754:
vertebrate evolution earlier than either birds or mammals, the cytochrome c of both mammals and birds should be equally different from the cytochrome c of fish. Similarly, all vertebrate cytochrome c should be equally different from the yeast protein." For example, the difference between the cytochrome c of a carp and a frog, turtle, chicken, rabbit, and horse is a very constant 13% to 14%. Similarly, the difference between the cytochrome c of a bacterium and yeast, wheat, moth, tuna, pigeon, and horse ranges from 64% to 69%. Together with the work of Emile
Zuckerkandl and Linus Pauling, the genetic equidistance result led directly to the formal postulation of the molecular clock hypothesis in the early 1960s.
1124:
49:
604:
978:) that can be used to express the uncertainty associated with divergence time estimates. Determining the shape and parameters of the probability distribution is not trivial, but there are methods that use not only the oldest fossil but a larger sample of the fossil record of clades to estimate calibration densities empirically. Studies have shown that increasing the number of fossil constraints increases the accuracy of divergence time estimation.
1033:
simultaneously, the risk of biased results is decreased. This approach has been improved upon by pairing it with different models. One current method of molecular clock calibration is total evidence dating paired with the fossilized birth-death (FBD) model and a model of morphological evolution. The FBD model is novel in that it allows for "sampled ancestors", which are fossil taxa that are the direct ancestor of a living taxon or
591:
1020:
calibrating from the known age of a node, expansion calibration uses a two-epoch model of constant population size followed by population growth, with the time of transition between epochs being the parameter of interest for calibration. Expansion calibration works at shorter, intraspecific timescales in comparison to node calibration, because expansions can only be detected after the
794:(New World Monkey) species (within experimental error). This meant that they had both accumulated approximately equal changes in albumin since their shared common ancestor. This pattern was also found for all the primate comparisons they tested. When calibrated with the few well-documented fossil branch points (such as no Primate fossils of modern aspect found before the
774:
differences between an outgroup (more distantly related) species and the faster-evolving species should be larger (since more molecular changes would have accumulated on that lineage) than the molecular differences between the outgroup species and the slower-evolving species. This method is known as the
1006:
dataset for both the extinct and the extant taxa. Unlike node calibration, this method reconstructs the tree topology and places the fossils simultaneously. Molecular and morphological models work together simultaneously, allowing morphology to inform the placement of fossils. Tip calibration makes
1032:
This approach to tip calibration goes a step further by simultaneously estimating fossil placement, topology, and the evolutionary timescale. In this method, the age of a fossil can inform its phylogenetic position in addition to morphology. By allowing all aspects of tree reconstruction to occur
949:
is used to constrain the minimum possible age for the node representing the most recent common ancestor of the clade. However, due to incomplete fossil preservation and other factors, clades are typically older than their oldest fossils. In order to account for this, nodes are allowed to be older
753:
of any two species is mostly conditioned by the time elapsed since the lines of evolution leading to these two species originally diverged. If this is correct, the cytochrome c of all mammals should be equally different from the cytochrome c of all birds. Since fish diverges from the main stem of
773:
proteins showed that approximately constant rates of change had occurred in all the lineages they assessed. The basic logic of their analysis involved recognizing that if one species lineage had evolved more quickly than a sister species lineage since their common ancestor, then the molecular
1019:
from a sample of extant genetic variation in the population using coalescent theory. Ancient population expansions that are well documented and dated in the geological record can be used to calibrate a rate of molecular evolution in a manner similar to node calibration. However, instead of
1187:
with time, but instead flattens out. Even at intermediate genetic distances, with phylogenetic data still sufficient to estimate topology, signal for the overall scale of the tree can be weak under complex likelihood models, leading to highly uncertain molecular clock estimates.
1057:
Sometimes only a single divergence date can be estimated from fossils, with all other dates inferred from that. Other sets of species have abundant fossils available, allowing the hypothesis of constant divergence rates to be tested. DNA sequences experiencing low levels of
950:
than the minimum constraint in node calibration analyses. However, determining how much older the node is allowed to be is challenging. There are a number of strategies for deriving the maximum bound for the age of a clade including those based on birth-death models, fossil
919:
studies—two areas of evolutionary biology where it is possible to sample sequences over an evolutionary timescale—the dates of the intermediate samples can be used to calibrate the molecular clock. However, most phylogenies require that the molecular clock be
1167:
techniques that explore a weighted range of tree topologies and simultaneously estimate parameters of the chosen substitution model. It must be remembered that divergence dates inferred using a molecular clock are based on statistical
1070:
In addition to such variation in rate with genomic position, since the early 1990s variation among taxa has proven fertile ground for research too, even over comparatively short periods of evolutionary time (for example
3006:
Marshall, D. C., et al. 2016. Inflation of molecular clock rates and dates: molecular phylogenetics, biogeography, and diversification of a global cicada radiation from
Australasia (Hemiptera: Cicadidae: Cicadettini).
1007:
use of all relevant fossil taxa during clock calibration, rather than relying on only the oldest fossil of each clade. This method does not rely on the interpretation of negative evidence to infer maximum clade ages.
914:
To use molecular clocks to estimate divergence times, molecular clocks need to be "calibrated". This is because molecular data alone does not contain any information on absolute times. For viral phylogenetics and
940:
by specifying time constraints for one or more nodes in the tree. Early methods of clock calibration only used a single fossil constraint (e.g. non-parametric rate smoothing), but newer methods (BEAST and
1066:
in bacteria, mammals, invertebrates, and plants. In the same study, genomic regions experiencing very high negative or purifying selection (encoding rRNA) were considerably slower (1% per 50 Myr).
1024:
of the species in question. Expansion dating has been used to show that molecular clock rates can be inflated at short timescales (< 1 MY) due to incomplete fixation of alleles, as discussed below
2551:
Douzery EJ, Delsuc F, Stanhope MJ, Huchon D (2003). "Local molecular clocks in three nuclear genes: divergence times for rodents and other mammals and incompatibility among fossil calibrations".
1183:. When enough time has passed, many sites have undergone more than one change, but it is impossible to detect more than one. This means that the observed number of changes is no longer
1155:. In particular, models that take into account rate variation across lineages have been proposed in order to obtain better estimates of divergence times. These models are called
1091:
have more fundamentally challenged the molecular clock hypothesis. According to Ayala's 1999 study, five factors combine to limit the application of molecular clock models:
1235:
has allowed the study of macroevolutionary processes in organisms that had limited fossil records. Phylogenetic comparative methods rely heavily on calibrated phylogenies.
2757:
892:
872:
848:
320:
962:
can be used to represent the uncertainty about the age of the clade. These calibration densities can take the shape of standard probability densities (e.g.
854:
in the population is then 1/N, since each copy of the gene is as good as any other. Every generation, each individual can have new mutations, so there are
3209:
2665:"Major analytical and conceptual shortcomings in a recent taxonomic revision of the Procellariiformes – A reply to Penhallurick and Wink (2004)"
635:
1095:
Changing generation times (If the rate of new mutations depends at least partly on the number of generations rather than the number of years)
3794:
3470:
2717:"Mitochondrial DNA evolution at a turtle's pace: evidence for low genetic variability and reduced microevolutionary rate in the Testudines"
243:
2140:"Expansion dating: calibrating molecular clocks in marine species from expansions onto the Sunda Shelf Following the Last Glacial Maximum"
3008:
554:
2773:
1083:
times, and many turtles have a molecular clock running at one-eighth the speed it does in small mammals, or even slower. Effects of
1045:
Bayesian methods can provide more appropriate estimates of divergence times, especially if large datasets—such as those yielded by
362:
4086:
3214:
1842:
1199:
that were both present as part of a polymorphism in the common ancestor. The inclusion of differences that have not yet become
1179:
The molecular clock runs into particular challenges at very short and very long timescales. At long timescales, the problem is
1059:
928:
record. There are two general methods for calibrating the molecular clock using fossils: node calibration and tip calibration.
68:
2785:
3090:
1278:
818:, which predicted a molecular clock. Let there be N individuals, and to keep this calculation simple, let the individuals be
815:
549:
3241:
496:
1254:
3588:
628:
357:
188:
2860:
Pascual-GarcĂa A, Arenas M, Bastolla U (November 2019). "The
Molecular Clock in the Evolution of Protein Structures".
798:), this led Sarich and Wilson to argue that the human-chimp divergence probably occurred only ~4–6 million years ago.
3789:
3385:
2500:
Ochman H, Wilson AC (1987). "Evolution in bacteria: evidence for a universal substitution rate in cellular genomes".
2453:"Phylogenomic datasets provide both precision and accuracy in estimating the timescale of placental mammal phylogeny"
795:
24:
2956:
Felsenstein J (2001). "Taking variation of evolutionary rates between sites into account in inferring phylogenies".
1947:
Zheng Y, Wiens JJ (April 2015). "Do missing data influence the accuracy of divergence-time estimation with BEAST?".
3425:
1220:
90:
2799:
Schwartz, J. H. & Maresca, B. (2006). "Do
Molecular Clocks Run at All? A Critique of Molecular Systematics".
1037:. This allows fossils to be placed on a branch above an extant organism, rather than being confined to the tips.
347:
315:
2632:
2613:
1203:
leads to a potentially dramatic inflation of the apparent rate of the molecular clock at very short timescales.
3204:
1797:"r8s: inferring absolute rates of molecular evolution and divergence times in the absence of a molecular clock"
1200:
1192:
899:
851:
569:
352:
3835:
3583:
1147:
Molecular clock users have developed workaround solutions using a number of statistical approaches including
621:
608:
2614:"Molecular systematics and biogeography of Antillean thrashers, tremblers, and mockingbirds (Aves: Mimidae)"
4091:
3578:
726:
with time, as estimated from fossil evidence. They generalized this observation to assert that the rate of
539:
4096:
3768:
3638:
1021:
595:
3978:
3958:
3145:
Morgan GJ (1998). "Emile
Zuckerkandl, Linus Pauling, and the molecular evolutionary clock, 1959-1965".
807:
95:
2284:"Time dependency of molecular rate estimates and systematic overestimation of recent divergence times"
3968:
3940:
3773:
3552:
2664:
1867:
1016:
461:
436:
416:
396:
73:
2813:
1159:
because they represent an intermediate position between the 'strict' molecular clock hypothesis and
4035:
3234:
2573:
1164:
1105:
Species-specific differences (due to differing metabolism, ecology, evolutionary history, ...)
971:
762:
451:
446:
421:
376:
342:
336:
325:
1108:
Change in function of the protein studied (can be avoided in closely related species by utilizing
1079:
have molecular clocks that on average run at half speed of many other birds, possibly due to long
3973:
3923:
3710:
3557:
2838:
1268:
995:
967:
564:
471:
466:
411:
372:
173:
2031:
Shapiro B, Drummond AJ, Rambaut A, Wilson MC, Matheus PE, Sher AV, et al. (November 2004).
1320:
4101:
3873:
3674:
3420:
3319:
2808:
2568:
476:
431:
253:
148:
945:) allow for the use of multiple fossils to calibrate molecular clocks. The oldest fossil of a
3475:
3189:(1965). "Evolutionary divergence and convergence in proteins". In Bryson V, Vogel HJ (eds.).
1212:
1135:) have long generation times and lower mutation rates, as expressed by short branches in the
1084:
990:, tip calibration is a method of molecular clock calibration in which fossils are treated as
574:
456:
401:
367:
280:
1015:
Demographic changes in populations can be detected as fluctuations in historical coalescent
3908:
3849:
3825:
3820:
2965:
2681:
2560:
2509:
2346:
2044:
1903:
1563:
1504:
1449:
1347:
1180:
1003:
942:
661:
426:
198:
83:
40:
2733:
2716:
2325:"The fossilized birth-death process for coherent calibration of divergence-time estimates"
2008:
1991:
1720:
1704:"A nonparametric approach to estimating divergence times in the absence of rate constancy"
1703:
994:
and placed on the tips of the tree. This is achieved by creating a matrix that includes a
877:
857:
833:
8:
3935:
3830:
3748:
3738:
3513:
3430:
3410:
3400:
3227:
1273:
1195:
of different sequences in the different populations. Instead, they represent alternative
1152:
1034:
963:
959:
895:
790:) albumin immunological cross-reactions suggested they were about equally different from
516:
506:
441:
406:
290:
193:
118:
53:
2969:
2685:
2564:
2513:
2350:
2048:
1907:
1659:
Philosophical
Transactions of the Royal Society of London. Series B, Biological Sciences
1567:
1508:
1495:
Sarich VM, Wilson AC (December 1967). "Immunological time scale for hominid evolution".
1453:
1351:
4060:
4040:
3918:
3903:
3888:
3810:
3609:
3573:
3375:
3170:
3133:
3096:
3047:
3022:
2989:
2933:
2906:
2826:
2697:
2645:
2594:
2533:
2477:
2452:
2428:
2405:
2393:
2369:
2336:
2324:
2256:
2231:
2181:"Late Glacial Demographic Expansion Motivates a Clock Overhaul for Population Genetics"
2112:
2087:
2068:
1972:
1929:
1772:
1747:
1725:
1679:
1654:
1587:
1536:
1520:
1418:
1311:
1263:
1160:
1148:
1088:
975:
874:
N new neutral mutations in the population as a whole. That means that each generation,
775:
746:
672:
559:
491:
275:
203:
168:
1813:
1796:
1472:
1437:
1370:
1335:
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3664:
3659:
3503:
3339:
3182:
3162:
3125:
3100:
3086:
3052:
2981:
2938:
2887:
2777:
2738:
2637:
2586:
2525:
2482:
2433:
2392:
Gavryushkina A, Heath TA, Ksepka DT, Stadler T, Welch D, Drummond AJ (January 2017).
2374:
2305:
2261:
2212:
2161:
2117:
2060:
2013:
1964:
1933:
1921:
1872:
1818:
1777:
1684:
1628:
1579:
1528:
1477:
1410:
1375:
1302:
1259:
1136:
1076:
937:
902:
in a population will accumulate at a clock-rate that is equal to the rate of neutral
827:
786:
707:
706:
The notion of the existence of a so-called "molecular clock" was first attributed to
260:
138:
128:
123:
3137:
2830:
2701:
2649:
2598:
2072:
1729:
1422:
936:
Sometimes referred to as node dating, node calibration is a method for time-scaling
4065:
4014:
3930:
3878:
3654:
3632:
3498:
3395:
3174:
3154:
3117:
3078:
3042:
3034:
2993:
2973:
2928:
2918:
2877:
2869:
2818:
2769:
2728:
2689:
2627:
2578:
2537:
2517:
2472:
2464:
2423:
2415:
2364:
2354:
2295:
2251:
2243:
2202:
2192:
2151:
2107:
2099:
2052:
2003:
1992:"Population growth makes waves in the distribution of pairwise genetic differences"
1976:
1956:
1911:
1892:"CladeDate : Calibration information generator for divergence time estimation"
1862:
1854:
1808:
1767:
1759:
1715:
1674:
1666:
1618:
1591:
1571:
1540:
1512:
1467:
1457:
1402:
1365:
1355:
1283:
791:
295:
1516:
3963:
3950:
3913:
3860:
3718:
3613:
3547:
3508:
3345:
2923:
2451:
dos Reis M, Inoue J, Hasegawa M, Asher RJ, Donoghue PC, Yang Z (September 2012).
1184:
1113:
723:
248:
238:
100:
1960:
1102:
is stronger in small populations, and so more mutations are effectively neutral)
3898:
3696:
3684:
3628:
3623:
3617:
3542:
3465:
3390:
2882:
2394:"Bayesian Total-Evidence Dating Reveals the Recent Crown Radiation of Penguins"
2329:
Proceedings of the
National Academy of Sciences of the United States of America
1442:
Proceedings of the
National Academy of Sciences of the United States of America
1340:
Proceedings of the
National Academy of Sciences of the United States of America
1244:
1216:
1109:
806:
The observation of a clock-like rate of molecular change was originally purely
758:
734:
was approximately constant over time and over different lineages (known as the
330:
233:
20:
16:
Technique to deduce the time in prehistory when two or more life forms diverged
3158:
3082:
2822:
2582:
2088:"Inference of human population history from individual whole-genome sequences"
1858:
1123:
48:
4080:
3815:
3733:
3728:
3679:
3669:
3309:
3186:
2672:
2641:
2618:
2138:
Crandall ED, Sbrocco EJ, Deboer TS, Barber PH, Carpenter KE (February 2012).
1925:
1306:
1191:
At very short time scales, many differences between samples do not represent
1128:
1099:
1046:
711:
653:
481:
153:
3038:
2873:
2419:
2359:
2300:
2283:
2197:
2180:
2156:
2139:
2056:
1916:
1891:
1763:
1623:
1606:
3893:
3691:
3405:
3380:
3365:
3331:
3291:
3166:
3129:
3056:
2985:
2942:
2891:
2843:
2781:
2590:
2486:
2468:
2437:
2378:
2309:
2265:
2247:
2216:
2165:
2121:
2064:
1968:
1876:
1822:
1781:
1688:
1670:
1632:
1414:
1379:
951:
811:
780:
750:
679:
501:
486:
270:
265:
183:
3023:"Quantitative prediction of molecular clock and ka/ks at short timescales"
2977:
2742:
2529:
2017:
1583:
1532:
1481:
1462:
1360:
1087:
are also likely to confound molecular clock analyses. Researchers such as
3593:
3490:
3452:
3435:
3370:
3351:
3284:
3264:
1249:
1072:
921:
916:
657:
511:
228:
178:
2774:
10.1002/(SICI)1521-1878(199901)21:1<71::AID-BIES9>3.0.CO;2-B
2230:
Ho SY, Tong KJ, Foster CS, Ritchie AM, Lo N, Crisp MD (September 2015).
2103:
749:, who wrote: "It appears that the number of residue differences between
3360:
3355:
3250:
2521:
1524:
1080:
999:
987:
719:
715:
691:
676:
665:
285:
208:
163:
143:
57:
3108:
Kumar S (August 2005). "Molecular clocks: four decades of evolution".
2839:"No Missing Link? Evolutionary Changes Occur Suddenly, Professor Says"
2207:
1554:
Kimura M (February 1968). "Evolutionary rate at the molecular level".
1393:
Kumar S (August 2005). "Molecular clocks: four decades of evolution".
4050:
4030:
3763:
3723:
3460:
3269:
1575:
1224:
1169:
1132:
955:
894:
new neutral mutations will become fixed. If most changes seen during
727:
669:
544:
158:
78:
3210:
Molecular clock explanation of the molecular equidistance phenomenon
3121:
2693:
1406:
4045:
3883:
3527:
3440:
2410:
1655:"The evolution of methods for establishing evolutionary timescales"
1173:
903:
823:
534:
133:
2341:
2032:
4055:
3279:
3274:
1228:
1196:
1140:
770:
766:
731:
695:
2715:
Avise JC, Bowen BW, Lamb T, Meylan AB, Bermingham E (May 1992).
958:
controls. Alternatively, instead of a maximum and a minimum, a
925:
819:
778:. Sarich and Wilson's paper reported, for example, that human (
675:. The biomolecular data used for such calculations are usually
590:
2391:
3480:
3219:
1232:
991:
946:
765:
in 1967 demonstrated that molecular differences among modern
2859:
2030:
1840:
3414:
3296:
2633:
10.1642/0004-8038(2001)118[0035:MSABOA]2.0.CO;2
2550:
2137:
822:(i.e. have one copy of each gene). Let the rate of neutral
701:
2798:
2450:
3314:
2904:
1746:
Drummond AJ, Suchard MA, Xie D, Rambaut A (August 2012).
1745:
1063:
687:
683:
2662:
2281:
2905:
Drummond AJ, Ho SY, Phillips MJ, Rambaut A (May 2006).
2322:
2282:
Ho SY, Phillips MJ, Cooper A, Drummond AJ (July 2005).
1312:"Molecular disease, evolution, and genic heterogeneity"
1052:
2611:
1841:
O'Reilly JE, Dos Reis M, Donoghue PC (November 2015).
1748:"Bayesian phylogenetics with BEAUti and the BEAST 1.7"
1211:
The molecular clock technique is an important tool in
3075:
The
Molecular Evolutionary Clock: Theory and Practice
880:
860:
850:. The probability that this new mutation will become
836:
2714:
2232:"Biogeographic calibrations for the molecular clock"
924:
using independent evidence about dates, such as the
3795:
Global Boundary Stratotype Section and Point (GSSP)
3215:
Date-a-Clade service for the molecular tree of life
2229:
1607:"Paleontological evidence to date the tree of life"
801:
652:is a figurative term for a technique that uses the
3181:
2907:"Relaxed phylogenetics and dating with confidence"
1310:
1301:
1163:'s many-rates model and are made possible through
886:
866:
842:
2323:Heath TA, Huelsenbeck JP, Stadler T (July 2014).
1836:
1834:
1832:
1336:"Primary Structure and Evolution of Cytochrome C"
4078:
1989:
2385:
1829:
1604:
1119:Changes in the intensity of natural selection.
3235:
3020:
2949:
2033:"Rise and fall of the Beringian steppe bison"
1883:
1741:
1739:
1139:, than the fast-evolving herbaceous bamboos (
1062:showed divergence rates of 0.7–0.8% per
629:
3193:. Academic Press, New York. pp. 97–166.
2499:
2444:
1843:"Dating Tips for Divergence-Time Estimation"
1652:
1648:
1646:
1644:
1642:
1494:
1435:
2955:
2853:
2612:Hunt JS, Bermingham E, Ricklefs RE (2001).
1295:
722:between different lineages changes roughly
3242:
3228:
3072:
2898:
2792:
1946:
1736:
1333:
1227:events, including those not documented by
636:
622:
3046:
2932:
2922:
2881:
2812:
2732:
2656:
2631:
2572:
2544:
2476:
2427:
2409:
2368:
2358:
2340:
2299:
2255:
2206:
2196:
2155:
2111:
2007:
1915:
1889:
1868:1983/ba7bbcf4-1d51-4b74-a800-9948edb3bbe6
1866:
1812:
1794:
1771:
1719:
1701:
1678:
1639:
1622:
1471:
1461:
1369:
1359:
1231:, such as the divergences between living
714:who, in 1962, noticed that the number of
3790:Global Standard Stratigraphic Age (GSSA)
2663:Rheindt, F. E. & Austin, J. (2005).
2493:
2085:
1438:"Rates of albumin evolution in primates"
1122:
1027:
1010:
702:Early discovery and genetic equidistance
2749:
2708:
2316:
2178:
1940:
1605:Benton MJ, Donoghue PC (January 2007).
1598:
1547:
4079:
3144:
3021:Peterson GI, Masel J (November 2009).
2605:
1553:
745:phenomenon was first noted in 1963 by
3223:
3107:
2755:
2734:10.1093/oxfordjournals.molbev.a040735
2277:
2275:
2133:
2131:
2009:10.1093/oxfordjournals.molbev.a040727
1949:Molecular Phylogenetics and Evolution
1721:10.1093/oxfordjournals.molbev.a025731
1392:
1319:. Academic Press, New York. pp.
1279:Neutral theory of molecular evolution
816:neutral theory of molecular evolution
3205:Allan Wilson and the molecular clock
1990:Rogers AR, Harpending H (May 1992).
1053:Non-constant rate of molecular clock
1315:. In Kasha, M., Pullman, B (eds.).
1255:Human mitochondrial molecular clock
931:
13:
3589:Adoption of the Gregorian calendar
3066:
2847:(Press release). 12 February 2007.
2272:
2128:
1436:Sarich VM, Wilson AC (July 1967).
981:
826:(i.e. mutations with no effect on
14:
4113:
3198:
3147:Journal of the History of Biology
1653:Donoghue PC, Yang Z (July 2016).
2457:Proceedings. Biological Sciences
1896:Methods in Ecology and Evolution
1221:phylogenetic comparative methods
802:Relationship with neutral theory
603:
602:
589:
47:
3471:English and British regnal year
3027:Molecular Biology and Evolution
3014:
3000:
2721:Molecular Biology and Evolution
2288:Molecular Biology and Evolution
2223:
2172:
2144:Molecular Biology and Evolution
2079:
2024:
1996:Molecular Biology and Evolution
1983:
1814:10.1093/bioinformatics/19.2.301
1788:
1752:Molecular Biology and Evolution
1708:Molecular Biology and Evolution
1611:Molecular Biology and Evolution
596:Evolutionary biology portal
3249:
3009:Systematic Biology 65(1):16–34
2958:Journal of Molecular Evolution
2553:Journal of Molecular Evolution
2502:Journal of Molecular Evolution
1695:
1488:
1429:
1386:
1327:
909:
555:Creation–evolution controversy
309:History of evolutionary theory
1:
4087:Evolutionary biology concepts
3584:Old Style and New Style dates
1795:Sanderson MJ (January 2003).
1517:10.1126/science.158.3805.1200
1334:Margoliash E (October 1963).
1289:
1223:. Estimation of the dates of
3536:Pre-Julian / Julian
2924:10.1371/journal.pbio.0040088
2086:Li H, Durbin R (July 2011).
540:Evolution as fact and theory
7:
3769:Geological history of Earth
3639:Astronomical year numbering
3191:Evolving Genes and Proteins
1961:10.1016/j.ympev.2015.02.002
1238:
1022:most recent common ancestor
10:
4118:
1040:
954:distribution analyses, or
736:molecular clock hypothesis
575:Nature-nurture controversy
18:
4023:
4007:
3991:
3949:
3941:Thermoluminescence dating
3859:
3848:
3836:Samarium–neodymium dating
3803:
3782:
3756:
3747:
3709:
3647:
3602:
3566:
3535:
3526:
3489:
3451:
3330:
3305:
3257:
3083:10.1007/978-3-030-60181-2
2823:10.1162/biot.2006.1.4.357
2758:"Molecular clock mirages"
2756:Ayala FJ (January 1999).
2583:10.1007/s00239-003-0028-x
1859:10.1016/j.tig.2015.08.001
1112:sequences or emphasizing
1017:effective population size
986:Sometimes referred to as
830:) in a new individual be
462:Evolutionary neuroscience
437:Evolutionary epistemology
417:Evolutionary anthropology
397:Applications of evolution
3655:Chinese sexagenary cycle
3110:Nature Reviews. Genetics
3073:Ho, S.Y.W., ed. (2020).
1902:(11). Wiley: 2331–2338.
1395:Nature Reviews. Genetics
1317:Horizons in Biochemistry
1157:relaxed molecular clocks
730:change of any specified
452:Evolutionary linguistics
447:Evolutionary game theory
422:Evolutionary computation
19:Not to be confused with
3869:Amino acid racemisation
3159:10.1023/A:1004394418084
2360:10.1073/pnas.1319091111
2179:Hoareau TB (May 2016).
2057:10.1126/science.1101074
1917:10.1111/2041-210x.13977
1269:Models of DNA evolution
1206:
565:Objections to evolution
472:Evolutionary psychology
467:Evolutionary physiology
412:Evolutionary aesthetics
391:Fields and applications
373:History of paleontology
3874:Archaeomagnetic dating
3386:Era of Caesar (Iberia)
2559:(Suppl 1): S201–S213.
2469:10.1098/rspb.2012.0683
2248:10.1098/rsbl.2015.0194
1671:10.1098/rstb.2016.0020
1144:
1127:Woody bamboos (tribes
888:
868:
844:
497:Speciation experiments
477:Experimental evolution
432:Evolutionary economics
254:Recent human evolution
112:Processes and outcomes
3774:Geological time units
3039:10.1093/molbev/msp175
2978:10.1007/s002390010234
2874:10.1093/sysbio/syz022
2420:10.1093/sysbio/syw060
2301:10.1093/molbev/msi145
2198:10.1093/sysbio/syv120
2157:10.1093/molbev/msr227
1890:Claramunt, S (2022).
1764:10.1093/molbev/mss075
1624:10.1093/molbev/msl150
1463:10.1073/pnas.58.1.142
1361:10.1073/pnas.50.4.672
1213:molecular systematics
1151:techniques and later
1126:
1085:small population size
1028:Total evidence dating
1011:Expansion calibration
889:
869:
845:
810:. Later, the work of
457:Evolutionary medicine
402:Biosocial criminology
368:History of speciation
281:Evolutionary taxonomy
244:Timeline of evolution
3826:Law of superposition
3821:Isotope geochemistry
2788:on 16 December 2012.
1702:Sanderson M (1997).
887:{\displaystyle \mu }
878:
867:{\displaystyle \mu }
858:
843:{\displaystyle \mu }
834:
743:genetic equidistance
427:Evolutionary ecology
41:Evolutionary biology
4092:Molecular evolution
3959:Fluorine absorption
3936:Luminescence dating
3831:Luminescence dating
3739:Milankovitch cycles
3579:Proleptic Gregorian
3411:Hindu units of time
2970:2001JMolE..53..447F
2883:20.500.11850/373053
2686:2005EmuAO.105..181R
2565:2003JMolE..57S.201D
2514:1987JMolE..26...74O
2463:(1742): 3491–3500.
2351:2014PNAS..111E2957H
2335:(29): E2957–E2966.
2104:10.1038/nature10231
2049:2004Sci...306.1561S
2043:(5701): 1561–1565.
1908:2022MEcEv..13.2331C
1568:1968Natur.217..624K
1509:1967Sci...158.1200S
1503:(3805): 1200–1203.
1454:1967PNAS...58..142S
1352:1963PNAS...50..672M
1274:Molecular evolution
1077:Tube-nosed seabirds
960:probability density
896:molecular evolution
529:Social implications
517:Universal Darwinism
507:Island biogeography
442:Evolutionary ethics
407:Ecological genetics
353:Molecular evolution
291:Transitional fossil
119:Population genetics
35:Part of a series on
4097:Molecular genetics
4061:Terminus post quem
4041:Synchronoptic view
4008:Linguistic methods
3969:Obsidian hydration
3904:Radiometric dating
3889:Incremental dating
3811:Chronostratigraphy
3077:. Springer, Cham.
2862:Systematic Biology
2522:10.1007/BF02111283
2398:Systematic Biology
2185:Systematic Biology
1847:Trends in Genetics
1665:(1699): 20160020.
1264:Y-chromosomal Adam
1172:and not on direct
1161:Joseph Felsenstein
1149:maximum likelihood
1145:
1089:Francisco J. Ayala
1060:negative selection
938:phylogenetic trees
906:in an individual.
898:are neutral, then
884:
864:
840:
784:) and chimpanzee (
776:relative rate test
747:Emanuel Margoliash
560:Theistic evolution
492:Selective breeding
204:Parallel evolution
169:Adaptive radiation
4074:
4073:
3987:
3986:
3844:
3843:
3705:
3704:
3660:Geologic Calendar
3522:
3521:
3092:978-3-030-60180-5
3033:(11): 2595–2603.
2801:Biological Theory
2098:(7357): 493–496.
1714:(12): 1218–1231.
1562:(5129): 624–626.
1260:Mitochondrial Eve
1153:Bayesian modeling
1137:phylogenetic tree
1098:Population size (
708:Émile Zuckerkandl
668:when two or more
646:
645:
337:Origin of Species
139:Natural selection
4109:
4066:ASPRO chronology
4015:Glottochronology
3931:Tephrochronology
3879:Dendrochronology
3857:
3856:
3754:
3753:
3553:Proleptic Julian
3543:Pre-Julian Roman
3533:
3532:
3328:
3327:
3244:
3237:
3230:
3221:
3220:
3194:
3178:
3141:
3104:
3061:
3060:
3050:
3018:
3012:
3004:
2998:
2997:
2964:(4–5): 447–455.
2953:
2947:
2946:
2936:
2926:
2902:
2896:
2895:
2885:
2857:
2851:
2848:
2834:
2816:
2796:
2790:
2789:
2784:. Archived from
2753:
2747:
2746:
2736:
2712:
2706:
2705:
2669:
2660:
2654:
2653:
2635:
2609:
2603:
2602:
2576:
2548:
2542:
2541:
2497:
2491:
2490:
2480:
2448:
2442:
2441:
2431:
2413:
2389:
2383:
2382:
2372:
2362:
2344:
2320:
2314:
2313:
2303:
2294:(7): 1561–1568.
2279:
2270:
2269:
2259:
2227:
2221:
2220:
2210:
2200:
2176:
2170:
2169:
2159:
2135:
2126:
2125:
2115:
2083:
2077:
2076:
2028:
2022:
2021:
2011:
1987:
1981:
1980:
1944:
1938:
1937:
1919:
1887:
1881:
1880:
1870:
1838:
1827:
1826:
1816:
1792:
1786:
1785:
1775:
1758:(8): 1969–1973.
1743:
1734:
1733:
1723:
1699:
1693:
1692:
1682:
1650:
1637:
1636:
1626:
1602:
1596:
1595:
1576:10.1038/217624a0
1551:
1545:
1544:
1492:
1486:
1485:
1475:
1465:
1433:
1427:
1426:
1390:
1384:
1383:
1373:
1363:
1331:
1325:
1324:
1314:
1299:
1284:Glottochronology
1114:silent mutations
998:dataset for the
932:Node calibration
893:
891:
890:
885:
873:
871:
870:
865:
849:
847:
846:
841:
808:phenomenological
650:molecular clock
638:
631:
624:
611:
606:
605:
598:
594:
593:
570:Level of support
363:Current research
348:Modern synthesis
343:Before synthesis
296:Extinction event
54:Darwin's finches
51:
32:
31:
25:Biological clock
4117:
4116:
4112:
4111:
4110:
4108:
4107:
4106:
4077:
4076:
4075:
4070:
4019:
4003:
3999:Molecular clock
3992:Genetic methods
3983:
3964:Nitrogen dating
3951:Relative dating
3945:
3914:Potassium–argon
3861:Absolute dating
3851:
3840:
3799:
3778:
3743:
3719:Cosmic Calendar
3711:Astronomic time
3701:
3643:
3598:
3562:
3548:Original Julian
3518:
3485:
3447:
3346:Ab urbe condita
3324:
3301:
3253:
3248:
3201:
3122:10.1038/nrg1659
3093:
3069:
3067:Further reading
3064:
3019:
3015:
3005:
3001:
2954:
2950:
2903:
2899:
2868:(6): 987–1002.
2858:
2854:
2837:
2814:10.1.1.534.4502
2797:
2793:
2754:
2750:
2713:
2709:
2694:10.1071/MU04039
2667:
2661:
2657:
2610:
2606:
2549:
2545:
2498:
2494:
2449:
2445:
2390:
2386:
2321:
2317:
2280:
2273:
2242:(9): 20150194.
2236:Biology Letters
2228:
2224:
2177:
2173:
2136:
2129:
2084:
2080:
2029:
2025:
1988:
1984:
1945:
1941:
1888:
1884:
1853:(11): 637–650.
1839:
1830:
1793:
1789:
1744:
1737:
1700:
1696:
1651:
1640:
1603:
1599:
1552:
1548:
1493:
1489:
1434:
1430:
1407:10.1038/nrg1659
1391:
1387:
1332:
1328:
1300:
1296:
1292:
1241:
1209:
1055:
1049:—are employed.
1043:
1030:
1013:
984:
982:Tip calibration
934:
912:
879:
876:
875:
859:
856:
855:
835:
832:
831:
804:
787:Pan troglodytes
718:differences in
704:
662:deduce the time
642:
601:
588:
587:
580:
579:
530:
522:
521:
392:
384:
383:
382:
310:
302:
301:
300:
249:Human evolution
239:History of life
223:
222:Natural history
215:
214:
213:
113:
105:
60:
28:
17:
12:
11:
5:
4115:
4105:
4104:
4099:
4094:
4089:
4072:
4071:
4069:
4068:
4063:
4058:
4053:
4048:
4043:
4038:
4036:New Chronology
4033:
4027:
4025:
4024:Related topics
4021:
4020:
4018:
4017:
4011:
4009:
4005:
4004:
4002:
4001:
3995:
3993:
3989:
3988:
3985:
3984:
3982:
3981:
3976:
3971:
3966:
3961:
3955:
3953:
3947:
3946:
3944:
3943:
3938:
3933:
3928:
3927:
3926:
3921:
3916:
3911:
3901:
3899:Paleomagnetism
3896:
3891:
3886:
3881:
3876:
3871:
3865:
3863:
3854:
3846:
3845:
3842:
3841:
3839:
3838:
3833:
3828:
3823:
3818:
3813:
3807:
3805:
3801:
3800:
3798:
3797:
3792:
3786:
3784:
3780:
3779:
3777:
3776:
3771:
3766:
3760:
3758:
3751:
3745:
3744:
3742:
3741:
3736:
3731:
3726:
3721:
3715:
3713:
3707:
3706:
3703:
3702:
3700:
3699:
3697:New Earth Time
3694:
3689:
3688:
3687:
3682:
3672:
3667:
3662:
3657:
3651:
3649:
3645:
3644:
3642:
3641:
3636:
3626:
3621:
3606:
3604:
3600:
3599:
3597:
3596:
3591:
3586:
3581:
3576:
3570:
3568:
3564:
3563:
3561:
3560:
3558:Revised Julian
3555:
3550:
3545:
3539:
3537:
3530:
3524:
3523:
3520:
3519:
3517:
3516:
3511:
3506:
3501:
3495:
3493:
3487:
3486:
3484:
3483:
3478:
3476:Lists of kings
3473:
3468:
3466:Canon of Kings
3463:
3457:
3455:
3449:
3448:
3446:
3445:
3444:
3443:
3438:
3433:
3428:
3418:
3408:
3403:
3398:
3393:
3391:Before present
3388:
3383:
3378:
3373:
3368:
3363:
3358:
3349:
3342:
3336:
3334:
3325:
3323:
3322:
3317:
3312:
3306:
3303:
3302:
3300:
3299:
3294:
3289:
3288:
3287:
3277:
3272:
3267:
3261:
3259:
3255:
3254:
3247:
3246:
3239:
3232:
3224:
3218:
3217:
3212:
3207:
3200:
3199:External links
3197:
3196:
3195:
3179:
3153:(2): 155–178.
3142:
3116:(8): 654–662.
3105:
3091:
3068:
3065:
3063:
3062:
3013:
2999:
2948:
2897:
2852:
2850:
2849:
2807:(4): 357–371.
2791:
2748:
2727:(3): 457–473.
2707:
2680:(2): 181–186.
2655:
2604:
2574:10.1.1.535.897
2543:
2508:(1–2): 74–86.
2492:
2443:
2384:
2315:
2271:
2222:
2191:(3): 449–464.
2171:
2150:(2): 707–719.
2127:
2078:
2023:
2002:(3): 552–569.
1982:
1939:
1882:
1828:
1807:(2): 301–302.
1801:Bioinformatics
1787:
1735:
1694:
1638:
1597:
1546:
1487:
1448:(1): 142–148.
1428:
1401:(8): 654–662.
1385:
1346:(4): 672–679.
1326:
1293:
1291:
1288:
1287:
1286:
1281:
1276:
1271:
1266:
1257:
1252:
1247:
1245:Charles Darwin
1240:
1237:
1217:macroevolution
1208:
1205:
1121:
1120:
1117:
1110:non-coding DNA
1106:
1103:
1096:
1054:
1051:
1042:
1039:
1029:
1026:
1012:
1009:
983:
980:
933:
930:
911:
908:
883:
863:
839:
814:developed the
803:
800:
759:Vincent Sarich
703:
700:
694:sequences for
644:
643:
641:
640:
633:
626:
618:
615:
614:
613:
612:
599:
582:
581:
578:
577:
572:
567:
562:
557:
552:
550:Social effects
547:
542:
537:
531:
528:
527:
524:
523:
520:
519:
514:
509:
504:
499:
494:
489:
484:
479:
474:
469:
464:
459:
454:
449:
444:
439:
434:
429:
424:
419:
414:
409:
404:
399:
393:
390:
389:
386:
385:
381:
380:
370:
365:
360:
355:
350:
345:
340:
333:
328:
323:
318:
312:
311:
308:
307:
304:
303:
299:
298:
293:
288:
283:
278:
276:Classification
273:
268:
263:
258:
257:
256:
246:
241:
236:
234:Common descent
231:
229:Origin of life
225:
224:
221:
220:
217:
216:
212:
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206:
201:
196:
191:
186:
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176:
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136:
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126:
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115:
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111:
110:
107:
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103:
98:
93:
87:
86:
81:
76:
71:
65:
62:
61:
52:
44:
43:
37:
36:
21:Chemical clock
15:
9:
6:
4:
3:
2:
4114:
4103:
4102:Phylogenetics
4100:
4098:
4095:
4093:
4090:
4088:
4085:
4084:
4082:
4067:
4064:
4062:
4059:
4057:
4054:
4052:
4049:
4047:
4044:
4042:
4039:
4037:
4034:
4032:
4029:
4028:
4026:
4022:
4016:
4013:
4012:
4010:
4006:
4000:
3997:
3996:
3994:
3990:
3980:
3977:
3975:
3972:
3970:
3967:
3965:
3962:
3960:
3957:
3956:
3954:
3952:
3948:
3942:
3939:
3937:
3934:
3932:
3929:
3925:
3922:
3920:
3917:
3915:
3912:
3910:
3907:
3906:
3905:
3902:
3900:
3897:
3895:
3892:
3890:
3887:
3885:
3882:
3880:
3877:
3875:
3872:
3870:
3867:
3866:
3864:
3862:
3858:
3855:
3853:
3850:Chronological
3847:
3837:
3834:
3832:
3829:
3827:
3824:
3822:
3819:
3817:
3816:Geochronology
3814:
3812:
3809:
3808:
3806:
3802:
3796:
3793:
3791:
3788:
3787:
3785:
3781:
3775:
3772:
3770:
3767:
3765:
3762:
3761:
3759:
3755:
3752:
3750:
3749:Geologic time
3746:
3740:
3737:
3735:
3734:Metonic cycle
3732:
3730:
3729:Galactic year
3727:
3725:
3722:
3720:
3717:
3716:
3714:
3712:
3708:
3698:
3695:
3693:
3690:
3686:
3683:
3681:
3678:
3677:
3676:
3673:
3671:
3670:ISO week date
3668:
3666:
3663:
3661:
3658:
3656:
3653:
3652:
3650:
3646:
3640:
3637:
3634:
3630:
3627:
3625:
3622:
3619:
3615:
3611:
3608:
3607:
3605:
3601:
3595:
3592:
3590:
3587:
3585:
3582:
3580:
3577:
3575:
3572:
3571:
3569:
3565:
3559:
3556:
3554:
3551:
3549:
3546:
3544:
3541:
3540:
3538:
3534:
3531:
3529:
3525:
3515:
3512:
3510:
3507:
3505:
3502:
3500:
3497:
3496:
3494:
3492:
3488:
3482:
3479:
3477:
3474:
3472:
3469:
3467:
3464:
3462:
3459:
3458:
3456:
3454:
3450:
3442:
3439:
3437:
3434:
3432:
3429:
3427:
3424:
3423:
3422:
3419:
3416:
3412:
3409:
3407:
3404:
3402:
3399:
3397:
3394:
3392:
3389:
3387:
3384:
3382:
3379:
3377:
3376:Byzantine era
3374:
3372:
3369:
3367:
3364:
3362:
3359:
3357:
3353:
3350:
3348:
3347:
3343:
3341:
3338:
3337:
3335:
3333:
3332:Calendar eras
3329:
3326:
3321:
3318:
3316:
3313:
3311:
3308:
3307:
3304:
3298:
3295:
3293:
3290:
3286:
3283:
3282:
3281:
3278:
3276:
3273:
3271:
3268:
3266:
3263:
3262:
3260:
3256:
3252:
3245:
3240:
3238:
3233:
3231:
3226:
3225:
3222:
3216:
3213:
3211:
3208:
3206:
3203:
3202:
3192:
3188:
3184:
3183:Zuckerkandl E
3180:
3176:
3172:
3168:
3164:
3160:
3156:
3152:
3148:
3143:
3139:
3135:
3131:
3127:
3123:
3119:
3115:
3111:
3106:
3102:
3098:
3094:
3088:
3084:
3080:
3076:
3071:
3070:
3058:
3054:
3049:
3044:
3040:
3036:
3032:
3028:
3024:
3017:
3010:
3003:
2995:
2991:
2987:
2983:
2979:
2975:
2971:
2967:
2963:
2959:
2952:
2944:
2940:
2935:
2930:
2925:
2920:
2916:
2912:
2908:
2901:
2893:
2889:
2884:
2879:
2875:
2871:
2867:
2863:
2856:
2846:
2845:
2840:
2836:
2835:
2832:
2828:
2824:
2820:
2815:
2810:
2806:
2802:
2795:
2787:
2783:
2779:
2775:
2771:
2767:
2763:
2759:
2752:
2744:
2740:
2735:
2730:
2726:
2722:
2718:
2711:
2703:
2699:
2695:
2691:
2687:
2683:
2679:
2675:
2674:
2666:
2659:
2651:
2647:
2643:
2639:
2634:
2629:
2625:
2621:
2620:
2615:
2608:
2600:
2596:
2592:
2588:
2584:
2580:
2575:
2570:
2566:
2562:
2558:
2554:
2547:
2539:
2535:
2531:
2527:
2523:
2519:
2515:
2511:
2507:
2503:
2496:
2488:
2484:
2479:
2474:
2470:
2466:
2462:
2458:
2454:
2447:
2439:
2435:
2430:
2425:
2421:
2417:
2412:
2407:
2403:
2399:
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2380:
2376:
2371:
2366:
2361:
2356:
2352:
2348:
2343:
2338:
2334:
2330:
2326:
2319:
2311:
2307:
2302:
2297:
2293:
2289:
2285:
2278:
2276:
2267:
2263:
2258:
2253:
2249:
2245:
2241:
2237:
2233:
2226:
2218:
2214:
2209:
2204:
2199:
2194:
2190:
2186:
2182:
2175:
2167:
2163:
2158:
2153:
2149:
2145:
2141:
2134:
2132:
2123:
2119:
2114:
2109:
2105:
2101:
2097:
2093:
2089:
2082:
2074:
2070:
2066:
2062:
2058:
2054:
2050:
2046:
2042:
2038:
2034:
2027:
2019:
2015:
2010:
2005:
2001:
1997:
1993:
1986:
1978:
1974:
1970:
1966:
1962:
1958:
1954:
1950:
1943:
1935:
1931:
1927:
1923:
1918:
1913:
1909:
1905:
1901:
1897:
1893:
1886:
1878:
1874:
1869:
1864:
1860:
1856:
1852:
1848:
1844:
1837:
1835:
1833:
1824:
1820:
1815:
1810:
1806:
1802:
1798:
1791:
1783:
1779:
1774:
1769:
1765:
1761:
1757:
1753:
1749:
1742:
1740:
1731:
1727:
1722:
1717:
1713:
1709:
1705:
1698:
1690:
1686:
1681:
1676:
1672:
1668:
1664:
1660:
1656:
1649:
1647:
1645:
1643:
1634:
1630:
1625:
1620:
1616:
1612:
1608:
1601:
1593:
1589:
1585:
1581:
1577:
1573:
1569:
1565:
1561:
1557:
1550:
1542:
1538:
1534:
1530:
1526:
1522:
1518:
1514:
1510:
1506:
1502:
1498:
1491:
1483:
1479:
1474:
1469:
1464:
1459:
1455:
1451:
1447:
1443:
1439:
1432:
1424:
1420:
1416:
1412:
1408:
1404:
1400:
1396:
1389:
1381:
1377:
1372:
1367:
1362:
1357:
1353:
1349:
1345:
1341:
1337:
1330:
1322:
1318:
1313:
1308:
1304:
1303:Zuckerkandl E
1298:
1294:
1285:
1282:
1280:
1277:
1275:
1272:
1270:
1267:
1265:
1261:
1258:
1256:
1253:
1251:
1248:
1246:
1243:
1242:
1236:
1234:
1230:
1226:
1222:
1218:
1214:
1204:
1202:
1198:
1194:
1189:
1186:
1182:
1177:
1175:
1171:
1166:
1162:
1158:
1154:
1150:
1142:
1138:
1134:
1130:
1129:Arundinarieae
1125:
1118:
1115:
1111:
1107:
1104:
1101:
1100:Genetic drift
1097:
1094:
1093:
1092:
1090:
1086:
1082:
1078:
1074:
1068:
1065:
1061:
1050:
1048:
1047:phylogenomics
1038:
1036:
1025:
1023:
1018:
1008:
1005:
1004:morphological
1002:along with a
1001:
997:
993:
989:
979:
977:
973:
969:
965:
961:
957:
953:
952:stratigraphic
948:
944:
939:
929:
927:
923:
918:
907:
905:
901:
897:
881:
861:
853:
837:
829:
825:
821:
817:
813:
809:
799:
797:
793:
789:
788:
783:
782:
777:
772:
768:
764:
760:
755:
752:
748:
744:
739:
737:
733:
729:
725:
721:
717:
713:
712:Linus Pauling
709:
699:
697:
693:
689:
685:
681:
678:
674:
671:
667:
663:
659:
655:
654:mutation rate
651:
639:
634:
632:
627:
625:
620:
619:
617:
616:
610:
600:
597:
592:
586:
585:
584:
583:
576:
573:
571:
568:
566:
563:
561:
558:
556:
553:
551:
548:
546:
543:
541:
538:
536:
533:
532:
526:
525:
518:
515:
513:
510:
508:
505:
503:
500:
498:
495:
493:
490:
488:
485:
483:
482:Phylogenetics
480:
478:
475:
473:
470:
468:
465:
463:
460:
458:
455:
453:
450:
448:
445:
443:
440:
438:
435:
433:
430:
428:
425:
423:
420:
418:
415:
413:
410:
408:
405:
403:
400:
398:
395:
394:
388:
387:
378:
374:
371:
369:
366:
364:
361:
359:
356:
354:
351:
349:
346:
344:
341:
339:
338:
334:
332:
329:
327:
326:Before Darwin
324:
322:
319:
317:
314:
313:
306:
305:
297:
294:
292:
289:
287:
284:
282:
279:
277:
274:
272:
269:
267:
264:
262:
259:
255:
252:
251:
250:
247:
245:
242:
240:
237:
235:
232:
230:
227:
226:
219:
218:
210:
207:
205:
202:
200:
197:
195:
192:
190:
187:
185:
182:
180:
177:
175:
172:
170:
167:
165:
162:
160:
157:
155:
154:Genetic drift
152:
150:
147:
145:
142:
140:
137:
135:
132:
130:
127:
125:
122:
120:
117:
116:
109:
108:
102:
99:
97:
94:
92:
89:
88:
85:
82:
80:
77:
75:
72:
70:
67:
66:
64:
63:
59:
55:
50:
46:
45:
42:
39:
38:
34:
33:
30:
26:
22:
3998:
3979:Stratigraphy
3924:Uranium–lead
3894:Lichenometry
3692:Winter count
3675:Mesoamerican
3603:Astronomical
3421:Mesoamerican
3406:Sothic cycle
3381:Seleucid era
3366:Bosporan era
3354: /
3344:
3292:Paleontology
3190:
3150:
3146:
3113:
3109:
3074:
3030:
3026:
3016:
3002:
2961:
2957:
2951:
2914:
2911:PLOS Biology
2910:
2900:
2865:
2861:
2855:
2844:ScienceDaily
2842:
2804:
2800:
2794:
2786:the original
2768:(1): 71–75.
2765:
2761:
2751:
2724:
2720:
2710:
2677:
2671:
2658:
2626:(1): 35–55.
2623:
2617:
2607:
2556:
2552:
2546:
2505:
2501:
2495:
2460:
2456:
2446:
2404:(1): 57–73.
2401:
2397:
2387:
2332:
2328:
2318:
2291:
2287:
2239:
2235:
2225:
2188:
2184:
2174:
2147:
2143:
2095:
2091:
2081:
2040:
2036:
2026:
1999:
1995:
1985:
1955:(1): 41–49.
1952:
1948:
1942:
1899:
1895:
1885:
1850:
1846:
1804:
1800:
1790:
1755:
1751:
1711:
1707:
1697:
1662:
1658:
1617:(1): 26–53.
1614:
1610:
1600:
1559:
1555:
1549:
1500:
1496:
1490:
1445:
1441:
1431:
1398:
1394:
1388:
1343:
1339:
1329:
1316:
1297:
1225:phylogenetic
1210:
1190:
1178:
1156:
1146:
1073:mockingbirds
1069:
1056:
1044:
1031:
1014:
985:
935:
913:
812:Motoo Kimura
805:
796:K-T boundary
785:
781:Homo sapiens
779:
763:Allan Wilson
756:
751:cytochrome c
742:
740:
735:
728:evolutionary
705:
658:biomolecules
649:
647:
502:Sociobiology
487:Paleontology
335:
271:Biogeography
266:Biodiversity
184:Coextinction
174:Co-operation
149:Polymorphism
74:Introduction
29:
3919:Radiocarbon
3594:Dual dating
3453:Regnal year
3431:Short Count
3371:Bostran era
3352:Anno Domini
3285:Big History
3265:Archaeology
1250:Gene orders
1000:extant taxa
972:exponential
917:ancient DNA
910:Calibration
757:Similarly,
512:Systematics
321:Renaissance
199:Convergence
189:Contingency
179:Coevolution
4081:Categories
3514:Vietnamese
3426:Long Count
3361:Anno Mundi
3356:Common Era
3258:Key topics
3251:Chronology
3187:Pauling LB
2917:(5): e88.
2411:1506.04797
2208:2263/53371
1290:References
1181:saturation
1081:generation
988:tip dating
956:taphonomic
922:calibrated
720:hemoglobin
716:amino acid
692:amino acid
677:nucleotide
670:life forms
666:prehistory
286:Cladistics
209:Extinction
194:Divergence
164:Speciation
144:Adaptation
58:John Gould
4051:Year zero
4031:Chronicle
3974:Seriation
3909:Lead–lead
3783:Standards
3764:Deep time
3724:Ephemeris
3610:Lunisolar
3574:Gregorian
3567:Gregorian
3528:Calendars
3491:Era names
3461:Anka year
3340:Human Era
3270:Astronomy
3101:231672167
2809:CiteSeerX
2762:BioEssays
2642:0004-8038
2569:CiteSeerX
2342:1310.2968
1934:252353611
1926:2041-210X
1170:inference
1133:Bambuseae
996:molecular
968:lognormal
904:mutations
900:fixations
882:μ
862:μ
838:μ
824:mutations
680:sequences
545:Dysgenics
261:Phylogeny
159:Gene flow
129:Diversity
124:Variation
4046:Timeline
3884:Ice core
3757:Concepts
3504:Japanese
3436:Tzolk'in
3401:Egyptian
3167:11620303
3138:14261833
3130:16136655
3057:19661199
2986:11675604
2943:16683862
2892:31111152
2831:28166727
2782:10070256
2702:20390465
2650:51797284
2599:23887665
2591:15008417
2487:22628470
2438:28173531
2379:25009181
2310:15814826
2266:26333662
2217:26683588
2166:21926069
2122:21753753
2073:27134675
2065:15567864
1969:25681677
1877:26439502
1823:12538260
1782:22367748
1730:17647010
1689:27325838
1633:17047029
1423:14261833
1415:16136655
1380:14077496
1309:(1962).
1239:See also
1193:fixation
1174:evidence
792:Ceboidea
767:Primates
724:linearly
696:proteins
673:diverged
609:Category
535:Eugenics
377:timeline
358:Evo-devo
316:Overview
134:Mutation
96:Evidence
91:Glossary
4056:Floruit
3804:Methods
3665:Iranian
3633:Islamic
3499:Chinese
3310:Periods
3280:History
3275:Geology
3175:5660841
3048:2912466
2994:9791493
2966:Bibcode
2934:1395354
2743:1584014
2682:Bibcode
2561:Bibcode
2538:8260277
2530:3125340
2510:Bibcode
2478:3396900
2429:5410945
2370:4115571
2347:Bibcode
2257:4614420
2113:3154645
2045:Bibcode
2037:Science
2018:1316531
1977:3895351
1904:Bibcode
1773:3408070
1680:4920342
1592:4161261
1584:5637732
1564:Bibcode
1541:7349579
1533:4964406
1525:1722843
1505:Bibcode
1497:Science
1482:4962458
1450:Bibcode
1348:Bibcode
1321:189–225
1307:Pauling
1229:fossils
1197:alleles
1141:Olyreae
1041:Methods
1035:lineage
828:fitness
820:haploid
771:albumin
732:protein
101:History
84:Outline
3852:dating
3648:Others
3614:Hebrew
3509:Korean
3320:Epochs
3173:
3165:
3136:
3128:
3099:
3089:
3055:
3045:
2992:
2984:
2941:
2931:
2890:
2829:
2811:
2780:
2741:
2700:
2648:
2640:
2597:
2589:
2571:
2536:
2528:
2485:
2475:
2436:
2426:
2377:
2367:
2308:
2264:
2254:
2215:
2164:
2120:
2110:
2092:Nature
2071:
2063:
2016:
1975:
1967:
1932:
1924:
1875:
1821:
1780:
1770:
1728:
1687:
1677:
1631:
1590:
1582:
1556:Nature
1539:
1531:
1523:
1480:
1473:335609
1470:
1421:
1413:
1378:
1371:221244
1368:
1219:, and
1185:linear
964:normal
926:fossil
607:
331:Darwin
3685:Aztec
3629:Lunar
3624:Solar
3618:Hindu
3481:Limmu
3441:Haab'
3396:Hijri
3171:S2CID
3134:S2CID
3097:S2CID
2990:S2CID
2827:S2CID
2698:S2CID
2668:(PDF)
2646:S2CID
2595:S2CID
2534:S2CID
2406:arXiv
2337:arXiv
2069:S2CID
1973:S2CID
1930:S2CID
1726:S2CID
1588:S2CID
1537:S2CID
1521:JSTOR
1419:S2CID
1201:fixed
976:gamma
947:clade
852:fixed
690:, or
69:Index
3680:Maya
3415:Yuga
3315:Eras
3297:Time
3163:PMID
3126:PMID
3087:ISBN
3053:PMID
2982:PMID
2939:PMID
2888:PMID
2778:PMID
2739:PMID
2638:ISSN
2587:PMID
2526:PMID
2483:PMID
2434:PMID
2375:PMID
2306:PMID
2262:PMID
2213:PMID
2162:PMID
2118:PMID
2061:PMID
2014:PMID
1965:PMID
1922:ISSN
1873:PMID
1819:PMID
1778:PMID
1685:PMID
1629:PMID
1580:PMID
1529:PMID
1478:PMID
1411:PMID
1376:PMID
1262:and
1233:taxa
1207:Uses
1165:MCMC
1131:and
992:taxa
761:and
741:The
710:and
682:for
648:The
79:Main
3155:doi
3118:doi
3079:doi
3043:PMC
3035:doi
2974:doi
2929:PMC
2919:doi
2878:hdl
2870:doi
2819:doi
2770:doi
2729:doi
2690:doi
2678:105
2673:Emu
2628:doi
2624:118
2619:Auk
2579:doi
2518:doi
2473:PMC
2465:doi
2461:279
2424:PMC
2416:doi
2365:PMC
2355:doi
2333:111
2296:doi
2252:PMC
2244:doi
2203:hdl
2193:doi
2152:doi
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