2001
- 248JournalA Middle Jurassic mammal from MadagascarJohn J. Flynn et al. — September 2, 1999
- 249JournalThe Cretaceous–Tertiary biotic transitionNorman MacLeod et al. — April 1997
- 250JournalThe Fossil Record of North American Mammals: Evidence for a Paleocene Evolutionary RadiationJohn Alroy — March 1999
- 251JournalQuantitative Analysis of the Timing of the Origin and Diversification of Extant Placental OrdersJ. David Archibald et al. — June 2001
- 252JournalPrimitive Early Eocene bat from Wyoming and the evolution of flight and echolocationNancy B. Simmons et al. — February 14, 2008
- 253Thewissen, Madar, Hussain (1996)Thewissen, Madar, Hussain — 1996
- 254evolution: plant timelineEncyclopædia Britannica, Inc. — 1996
- 256JournalProgress in understanding angiosperm history, success, and relationships: Darwin's abominably 'perplexing phenomenon'William L. Crepet — November 21, 2000
- 257JournalEusociality: Origin and consequencesEdward O. Wilson et al. — September 20, 2005
- 258JournalAncestral Monogamy Shows Kin Selection Is Key to the Evolution of EusocialityWilliam O. H. Hughes et al. — May 30, 2008
- 259JournalThe evolution of eusociality in molerats (Bathyergidae): a question of risks, numbers, and costsBarry G. Lovegrove — January 1991
- 260Labandeira, Eble (1999)Labandeira, Eble — 1999
- 261Darwin & The Descent of ManStephen Montgomery — Christ's College — 2009
- 262JournalA new hominid from the Upper Miocene of Chad, Central AfricaMichel Brunet et al. — July 11, 2002
- 263JournalEnvironment and Behavior of 2.5-Million-Year-Old Bouri HominidsJean de Heinzelin et al. — April 23, 1999
- 264JournalVariation in hominid brain size: How much is due to method?Carmen De Miguel et al. — 2001
- 265Leakey (1994) p. 87–89Leakey — 1994
- 266JournalAnatomy, behavior, and modern human originsRichard Klein — 1995
- 267JournalWhy did modern human populations disperse from Africa ca. 60,000 years ago? A new modelPaul Mellars — June 20, 2006
- 268Benton, 2005a p. Chapter 6: "Tetrapods of the Triassic"Benton, 2005a
- 269Extinction!Norman MacLeod — 1999
- 270JournalCyclic and secular variation in microfossil biomineralization: clues to the biogeochemical evolution of Phanerozoic oceansRonald E. Martin — June 1995
- 271JournalSecular Increase in Nutrient Levels through the Phanerozoic: Implications for Productivity, Biomass, and Diversity of the Marine BiosphereRonald E. Martin — June 1996
- 272JournalCycles in fossil diversityRobert A. Rohde et al. — March 10, 2005