"Although ubiquitous, this approach has an inherent circularity and is subject to possible error because it relies on the accuracy of the ages of fossils." — Kevin Langergraber and colleagues, on the old fossil-calibrated mutation rate, PNAS, 20121
"To say that God could have created the pattern is merely ad hoc." — Todd Charles Wood, young-earth creationist biologist, 20062
Summary
Humans and chimpanzees differ at about 1.2 % of the DNA letters that can be lined up between them, roughly 35 million single-letter differences.3 How long would mutation take to make them? The answer depends on the mutation rate, and for twenty years the field has had two rates that disagree by a factor of two. This article works the arithmetic for both, then asks what a young-earth model has to say instead, and whether the evidence favors it.
- The old rate was partly circular. The "phylogenetic" rate of about 1 × 10-9 changes per DNA letter per year was set by dividing the human-chimp difference by an assumed divergence date of about 7 million years.1 The "pedigree" rate is counted directly, by sequencing parents and children: 1.20 × 10-8 per letter per generation in 78 Icelandic families.4 At human generation lengths of 25 to 29 years that is 0.41 to 0.48 × 10-9 per year, about half the old rate.5
- Half the rate means about twice the date. On the old rate, the leading genome model put the human-chimp split at 3.7 million years; on the measured rate, at about 6.6 Published estimates that use measured rates now run from 5.5-6.3 million years, from Yoo and colleagues in 2025 on complete ape genomes, through 6.5-9.3 from Moorjani and colleagues in 2016, to 7-13 from Langergraber and colleagues in 2012.178
- No single steady rate fits every fossil calibration, and the mainstream says so. The family rate fixes the human-chimp split but pushes the human-orangutan divergence to 34-46 million years, against a fossil limit under 18.9 The usual repair is a slowdown in the yearly rate as ape generations lengthened. Part of that repair is now measured: chimpanzee, gorilla and orangutan families mutate about 1.48 times faster per year than human families.10 This is a real weakness of the clock. It does not help a young earth: the youngest of these dates, 3.75 million years, is 500 times the roughly 7,500 years since creation on the Septuagint, the longest biblical chronology.
- About a third of the genome does not follow the family tree. In 30 % of the genome, gorilla is closer to human or to chimpanzee than those two are to each other, split evenly between the two patterns, rarer near genes and rarer on the X chromosome.666 The 2025 complete genomes raise it to 39.5 %.8 On common descent this is "incomplete lineage sorting", and with the measured mutation rate it implies an ancestral population of tens of thousands, about 200,000 in the newest estimate. On separate creation the three genomes would have had to be made with this mosaic already in them.
- Young-earth geneticists accept the measured differences and say most were created. Robert Carter writes that dating the ape lineages is only "uncovering the differences God built into their initial genomes."11 The arithmetic leaves no other option. At measured rates, and even allowing for the patriarchs' great ages, mutation since creation adds about 7,000 to 9,000 changes to each lineage, depending on the chronology and the generation length. Both lineages together make 0.04 to 0.05 % of the 35 million differences. Mutation would have to run about 1,900 to 2,500 times faster to make them all.12
- The "created" differences carry the fingerprints of mutation. A quarter of the human-chimp differences sit at CpG sites, which are 2 % of the DNA,3 and CpG sites are the hottest spots for new mutations in families.4 The Y chromosome differs most and the X least, in the ratio a male-biased mutation process predicts,3 and fathers pass on about four times as many new mutations as mothers.4 A designer could have made created differences look like this. No young-earth model predicts it, and Wood's warning about ad hoc appeals applies.
- The waiting-time argument is the young-earth side's strongest point, and it proves less than is claimed. John Sanford's simulations find 84 million years to establish one specific two-letter DNA string in a hominin population, and 2 billion years for five letters.13 Mainstream mathematicians agree in substance: 216 million years for one prespecified pair of regulatory mutations in humans.14 But the wait shrinks in proportion to the number of acceptable targets. Inside 6 million years a two-letter string needs 15 equally good alternatives; a five-letter string needs 385.15 Nobody on either side has counted how many human-specific changes needed one exact multi-letter target.
Verdict. The family-measured mutation rate is real, it roughly doubles the dates, and it leaves the clock needing a rate slowdown that is plausible and partly measured but not proven. A young-earth model must say that more than 99.9 % of the single-letter differences were created. The pattern of those differences is the pattern mutation and shared ancestry predict: the CpG excess, the sex-chromosome ratio, and the mosaic of incomplete lineage sorting. A created-differences model fits that pattern only by stipulation. On this evidence the mutation reading is better supported. The waiting-time problem is a real constraint on prespecified multi-letter changes, but it has not been shown to apply to the changes that actually separate the two species.
Part 1: Two clocks that disagree by a factor of two
A mutation rate is the chance that one DNA letter changes in one generation. Turning it into a clock needs two more numbers: how many letters differ between two species, and how many years a generation lasts. Because both lineages mutate after they part, the time since their last shared ancestor is the difference divided by twice the yearly rate.5
The phylogenetic rate was set by the date it was later used to find
Before 2010 almost every human-chimp date used a rate of about 1 × 10-9 per letter per year. Langergraber and colleagues explain where it came from. It was "derived from the observed DNA sequence difference of around 1.3% between the human and chimpanzee genomes ... and an assumed DNA sequence divergence between these species at 7 Ma," with a second calibration from human and macaque at an assumed 25 million years.1 The arithmetic checks: 1.3 % divided by twice 7 million years is 0.93 × 10-9.5
A date found with this rate is therefore partly the date that went into it. Langergraber's own team calls the method circular. The same paper also records the clock's best success. In 1967 Vincent Sarich and Allan Wilson used immunological data to put the human-ape split at only about 5 million years, "some three to four times more recently than had been assumed on the basis of the fossil record." DNA studies later put it at 4 to 6 million years again and again.1
The pedigree rate is counted, not assumed
Since 2010, whole-genome sequencing of parents and children has let geneticists count new mutations directly. Augustine Kong and colleagues at deCODE genetics sequenced 78 Icelandic families and found 4,933 new mutations, a rate of 1.20 × 10-8 per letter per generation.4 The count rose with the father's age by about two mutations per year of his age.4 By 2012 four such family studies gave 0.97 to 1.36 × 10-8 per generation.1 The largest since, of 1,548 Icelandic families in 2017, gave 1.29 × 10-8 per generation, or 0.43 × 10-9 per year.16
The rate per generation is not yet a clock. It has to be divided by the length of a generation, and that is not one number.
| Generation length used | Source | Yearly rate at 1.20 × 10-8 per generation |
|---|---|---|
| 25 years, the older genetics convention | Langergraber 2012 says genetics "commonly used a generation time of 20 or 25 y"1 | 0.48 × 10-9 |
| 26.9 years, human average over 250,000 years | Wang and colleagues 2023, from mutation patterns17 | 0.45 × 10-9 |
| 29 years, living humans | Fenner 2005, cited by Langergraber1 | 0.41 × 10-9 |
| 24.6 years, wild chimpanzees | Langergraber 2012, 226 offspring with known parents1 | 0.49 × 10-9 |
Every row is about half the phylogenetic rate. Moorjani and colleagues put it plainly: the family rate "is approximately twofold lower than the mutation rate inferred from the number of substitutions observed between primates."7 Aylwin Scally and Richard Durbin, reviewing the change in 2012, called the measured value "approximately half of that previously derived from fossil calibration."18 Generation length alone cannot close the gap. They calculated that a generation of 10 to 15 years would be "needed for the de novo and phylogenetic rates to match," which "would be shorter than any other living great ape and less than half of the value measured in present-day humans."18
A third method gives a faster rate
Counting new single-letter mutations is not the only direct route. James Sun and colleagues counted 2,058 new mutations at short repeated DNA segments in 85,289 Icelanders, then used the known behaviour of those segments to infer the single-letter rate. They got 1.82 × 10-8 per generation, 50 % above the family counts.19 The authors suggested the whole-genome studies might be missing some real mutations; the whole-genome studies have since grown much larger and kept their lower value.16 The most complete count so far, published by David Porubsky and colleagues in 2025, read a four-generation family of 28 people with five sequencing technologies. It covered 91.9 % of the 22 ordinary chromosomes, including the repeats and centromeres earlier studies could not read, and found 1.17 × 10-8 per letter per generation.20 New mutations were commoner in those repeats, but not common enough to raise the genome-wide rate. So the gap is not explained by mutations hidden in hard-to-read DNA. The disagreement between methods is itself one of the uncertainties in the dates below.
The other apes mutate faster per year
If the human rate per year had always been what it is now, the human, chimpanzee and gorilla lineages should show the same amount of change since they parted. They nearly do. Moorjani's team found substitution rates "∼2% higher in chimpanzees and ∼7% higher in the gorilla than in humans."7 Yet Søren Besenbacher and colleagues, sequencing chimpanzee, gorilla and orangutan families, found these apes now mutate about 1.48 times faster per year than humans.10 The two findings fit only if the human slowdown is recent, and that is how Besenbacher reads it: "an appreciable slowdown in the yearly mutation rate in the human lineage that is likely to be recent as genome comparisons almost adhere to a molecular clock." The favoured cause is life history. Most new mutations are counted per generation, so a species that breeds younger collects more of them per year, and Scally and colleagues note that ape body sizes and generation times grew over this period.6 Generation length alone is not enough: as shown above, it would need 10- to 15-year generations. Guy Amster and Guy Sella's 2016 model adds "the age of onset of male puberty and rates of spermatogenesis," since most new mutations arise in fathers. With those, they report, it "largely bridges the gap reported between estimates of split times based on fossil and molecular evidence," and the split can be as recent as 6.6 million years.21 The price is inputs nobody can measure: how old ancestral apes were at puberty and at fatherhood millions of years ago.
Part 2: From a difference to a date
Two clocks run inside every comparison
A human and a chimpanzee DNA sequence did not start to differ on the day the two populations parted. The ancestral population already carried variation, so two gene copies picked from it had usually split long before. Langergraber and colleagues state it this way: the differences seen today are "the sum of two values," those that built up after the two lineages stopped interbreeding and "the sequence differences that correspond to the diversity in the common ancestor of both lineages."1 The 2005 chimpanzee genome paper splits the time the same way, into the time since speciation and the coalescence time in the ancestral population, which it put on the order of 1 to 2 million years.3
So a date needs three things: the difference, the yearly rate, and the size of the ancestral population. For two gene copies in a population of N breeding individuals, the average wait back to their common ancestor is 2N generations. The split time is the total divergence time minus that wait.5
The arithmetic, at each rate
Divergence of 1.23 %, the 2005 consortium's figure,3 gives these times since the last shared ancestor of the DNA, before any allowance for the ancestral population:
| Yearly rate (× 10-9) | Where it comes from | Divergence time | Split if ancestral N = 50,000 | Split if N = 100,000 |
|---|---|---|---|---|
| 1.00 | old fossil calibration | 6.2 million years | 3.7 | 1.3 |
| 0.64 | ape families, 1.48 × the human rate | 9.7 | 7.2 | 4.8 |
| 0.51 | 1.25 × 10-8 over 24.5 years, the input Yoo 2025 used | 12.1 | 9.6 | 7.2 |
| 0.45 | 1.20 × 10-8 over 26.9 years | 13.8 | 11.3 | 8.9 |
| 0.41 | 1.20 × 10-8 over 29 years | 14.9 | 12.4 | 10.0 |
All figures are in millions of years, with a 24.5-year generation in the ancestor.5 Two things follow.
First, the old rate leaves almost no room for an ancestral population. At 1 × 10-9 per year, a 6-million-year split allows an ancestral population of only about 3,000, and a 7-million-year split is impossible. The measured rates need a large ancestral population for a 6 to 7 million year split: about 100,000 to 180,000. The 2025 complete-genome study estimated 198,000.8
Second, rate and population size trade against each other. A slower rate makes the total divergence older, and a larger ancestral population takes more of that time away from the split. Much of the spread between published dates comes from different choices of these two inputs.
What the published studies found
| Study | Rate and method | Human-chimp split |
|---|---|---|
| Scally and colleagues 2012 | old rate, 1 × 10-9 per year | 3.7 million years6 |
| Sun and colleagues 2012 | repeat-segment rate, 0.68 × 10-9 per year | 4.97 (3.75 to 6.57)19 |
| Yoo and colleagues 2025 | 1.25 × 10-8 per generation, complete genomes | 5.5 to 6.38 |
| Scally and colleagues 2012 | measured rate, 0.5-0.6 × 10-9 per year | about 66 |
| Besenbacher and colleagues 2019 | ape family rates, 1.48 × the human rate | 6.6 (5.9 to 7.3)10 |
| Moorjani and colleagues 2016 | CpG changes only, which track years better | 7.9 (6.5 to 9.3)7 |
| Langergraber and colleagues 2012 | family rates and measured ape generation lengths | 7 to 131 |
| Sun and colleagues 2012 | their model rerun with Kong's family rate | 8.32 to 11.89 |
Langergraber's team recalculated older published splits with the family rates. Scally's 3.69 million years became 6.78 to 11.03. Others moved from 4.2-4.5 to between 7.8 and 13.5.1
The orangutan problem
The family rate fixes one fossil problem and creates another. On the old rate, the human-chimp split fell after fossils such as Sahelanthropus, dated 6.8 to 7.2 million years, that many paleontologists place on the human side.19 The slower rate moves the split back past them. Whether Sahelanthropus stood on the human line at all is still argued, which makes it a soft calibration either way; that dispute is worked in Sahelanthropus, Orrorin, and Ardipithecus. But Sun's team found that the same slow rate puts the human-orangutan divergence at 34 to 46 million years, "so much older than the upper bound from the fossil record of <18 Mya on human-orangutan speciation that the date is implausible."9
Scally's team proposed the repair now usually accepted: a rate near 1 × 10-9 in the early great apes, slowing as body size and generation length grew in every descendant line. They admitted the cost: Sahelanthropus and an early gorilla fossil "can be accommodated ... only if most of the decrease occurred early in great ape evolution."6 Sun's team was blunter. The slowdown "would need to have been coincidental in both lineages," and "extraordinarily dramatic: about 3-fold in both lineages."9 Besenbacher's ape families have since measured part of that difference directly, and with their faster ape rates the dates come out "much more in line with the fossil record and the biogeography."10
Scally summed up the state of the field in one sentence: "the inability to observe past mutation rates means that the timing of events from genetic data remains uncertain."6 Geneticists said as much in public. At a 2015 meeting in Leipzig on the human mutation rate, David Reich of Harvard Medical School told Nature: "The fact that the clock is so uncertain is very problematic for us," and "It means that the dates we get out of genetics are really quite embarrassingly bad and uncertain." The same report noted that the slow clock "gave nonsensical results when extended further back in time," such as an ape-monkey ancestor that "could have encountered dinosaurs," and that many authors were "presenting multiple dates" to hedge. Reich still advised using the slower rate, with a warning: "This is not the speed of light. This is not physics."22 Young-earth critics quote Reich's words, and on the uncertainty they are right.23 A skeptic should grant the point. So should a creationist reading it as support for a young earth, because every candidate date in this dispute lies between 3.75 and 13 million years. On the Masoretic text's roughly 6,000 years since creation that is 600 to 2,200 times too old; on the Septuagint's roughly 7,500, 500 to 1,700 times. The disagreement is a factor of about three and a half. The gap to a young earth is a factor of a thousand.
Part 3: Incomplete lineage sorting, a third of the genome off the family tree
What it is
Take any stretch of DNA and ask which two of human, chimpanzee and gorilla match best. Most of the time the answer is human and chimpanzee. But in 30 % of the genome, "gorilla is closer to human or chimpanzee than the latter are to each other."6 The complete genomes of 2025 put it at 39.5 % of the autosomes and 24 % of the X chromosome.8
Common descent explains this without strain. The population ancestral to humans and chimpanzees carried many variants, some of which it had inherited from the older population that also gave rise to gorillas. If the time between the gorilla split and the chimpanzee split was short compared with the size of that population, many variants were still unsorted when humans and chimpanzees parted. Each lineage then kept one version or the other more or less at random. Where the human line kept the version the gorilla line also kept, human and gorilla match.
The proportion follows a simple rule. Two gene copies fail to find their common ancestor in the population between the two splits with a chance of e-t/2N, where t is the time between the splits in generations and N the population size. When they fail, two times in three the resulting tree disagrees with the species tree.24 A 30 % disagreement means the time between splits was only about 0.8 times twice the population size. If that population had held 10,000 individuals and lasted 4 million years, the disagreement would be 0.007 %, not 30 %.
ILS fixes only that ratio. It does not fix years, because a population of 100,000 lasting 3.5 million years gives the same share as one of 50,000 lasting 1.7 million. The mutation rate sets the years, which is why the rate dispute and the population-size estimates move together. With the measured rates, published estimates of the ancestral population run from about 50,000 to about 200,000.785
It passes three checks a random mosaic would not
ILS is not a fudge factor added afterward. It predicts specific patterns, and the genomes show them.
- The two odd trees should be equally common. Chance sorting has no favourite. Scally's team found "no significant difference between the number sorting as ((H,G),C) and ((C,G),H)."6 Interbreeding between two of the species after they parted would have favoured one of the two.
- It should be rarer where natural selection shrinks the effective population. Selection on genes removes variation from the DNA around them. ILS drops to 22 % inside coding exons, and "the suppression of ILS extends out to several hundred kbp from coding genes, evident even in raw site patterns before any model inference."6
- It should be rarer on the X chromosome. A population carries three X chromosomes for every four copies of each other chromosome, and selection acts on the single X in males. Scally's team found less ILS on X;6 the complete genomes give 24 % on X against 39.5 % on the other chromosomes.8
The same population variation also accounts for a large share of the human-chimp difference itself. On the mainstream models, between a fifth and three fifths of the 1.2 % was already present as variation before the two lineages parted.5 Nathaniel Jeanson asked evolutionists about such "preexisting differences": "When can preexisting differences be invoked, and when should they not be invoked? What testable predictions does the hypothesis of preexisting differences make?"25 It is a fair question, and ILS is the answer. The preexisting variation of the mainstream model must show up as gene trees that disagree in a known share, evenly between the two alternatives, less near genes and less on X. It does.
What a separate-creation model would need
On a young-earth model humans, chimpanzees and gorillas were created as separate kinds, with no shared ancestral population. The pattern then has to have been built into the created genomes: in about a third of the DNA, human made to match gorilla against chimpanzee or chimpanzee to match gorilla against human, in equal measure, less often near genes and on the X chromosome.
Jeffrey Tomkins and Jerry Bergman, writing in the Journal of Creation in 2013, take ILS as evidence against common descent. They write that ILS "across clearly different and unrelated kinds of organisms represents a mosaic of DNA sequence patterns that cannot be explained by common ancestry."26 On the gorilla genome they say that "no consensus path of common ancestry between humans and various apes exists, and no coherent model of primate evolution could be achieved."26 That misreads the paper they quote. Scally's team fit the 30 % inside one model with one set of speciation times and population sizes, and wrote that "whole genome comparisons can be strongly conclusive about the ordering of speciation events."6 What the genome left uncertain was the timing, not the tree.
No young-earth writer found in this research has offered a created-kinds model that predicts the size of the ILS share, its even split, or its drop near genes and on the X. Jeanson's Replacing Darwin, the fullest young-earth treatment of nuclear DNA differences between species, does not discuss it.27 A designer could have made the three genomes this way. But as Todd Wood wrote of the human-chimp pattern in general, "To say that God could have created the pattern is merely ad hoc."2
Part 4: The young-earth position, in its own words, and its price
Young-earth geneticists accept the measured differences
None of the young-earth sources examined here disputes that humans and chimpanzees differ at tens of millions of DNA letters. Their disagreement is about where the differences came from.
Nathaniel Jeanson, whose 2017 book Replacing Darwin gives the fullest young-earth treatment, accepts the measured rates and the measured differences and runs the evolutionary prediction himself. "Traditionally, evolutionists have put the split time between these species at 4.5 to 7 million years ago." At the measured human and chimpanzee rates that predicts "only around 8.4 million" to "only around 13.1 million" differences, while "over 26 million single base pair differences separate the human and chimpanzee nuclear DNA sequences."25 He then notes that the date moved to fit: "The human-chimpanzee divergence time was revised in light of the mutation rates in these species."25
That observation is correct, and it is the same gap Langergraber and Scally addressed in 2012. Two corrections are needed to his arithmetic. First, his prediction counts only the time since the split and leaves out the variation already present in the ancestral population, which the 2005 chimpanzee genome paper had put at 1 to 2 million years' worth of divergence.3 Second, the "11 to 17 million years" he reports as the revised split25 is, in the chimpanzee family study it matches, "an average time to the most common ancestor," the total divergence time rather than the split.28 A total divergence near 13 million years, with the split several million years younger, is what Part 2's arithmetic gives at that rate. Jeanson's second argument, that the fast family rate of mitochondrial DNA predicts far more human-chimp mitochondrial differences than the 1,483 observed, turns on why short-term and long-term mitochondrial rates differ. That question belongs to the Mitochondrial Eve article.
His own answer is creation: "A creationist would respond by saying that these differences were created."25 He sums up: "it appears that mtDNA differences are due to a constant rate of mutation over 6,000 years, whereas the majority — if not vast majority — of nuclear DNA differences are due to preexisting DNA differences that were placed in the ancestors of modern species 6,000 years ago."25
Robert Carter of Creation Ministries International takes the same view of the ape dates: "As far as timing the rise of the great ape lineages, all they are doing is uncovering the differences God built into their initial genomes. One of the basic premises of creation thought is that God designed hierarchically ... so some pairs of created groups are necessarily more similar than others and uncovering those differences does nothing to address the creation-evolution debate."11
Todd Wood, the young-earth biologist who has written most carefully about the chimpanzee genome, is more cautious. He accepts the consortium's figures, including its estimate that the fixed difference may be "as low as 1.06%."2 He floats, and does not endorse, the idea that God "created humans and chimpanzees with identical genomes," calling it "an intriguing possibility" that known biochemical differences "may well rule ... out."2 And he names the problem young-earth genetics has not solved: "Because many differences between the human and chimpanzee genomes may have been created rather than arising during post-creation history, we are left with the challenge of distinguishing created differences from differences that arose during the history of the two baramins."2 His closing judgment: "The genome revolution, exciting though it is, is not an obvious victory for creationism."2
John Sanford frames the differences as information the human line would have had to gain: "Chimp and human genomes are minimally 5 % divergent [1], representing about 150 million nucleotide differences."13 That figure counts every letter inside insertions and deletions as a separate difference. Wood made the correction in 2006 against an earlier creationist argument: "An insertion or deletion of 1000 nucleotides is only one mutational event, even though the total difference is 1000 nucleotides."2 The single-letter differences number about 35 million; the insertion and deletion events about 5 million.3
The price: more than 99.9 % created
The arithmetic leaves a young-earth model no middle ground. Take the measured human rate, about 35 new single-letter changes per copy of the genome per generation. Give the patriarchs of Genesis 5 and 11 the extra mutations that older fathers pass on today, about two more per child for each year of the father's age, run in a straight line up to Noah's 502 years. From Adam to Abraham that is 19 births in 1,948 years. Then add about 145 to 168 ordinary generations to the present.412
| Step | New mutations on one copy of the genome |
|---|---|
| Adam to Abraham, father's-age effect included | about 2,000 |
| Abraham to today, 25 to 29 years per generation | about 5,000 to 5,800 |
| Total per lineage | about 7,000 to 7,800 |
| Human and chimpanzee lineages together | about 14,000 to 15,700 |
| Share of the 35 million single-letter differences | 0.04 to 0.045 % |
The Septuagint's longer ages before Abraham raise the patriarchs' share from about 2,000 to about 3,400, and the total to about 8,400 to 9,200 per lineage, or 0.05 % of the differences. To make all the differences by mutation in this time, the rate would have to run about 1,900 to 2,500 times faster than it is measured in families today, in both species, on every line. That would be about 130,000 to 175,000 new mutations in every child, against about 60 to 70 today.
Two young-earth geneticists have floated a steeper effect of a father's age, and it does not change the answer. Robert Carter and Matthew Powell, extrapolating from a 2000 review of human mutation by the geneticist James Crow, wrote that "a man 500 years of age would donate approximately 10,000 mutations to a child." On Kong's straight line, they note, Noah "would only contribute slightly more than 1,000 mutations."29 Giving 10,000 new mutations to every one of the ten pre-Flood births, not to Noah alone, still accounts for only 0.6 % of the differences.12
So Wood's "identical genomes" possibility is ruled out by the numbers as well as by the biochemistry he mentions, and Jeanson and Carter are right that, on a young earth, almost all of the differences must have been there from the start. The young-earth claim to test is therefore not "mutation was faster." It is "more than 99.9 % of the single-letter differences between humans and chimpanzees were created." Part 5 tests it.
Part 5: Created differences or mutations? The fingerprints
Mutation is not random in where it strikes or what it does. It has habits, measured directly in families. If the human-chimp differences were made by mutation, they should share those habits. If they were created, nothing requires them to. Three habits can be checked against published numbers.
The CpG hotspot
Where a C is followed by a G in DNA, the C usually carries a methyl tag, and a methylated C decays to T far more often than any other letter changes. Kong's family study found transitions at CpG sites occurring at 18.2 times the rate at other sites, "ostensibly because spontaneous oxidative deamination of methylated cytosines leads to an increase in transition mutations."4
The human-chimp differences show the same hotspot. CpG sites "account for 25.2% of all substitutions while constituting only 2.1% of all aligned bases," a twelvefold excess and about 9 million of the 35 million differences.35 Two finer details match too. Changes at CpG sites behave more like a clock in years than other changes do, which Moorjani's team attributes to "their nonreplicative origin": they come from chemical decay over time rather than from copying errors.7 And the 2005 consortium found the male excess at CpG sites much smaller than elsewhere, about 2 against 3 to 6, which it read as "roughly equal rates of CpG to TpG transitions in the male and female germ line."3 Decay that runs with time should hit eggs and sperm alike; copying errors pile up in sperm, which go through far more cell divisions.
The sex chromosomes
Fathers pass on more new mutations than mothers. In Kong's families the ratio was 3.9 to 1;4 in a chimpanzee family studied by Oliver Venn and colleagues, "seven to eight times."28 This bias has a predictable effect on long-term change. The Y chromosome is always in a male, the X spends two thirds of its time in females, and the other chromosomes half. So with a male bias the Y should change fastest and the X slowest.
The human-chimp differences do exactly that: "a mean divergence of 1.9% for chromosome Y and 0.94% for chromosome X," against 1.23 % overall.3
| Y relative to other chromosomes | X relative to other chromosomes | |
|---|---|---|
| Observed human-chimp difference | 1.54 | 0.76 |
| Predicted from human families (male bias 3.2 to 3.9) | 1.52 to 1.59 | 0.80 to 0.83 |
| Predicted from the chimpanzee family (male bias 7.8) | 1.77 | 0.74 |
| Predicted with no male bias | 1.00 | 1.00 |
Both observed ratios fall inside the range the family-measured biases predict.30 The match is not perfect. The X is somewhat less different than the human bias alone predicts, which Scally's team put down to the smaller ancestral population of X chromosomes, and after correcting for it they estimated a male bias of 2.3.6
Regional variation
Mutation rates vary along the chromosomes, and so do the human-chimp differences. The 2005 consortium concluded that "the large-scale variation in the human–chimpanzee divergence rate primarily reflects regional variation in mutation rate."3 The direct test came later, by comparing where new mutations land in families with where the human-chimp differences are densest. Thomas Smith, Peter Arndt and Adam Eyre-Walker found the two correlated in all three large family datasets, "as expected," but more weakly than mutation alone predicts: 0.16 to 0.31, against 0.30 to 0.68 expected.31 They traced part of the gap to recombination, which also biases which variants become fixed. This check counts for mutation, but less strongly than the other two.
What these fingerprints do and do not show
They show that the human-chimp differences have the statistical character of mutations, in three independent respects, each measured in families without assuming any ancestry. They do not by themselves prove common descent. A model in which separately created kinds then diverged by mutation would also produce them, but Part 4 showed a young-earth timescale leaves room for only 0.05 % of the differences to arise that way.
So the young-earth model must say that God created the differences with the fingerprints of mutation already in them: a twelvefold CpG excess, a Y-to-X ratio matching the male bias of fathers' sperm, and the regional pattern of mutation rates. That is possible for a Creator. It is also exactly what Wood called "merely ad hoc,"2 and it gives up the one test that could separate created from mutated differences, the test Wood said young-earth genetics most needed.2 No young-earth source examined here addresses the CpG excess or the sex-chromosome ratio.27
What fits the created-differences view better
The created-differences view is not without support. Four findings sit awkwardly with a simple mutational clock.
- The yearly rate is not constant. As Part 2 showed, no single steady rate fits both the human-chimp and the human-orangutan fossils. The repair, a slowdown as apes reached puberty and fatherhood later, is reasonable and partly measured in living apes, but its timing and the ancestral ages it needs are inferred, not observed.
- The direct methods disagree. Repeat-segment counting gives a rate 50 % higher than whole-genome family counting.19
- The regional match is weaker than mutation alone predicts.31
- Single-letter changes are not the whole difference. Insertions, deletions and rearranged segments cover far more DNA than single-letter changes do, and the 2025 complete genomes found 12.5 to 27.3 % of each ape genome without a simple one-to-one match. Those differences are the subject of How Similar Are Human and Chimp DNA?. This article prices only the single-letter changes.
None of these four is evidence that the differences were created. Each is a place where the mutational account has a free parameter or an unexplained residue.
Part 6: The waiting-time problem on the human line
The questions so far concern neutral changes, which make up most of the human-chimp difference and need no selection. The waiting-time argument asks something else: how long a hominin population must wait for a specific useful change that needs several DNA letters at once.
The young-earth case, and the mainstream agreement
John Sanford and colleagues modified the simulation program Mendel's Accountant and "simulated a classic pre-human hominin population of at least 10,000 individuals, with a generation time of 20 years, and with very strong selection." Their result: "To establish a string of two nucleotides required on average 84 million years. To establish a string of five nucleotides required on average 2 billion years."13 Their full table gives 1.53 million years for one specified letter, 84.1 million for two, 2.31 billion for five and 18.5 billion for eight.13 Sanford concludes that "Routine establishment of specific beneficial strings of two or more nucleotides becomes very problematic."13
Mainstream mathematics agrees on the core. Rick Durrett and Deena Schmidt, writing in Genetics in 2008, worked out how long a human-sized population waits for one prespecified pair of mutations that switches off one gene-control site and creates another: "Multiplying by 25 years per generation gives 216 million years." The same paper notes that a new control site needing only one mutation appears "in an average of 60,000 years."14
The price
The human lineage has had 5.5 to 13 million years since the split, on the dates of Part 2. Against that window:
| Change needed | Sanford's waiting time | Fits in 6 million years? | Equally good targets needed to fit in 6 million | ... in 13 million |
|---|---|---|---|---|
| 1 specified letter | 1.53 million years | yes | 1 | 1 |
| 2 specified letters | 84 million years | no | 15 | 7 |
| 5 specified letters | 2.31 billion years | no | 385 | 178 |
| 8 specified letters | 18.5 billion years | no | 3,084 | 1,424 |
| Durrett-Schmidt prespecified pair | 216 million years | no | 36 | 17 |
The last two columns use a property both sides accept: if any of k different strings or places would do the job equally well, the expected wait falls by about k.15 Sanford says so himself. He lists the ways a real target could be easier than his, including that "the string should be able to arise anywhere," and concludes, "In all these special circumstances the waiting time will certainly be shorter than this study indicates."13 He also reports a published model in which a five-letter string allowed anywhere among 20,000 human gene promoters takes only about 7,000 years. He rejects it on biological grounds: "biologically it is simply not reasonable to assume that a specific short string can be, by itself, beneficial almost anywhere in the genome, with minimal contextual constraint."3213
What the argument shows
The arithmetic is not in dispute. A change that needs two or more particular letters at one particular place, with no benefit until all are present, is very unlikely on the human line in 6 or even 13 million years. A single specified letter is not limited: Sanford's own model gives it 1.5 million years.
What is not known is how many human-specific traits needed changes of the hard kind. If most needed one of hundreds of possible strings, or could be built one useful letter at a time, the waiting time vanishes. If even a few needed one exact multi-letter string with useless intermediates, it bites. No published study on either side has gone through the human-specific changes and counted. Until someone does, the waiting-time argument establishes a constraint, not a refutation.
The separate question of how many beneficial changes selection could fix in the time, Haldane's dilemma, and the full technical dispute over Sanford's simulations are in Haldane's Dilemma and the Waiting-Time Problem.
Objections and responses
"Molecular clocks assume common ancestry, so any date they give is circular." Partly true, and the mainstream says so about the old rate: Langergraber's team called fossil calibration "an inherent circularity."1 The family rate is different. It is counted in living parents and children and assumes nothing about ancestry. Turning it into a split date does assume the two lineages shared an ancestor, so a date cannot by itself prove shared ancestry. That is why this article tests the pattern of the differences in Parts 3 and 5, not only their number.
"The rates disagree by a factor of two or more, so the clock is worthless." The spread among published human-chimp splits is about three-and-a-half-fold, from 3.75 to 13 million years. The gap between any of them and a young earth is about a thousandfold. A clock that cannot tell 4 million years from 13 can still tell either from 7,500.
"Rates measured in families run far faster than fossil-calibrated rates, so the long ages are wrong." Jeffrey Tomkins and Jerry Bergman, writing in the Journal of Creation in 2015, call counting in families "the most empirical way to measure genetic change" and say these counts "greatly exceed the more speculative and spurious rates achieved from paleontology."23 That is true of the fast-changing control region of mitochondrial DNA, as the Mitochondrial Eve article shows. For the nuclear DNA that holds the 35 million differences, it runs the other way. The family rate is about half the fossil rate,18 so the most empirical rate makes the human-chimp split older, not younger. Tomkins and Bergman also argue that "pervasive genome-wide biochemical functionality" undercuts the neutral theory on which clocks rest.23 If most of the differing letters did have functions, the dates would grow longer still. Changes with no effect pile up at the rate mutations arise.21 Changes that damage a function are mostly weeded out, so working DNA collects differences more slowly than mutation supplies them, and more time is needed to reach the same count.
"The evolutionists simply moved the date to fit the new rate." Jeanson is right that the date was "revised in light of the mutation rates."25 Revising a date when a better measurement arrives is how measurement works. The real cost is elsewhere: the revision broke the orangutan calibration, and the repair, a slowdown in the yearly rate, is a free parameter that has been measured only in living apes.910
"Young-earth geneticists trust family-measured rates for Mitochondrial Eve. Consistency demands the same rate here." They do, and the consistency holds. The same family rate that young-earth writers use for human mitochondrial DNA gives at most about 0.05 % of the human-chimp differences in 6,000 to 7,500 years. Jeanson and Carter accept this, which is why they say the differences were created. The tension is not inconsistency. It is that the created differences look like mutations.
"God would naturally use the same chemistry and the same design rules for both species, so created differences would look like mutations." Shared chemistry explains shared genes. It does not explain why the differences pile up at the one site that methyl-C decay attacks, or why the Y differs twice as much as the X in the ratio set by fathers' sperm. Those are signatures of a process acting over time, not of a design rule. A Creator could have reproduced them, but the explanation then predicts nothing that was not already observed, which is Wood's objection to ad hoc design.2
"Incomplete lineage sorting shows the tree of life is false." ILS needs no transfer of DNA between species after they parted; Tomkins and Bergman assume it does.26 It needs only that variants present before the split were kept unevenly. It predicts an even split between the two odd trees and a drop near genes and on the X, and the genomes show both.6
"Humans and chimpanzees differ by 5 %, 150 million letters, far too many for mutation." The 5 % counts every letter inside an insertion or deletion as a separate change. Wood corrected the same mistake in 2006.2 The single-letter differences number about 35 million, the insertion and deletion events about 5 million.3 On the mainstream model, the single-letter count is what the measured rate predicts once the ancestral population is counted, as Part 2 shows.
"The waiting-time problem proves humans could not have evolved from apes." It proves that changes needing a prespecified multi-letter string at one place could not arise in the time. Whether human evolution needed many such changes is not established by Sanford, who concedes the waits shorten whenever the string could arise in more places or more forms.13
From the skeptic's side: "The young-earth model makes no testable prediction here at all." It can. Wood posed the test: distinguish created differences from those that "arose during the history of the two baramins."2 A created-differences model that predicted, in advance, a spectrum of differences unlike mutation's, and then found it, would be strong evidence. The spectrum found so far is mutation's.
Issues and unanswered questions
- Besenbacher's faster ape rates rest on ten families: seven chimpanzee, two gorilla and one orangutan. The orangutan interval, 0.98 to 1.85 times the human rate, includes no difference at all.10 Method matters too. When five laboratories analysed the same family of rhesus macaques in 2022, their rates ran from 0.46 to 0.85 × 10-8 per letter per generation, "almost a twofold variation" from processing choices alone.33 Besenbacher's team guarded against this by running the apes through "essentially the same" pipeline it had used for humans, and by checking that about half of the new mutations found in a young ape passed on to its own offspring, as real mutations should.10 The slowdown that rescues the orangutan calibration is measured, but thinly.
- The two direct methods still disagree. Repeat-segment counting gives 1.82 × 10-8 per generation and whole-genome counting about 1.2 × 10-8. Sun's team suggested the genome counts miss real mutations;19 the genome counts have since grown much larger without rising, and the 2025 four-generation family, read into the repeats and centromeres, still gave 1.17 × 10-8.20 No paper found here reconciles them.
- The transition-to-transversion ratio of human-chimp differences was not found stated in any source read, so it could not be compared with the 67.8 % transitions seen in families.4
- The regional correlation is half of what mutation alone predicts. Smith and colleagues attribute the rest partly to recombination and biased gene conversion.31 A young-earth reader could press this point harder than the mainstream does.
- No created-kinds model of ILS exists. A model that produced the 30 to 40 % mosaic, its even split, and its drop near genes and on the X from designed starting genomes would be a real contribution. None was found.
- Nobody has counted the hard changes. The waiting-time dispute turns on how many human-specific traits needed one exact multi-letter string. Neither side has done the count.
- Jeanson's "11 to 17 million years" is cited in his book to "the authors of the chimpanzee genome paper." The figure matches Venn and colleagues' 2014 chimpanzee family study, which calls it an average time to the common ancestor. The 2005 chimpanzee genome paper does not contain it.2825
Verdict
The family-measured mutation rate is the best-measured number in this debate, and it is about half the old fossil-calibrated rate. With it, the human-chimp split moves from about 4 million years to between 5.5 and 13 million, depending on the generation length and the ancestral population assumed. The revision fixes one fossil conflict and creates another, which mainstream geneticists repair with a slowdown in the yearly rate that is plausible and partly measured. That is a real weakness, and young-earth critics are right to press it. It moves the dates by a factor of about three, never toward thousands of years.
A young-earth model cannot get the differences from mutation. At measured rates, even allowing for the patriarchs' great ages, mutation since creation makes 0.04 to 0.05 % of them. Jeanson and Carter accept this and say the differences were created. That claim is coherent, but it meets three findings that it does not predict: the differences sit at mutation's CpG hotspot, the sex chromosomes differ in the ratio set by fathers' male-biased mutation, and a third of the genome shows the gorilla mosaic that a large shared ancestral population leaves behind, in the proportions and places that theory expects. The mutational reading predicts all three. The created reading can only accommodate them.
The waiting-time argument is the young-earth side's strongest point. It shows that changes needing exact multi-letter strings are very unlikely on the human line. It has not shown that the human line needed many such changes.
On the evidence available, the human-chimp differences are better explained as mutations accumulated over millions of years than as created differences. The young-earth case here rests on Wood's unmet challenge, a way to tell created differences from mutated ones, and on a count of waiting-time targets that nobody has made.
Sources
- 1.
Langergraber, Kevin E. et al. “Generation times in wild chimpanzees and gorillas suggest earlier divergence times in great ape and human evolution.” Proceedings of the National Academy of Sciences 109(39):15716–15721. Sep 25, 2012. A field and genetics study from the Max Planck Institute for Evolutionary Anthropology, with Svante Pääbo and Linda Vigilant as senior authors. It measured generation lengths from parentage records of 226 wild chimpanzee and 105 mountain gorilla offspring, combined them with family-measured human mutation rates, and recalculated published ape split dates without fossil calibration, putting the human-chimp split at 7 to 13 million years. The article uses it for the origin of the old 1 × 10-9 rate, the generation lengths and the recalculated dates.
- 1a.
Page 15716: “Recently, researchers have commonly used a mutation rate of 1 × 10−9 mutations per site per year (e.g., refs. 4, 6, 8, 15) derived from the observed DNA sequence difference of around 1.3% between the human and chimpanzee genomes (8, 15, 16) and an assumed DNA sequence divergence between these species at 7 Ma, as well as from an observed sequence difference of 6.46% between the human and macaque genomes (17) and an assumption of their DNA sequence divergence at 25 Ma.” “Although ubiquitous, this approach has an inherent circularity and is subject to possible error because it relies on the accuracy of the ages of fossils.” ↩ ↩ ↩ ↩
- 1b.
Page 15716: “Sarich and Wilson used immunological data to propose that humans and African great apes diverged only about 5 million y ago, some three to four times more recently than had been assumed on the basis of the fossil record (1).” “this divergence has since been repeatedly estimated from DNA sequence data at 4–6 million years ago (Ma) (3–8).” ↩
- 1c.
Page 15716: “Genetic studies of humans have commonly used a generation time of 20 or 25 y”. ↩
- 1d.
Page 15716: “The total amount of sequence differences observed today between two evolutionary lineages can be expressed as the sum of two values: the sequence differences that accumulated since gene flow ceased between the lineages (“split time”) and the sequence differences that correspond to the diversity in the common ancestor of both lineages.” ↩
- 1e.
Page 15717, table 1: Average age of parents at birth: chimpanzees, 24.63 years from 226 offspring in eight communities; mountain gorillas, 19.28 years from 105 offspring; humans, taken from Fenner 2005, 28.6 years for hunter-gatherers and 29.1 for countries. ↩
- 1f.
Page 15718: “DNA sequencing of human families has recently yielded four direct estimates of mutation rates ranging from 0.97 × 10−8 to 1.36 × 10−8/site/generation (19–21). When considering the average present-day human generation time of 29 y, this results in rates ranging from 0.33 to 0.47 × 10−9/site/year.” ↩ ↩
- 1g.
Page 15718: “We estimate the split time between the human and chimpanzee lineages at between 7 and 13 million years, whereas previous estimates range from 4 to 6 million years.” ↩ ↩
- 1h.
Page 15718, table 2: Human-chimp splits first published at 1.0 × 10-9 per year and recalculated: Scally and colleagues 2012, 3.69 to 6.78–11.03 million years; Hobolth and colleagues 2011, 4.22 to 7.76–12.62; Dutheil and colleagues 2009, 4.38 to 8.05–13.09; Prüfer and colleagues 2012, 4.5 to 8.27–13.45. ↩
- 1a.
- 2.
Wood, Todd Charles. “The chimpanzee genome and the problem of biological similarity.” Occasional Papers of the Baraminology Study Group 7:1–18. Feb 20, 2006. A review by a young-earth creationist biologist, then at Bryan College's Center for Origins Research, of what the 2005 chimpanzee genome means for creationism. It accepts the measured differences, rejects several popular creationist arguments against them, and concludes that creation biology needs an explanation of the pattern of similarity rather than an appeal to a common designer. The article uses it as the most careful young-earth treatment of the question.
- 2a.
Page 7: “The question is not how similarity arose but why this particular pattern of similarity arose. To say that God could have created the pattern is merely ad hoc.” ↩ ↩ ↩ ↩
- 2b.
Page 4: “the fixed differences between the two genomes may be as low as 1.06%”. ↩
- 2c.
Page 10: “An insertion or deletion of 1000 nucleotides is only one mutational event, even though the total difference is 1000 nucleotides.” ↩ ↩
- 2d.
Page 12: “If the fixed nucleotide mismatches between the chimpanzee and human genomes are 1.06%, then the original nucleotide identity could be as high as 99%. At that high level of similarity, perhaps it is not impossible to believe that God created humans and chimpanzees with identical genomes. The known differences between human and chimpanzee biochemistry (see Varki 2000; Varki and Atheide 2005) may well rule this out, but it is an intriguing possibility.” ↩
- 2e.
Page 13: “Because many differences between the human and chimpanzee genomes may have been created rather than arising during post-creation history, we are left with the challenge of distinguishing created differences from differences that arose during the history of the two baramins.” ↩ ↩ ↩
- 2f.
Page 14: “The genome revolution, exciting though it is, is not an obvious victory for creationism.” ↩
- 2a.
- 3.
Chimpanzee Sequencing and Analysis Consortium. “Initial sequence of the chimpanzee genome and comparison with the human genome.” Nature 437:69–87. Sep 1, 2005. The first chimpanzee genome, from one male chimpanzee, compared letter by letter with the human genome. It counted about 35 million single-letter differences and 5 million insertions and deletions, and it described how the differences vary along the chromosomes, by chromosome and by type of site. The article uses its counts, its sex-chromosome and CpG figures, and its account of what the divergence time contains.
- 3a.
Page 69: “approximately thirty-five million single-nucleotide changes, five million insertion/deletion events, and various chromosomal rearrangements”. “Single-nucleotide substitutions occur at a mean rate of 1.23% between copies of the human and chimpanzee genome, with 1.06% or less corresponding to fixed divergence between the species.” ↩ ↩ ↩ ↩
- 3b.
Page 71: The paper divides the divergence time of each stretch of DNA into two parts: the time since speciation, which it gives as about 6 to 7 million years, and the coalescence time within the common ancestral population, which it says "may be on the order of 1–2 million years". ↩ ↩
- 3c.
Page 71: “The most striking outliers are the sex chromosomes, with a mean divergence of 1.9% for chromosome Y and 0.94% for chromosome X. The likely explanation is a higher mutation rate in the male compared with female germ line”. “is consistent with roughly equal rates of CpG to TpG transitions in the male and female germ line”. The paper's estimates of the male-to-female ratio are 3 to 6 from the sex chromosomes, and about 2 from CpG sites alone. ↩ ↩ ↩
- 3d.
Page 71: “we suggest that the large-scale variation in the human–chimpanzee divergence rate primarily reflects regional variation in mutation rate.” ↩
- 3e.
Page 72: “Sites containing CpG dinucleotides in either species show a substantially elevated divergence rate of 15.2% per base; they account for 25.2% of all substitutions while constituting only 2.1% of all aligned bases.” ↩ ↩
- 3a.
- 4.
Kong, Augustine et al. “Rate of de novo mutations and the importance of father's age to disease risk.” Nature 488:471–475. Aug 23, 2012. A whole-genome sequencing study of 78 Icelandic parent-child trios by deCODE genetics. It found 4,933 new mutations, a rate of 1.20 × 10-8 per letter per generation, about two more per year of the father's age, and almost four times as many from fathers as from mothers. The article uses it for the family rate and for the pattern of new mutations: their CpG hotspot, their transition share and their paternal bias.
- 4a.
Section Abstract: “Here we show that in our samples, with an average father’s age of 29.7, the average de novo mutation rate is 1.20×10−8 per nucleotide per generation.” Section Mutations by type and by chromosome: “From that, 4933 mutations correspond to a germline mutation rate of 1.20×10−8 per”. ↩ ↩
- 4b.
Section Parent of origin and father’s age: “The number of mutations increases with father’s age (P = 3.6×10−19) with an estimated effect of 2.01 mutations per year”. ↩ ↩ ↩
- 4c.
Section Parent of origin and father’s age: “In the five trios, the average number of paternal and maternal mutations is 55.4 and 14.2 respectively”. “the mean number of paternal mutations is substantially higher than the mean number of maternal mutations (ratio = 3.9)”. ↩ ↩ ↩
- 4d.
Section Mutations by type and by chromosome: “Approximately two-thirds (3344/4933 = 67.8%) are transitions. Moreover, there is clear difference between mutation rates at CpG and non-CpG sites. CpG dinucleotides are known to be mutational hotspots in mammals, ostensibly because spontaneous oxidative deamination of methylated cytosines leads to an increase in transition mutations”. “The observed rate of transitions here is 18.2 times that at non-CpG sites”. ↩ ↩ ↩
- 4a.
- 5.
This site's arithmetic. Divergence time is T = d / 2μ, where d is the share of letters that differ and μ the yearly rate per letter; the 2 is there because both lineages mutate. Split time is T minus 2Ng, the average time two gene copies take to find a common ancestor in an ancestral population of N breeding individuals with generations of g years. With d = 1.23 %, a rate of 1 × 10-9 gives T = 6.15 million years; 1.20 × 10-8 per generation over 25, 26.9 or 29 years gives 0.48, 0.45 or 0.41 × 10-9 per year and T = 12.8, 13.8 or 14.9 million years. At 1 × 10-9 a 6-million-year split leaves room for N of only about 3,000, and 7 million is impossible; at 0.41 to 0.51 × 10-9 a 6 to 7 million year split needs N of about 100,000 to 180,000. The 1 × 10-9 rate itself is 1.3 % divided by twice 7 million years, 0.93 × 10-9. On Yoo and colleagues' own inputs (1.25 × 10-8 per generation, 24.5-year generations, N = 198,000, splits at 5.5-6.3 and 10.6-10.9 million years), 61 to 64 % of today's divergence predates the split, and the same inputs predict 35.6 to 40.5 % discordant gene trees against the 39.5 % reported; with the family rate and N of 50,000 to 100,000 the share predating the split is 19 to 36 %. CpG sites carry 25.2 % of the differences on 2.1 % of the letters, a twelvefold excess, about 8.8 million of 35 million. The calculations are in the file research/split-arithmetic.ts beside this article, which first reproduces Kong's rate from his counts and Langergraber's recalibrated dates of 6.78 and 11.03 million years. ↩ ↩ ↩ ↩ ↩ ↩ ↩ ↩
- 6.
Scally, Aylwin et al. “Insights into hominid evolution from the gorilla genome sequence.” Nature 483:169–175. Mar 8, 2012. The first gorilla genome, from the Wellcome Trust Sanger Institute and partners, compared with the human, chimpanzee and orangutan genomes using a model, CoalHMM, that infers the family tree at each position along the DNA. It found that 30 % of the genome does not follow the species tree, and it showed how the human-chimp split date depends on which mutation rate is assumed. The article uses it for incomplete lineage sorting and for the dates.
- 6a.
Section Summary: “In 30% of the genome, gorilla is closer to human or chimpanzee than the latter are to each other; this is rarer around coding genes, indicating pervasive selection throughout great ape evolution, and has functional consequences in gene expression.” ↩ ↩
- 6b.
Section Speciation of the great apes: “CoalHMM’s results would correspond to speciation time estimates THC and THCG of 3.7 and 5.95 Mya respectively (Fig. 1b).” “indicate that a lower value of 0.5-0.6 × 10−9 bp−1y−1 is plausible (based on average hominine generation times of 20 to 25 y). This would give substantially older estimates of approximately 6 and 10 Mya for THC and THCG, potentially in better agreement with the fossil record.” The first figure uses the old rate of 10-9 per year. ↩ ↩ ↩
- 6c.
Section Speciation of the great apes: “This is possible if we allow for mutation rates changing over time, with a mutation rate of around 1 × 10−9 bp−1y−1 in the common ancestor of great apes, decreasing to lower values in all extant species (Fig. 1b).” “A decrease within the great apes is also a predicted consequence of the observed increase in body sizes over this time period and the association of small size with shorter generation times in other primates”. “However we note that Sahelanthropus and Chororapithecus remain difficult to incorporate in this model, and can be accommodated as hominin and gorillin genera only if most of the decrease occurred early in great ape evolution.” ↩ ↩
- 6d.
Section Speciation of the great apes: “In summary, although whole genome comparisons can be strongly conclusive about the ordering of speciation events, the inability to observe past mutation rates means that the timing of events from genetic data remains uncertain.” ↩ ↩
- 6e.
Section Incomplete lineage sorting and selection: “Across the genome we find 30% of bases exhibiting ILS, with no significant difference between the number sorting as ((H,G),C) and ((C,G),H).” “Within coding exons mean ILS drops to 22%, and the suppression of ILS extends out to several hundred kbp from coding genes, evident even in raw site patterns before any model inference (Fig. 2b).” ↩ ↩ ↩ ↩
- 6f.
Section Incomplete lineage sorting and selection: “As expected, we see reduced ILS (Fig. 2a) and HC sequence divergence dHC (Fig. SF6.1) on the X chromosome, corresponding to a difference in Ne between X and the autosomes within the ancestral HC population.” “Incorporating the ancestral Ne estimates from CoalHMM, we estimate a ratio of 0.87 ± 0.09 between average mutation rates on X and the autosomes on the HC lineage, corresponding to a male/female mutation rate bias α = 2.3 ± 0.4”. ↩ ↩ ↩
- 6a.
- 7.
Moorjani, Priya et al. “Variation in the molecular clock of primates.” Proceedings of the National Academy of Sciences 113(38):10607–10612. Sep 20, 2016. A comparison of whole genomes from 10 primate species that measured how fast different kinds of DNA change ran along each lineage. It found small differences among the apes and that changes at CpG sites keep the steadiest time, and it used them alone to date the human-chimp split at about 7.9 million years. The article uses its dates, its statement of the twofold rate gap and its CpG finding.
- 7a.
Page 10607: “is approximately twofold lower than the mutation rate inferred from the number of substitutions observed between primates”. ↩
- 7b.
Page 10607: “Within apes, rates are ∼2% higher in chimpanzees and ∼7% higher in the gorilla than in humans.” ↩
- 7c.
Page 10607: “transitions at CpG sites exhibit a more clocklike behavior than do other types, presumably because of their nonreplicative origin.” ↩
- 7d.
Page 10611: “we estimate that humans diverged from chimpanzees ∼12.1 Mya and from gorillas ∼15.1 Mya (Fig. 4).” Pages 10611–10612: “the human–chimpanzee split time is ∼7.9 Mya”. “Accounting for this uncertainty provides estimates of human–chimpanzee split time in the range of 6.5–9.3 Mya”. “We note that there is substantial uncertainty in estimates of ancestral population size of apes, with previous estimates ranging between 50,000 and 100,000 (43–45).” ↩ ↩ ↩
- 7a.
- 8.
Yoo, DongAhn et al. “Complete sequencing of ape genomes.” Nature 641:401–418. May 8, 2025. A large consortium paper that sequenced the chromosomes of six ape species end to end with no gaps. It dated the ape splits with a model that also estimates incomplete lineage sorting and ancestral population sizes, using a mutation rate of 1.25 × 10-8 per generation. The article uses its split dates, its ILS shares and its ancestral population sizes.
- 8a.
Page 403: “Our analyses dated the human–chimpanzee split between 5.5 and 6.3 million years ago (Ma; minimum to maximum estimate of divergence), the African ape split at 10.6–10.9 Ma and the orangutan split at 18.2–19.6 Ma (Fig. 2a). We inferred ILS for an average of 39.5% of the autosomal genome and 24% of the X chromosome.” “we estimated that the human–chimpanzee–bonobo ancestral population size (average Ne = 198,000) is larger than that of the human–chimpanzee–gorilla ancestor (Ne = 132,000)”. ↩ ↩ ↩ ↩ ↩ ↩ ↩
- 8b.
Section Supplementary Information, Note VI: “In our analysis we considered any genomic region in ILS status when the reconstructed phylogeny as different from ((A,B)),C,D).” “For all the analyses we used a mutation rate of 1.25×10-8 and the average generation time across the species for specific node”. The stray parenthesis is the supplement's. The generation times listed for the human-chimpanzee-bonobo and human-chimpanzee-gorilla nodes are 24.5 and 21.75 years. ↩
- 8a.
- 9.
Sun, James X. et al. “Supplementary Note for A direct characterization of human mutation based on microsatellites.” Nature Genetics 44. 2012. The 70-page supplement to the same study. Its chapter 10 reruns the dating with Kong and colleagues' whole-genome family rate in place of the repeat-segment rate and sets the results against the fossil record. The article uses it for the human-orangutan conflict and for the authors' view of the slowdown hypothesis.
- 9a.
Page 66: “the inferred date of human-chimpanzee speciation is 8.32-11.8 Mya”. “The inferred human-orangutan genetic divergence date of 34.0-46.2 Mya is so much older than the upper bound from the fossil record of <18 Mya on human-orangutan speciation that the date is implausible”. ↩ ↩ ↩
- 9b.
Page 66: “However, this scenario also requires us to hypothesize a combination of unlikely events: (a) the slowdown would need to have been coincidental in both lineages to explain the observations, and (b) the slowdown would also have to have been extraordinarily dramatic: about 3-fold in both lineages in the period ancestral to human-chimpanzee divergence to produce as extreme an effect as is observed.” ↩ ↩
- 9a.
- 10.
Besenbacher, Søren et al. “Direct estimation of mutations in great apes reconciles phylogenetic dating.” Nature Ecology & Evolution 3:286–292. 2019. A family study of great apes from Aarhus University and partners: ten parent-offspring trios, seven chimpanzee, two gorilla and one orangutan, with 397 new mutations. Extending the human parental-age model to these apes, it found their yearly rates about 1.5 times the human rate and used them to date the human-chimp split at 6.6 million years. The article uses its rates and dates, and notes how few families they rest on.
- 10a.
Section Abstract: “The yearly trio-based mutation rate estimate of around 0.43 × 10−9 is markedly lower than previous indirect estimates of about 1 × 10−9 per year from phylogenetic comparisons of the great apes calibrated by fossil evidence.” “and by a factor of 1.48 ± 0.08 for the three species combined. These estimates suggest an appreciable slowdown in the yearly mutation rate in the human lineage that is likely to be recent as genome comparisons almost adhere to a molecular clock. If the nonhuman rates rather than the human rate are extrapolated over the phylogeny of the great apes, we estimate divergence and speciation times that are much more in line with the fossil record and the biogeography.” ↩ ↩ ↩ ↩
- 10b.
Section Discussion: “The human-chimpanzee separation time of 6.6 million years (95% confidence interval, 5.9–7.3 million years) that we estimated is consistent with Ardipithecus”. “A species separation time of 15.9 million years ago (95% confidence interval, 14.2–17.6 million years ago) with orangutan is well within the time interval between the age of Sivapithecus (12.2 million years old) and Proconsul (23 million years old)”. ↩
- 10c.
Figure 3: Yearly rates relative to humans, with 95 % intervals: chimpanzee 1.50 (1.29–1.70), gorilla 1.51 (1.19–1.84), orangutan 1.42 (0.98–1.85). Table 1 lists the ten families: seven chimpanzee, two gorilla, one orangutan. ↩
- 10d.
Section Results: “Our mutation rate pipeline is essentially the same as the one we previously used in humans, and which we showed has a very low false-positive rate”. “We also found that the number of transmitted mutations to the grandchild is not significantly different from 50%”. ↩
- 10a.
- 11.
Carter, Robert W. “Genealogical vs phylogenetic mutation rates: answering a challenge.” Proceedings of the Ninth International Conference on Creationism, 168–180. 2023. A conference paper by a young-earth geneticist at Creation Ministries International, defending the use of fast family-measured mitochondrial rates for a young human timescale and modelling mutation accumulation with Mendel's Accountant. In passing it states how a young-earth model reads the great-ape dates. The article uses that statement.
- 11a.
Page 172: “As far as timing the rise of the great ape lineages, all they are doing is uncovering the differences God built into their initial genomes. One of the basic premises of creation thought is that God designed hierarchically (Cserhati and Carter 2020), so some pairs of created groups are necessarily more similar than others and uncovering those differences does nothing to address the creation-evolution debate.” ↩ ↩
- 11a.
- 12.
This site's arithmetic. New mutations per child follow Kong and colleagues' straight-line model, 63.2 at a father's age of 29.7 plus 2.01 for each further year;4 any one copy of the genome carries half of them. Fathers' ages at the son's birth are those of the Masoretic text: 130, 105, 90, 70, 65, 162, 65, 187, 182 and 502 in Genesis 5 (Noah at Shem's birth, from Genesis 11:10), then 100, 35, 30, 34, 30, 32, 30, 29 and 70 in Genesis 11, for 1,948 years from Adam to Abraham. That gives about 2,000 new mutations per lineage. From Abraham to today is about 4,200 years: 145 generations at 29 years or 168 at 25, each adding 1.20 × 10-8 × 2.9 billion letters, about 35. Totals: 7,000 to 7,800 per lineage, 14,000 to 15,700 for the human and chimpanzee lineages together, 0.040 to 0.045 % of 35 million. The Septuagint's ages, the longest biblical chronology, are 230, 205, 190, 170, 165, 162, 165, 167, 188 and 502 in Genesis 5, then 100, 135, 130 (the second Cainan), 130, 134, 130, 132, 130, 79 and 70, for 3,314 years and 20 births to Abraham. They give about 3,400 patriarchal mutations, 8,400 to 9,200 per lineage, 0.048 to 0.053 % of the differences, and a needed speed-up of 1,900 to 2,100 times, against 2,200 to 2,500 on the Masoretic ages. Today's rate on both copies of 2.9 billion letters is about 70 new mutations per child, so a speed-up of 1,900 to 2,500 times means about 130,000 to 175,000. The straight-line extension to fathers aged several centuries is generous to the young-earth side; 76 of Kong's 78 fathers were between 18 and 40.5. The Carter-Powell case gives each of the ten births of Genesis 5 the full 10,000 new mutations they estimate for a 500-year-old father, all of them on the one lineage traced, with no halving, and the later births their ordinary share: about 106,000 per lineage, 0.6 % of the differences. ↩ ↩ ↩
- 13.
Sanford, John et al. “The waiting time problem in a model hominin population.” Theoretical Biology and Medical Modelling 12:18. 2015. A computer simulation of how long a hominin population needs to establish a short, specific string of DNA letters. It modified Mendel's Accountant so that a starting string and a target string could be set, simulated a population of at least 10,000 with 20-year generations and very strong selection, and found that a two-letter string takes on average 84 million years and a five-letter string 2 billion years. Sanford, a former Cornell plant geneticist, is the leading young-earth writer on genetics. The article uses its results and its own statement of when the waits shorten.
- 13a.
Page 1: “We simulated a classic pre-human hominin population of at least 10,000 individuals, with a generation time of 20 years, and with very strong selection (50 % selective elimination).” “To establish a string of two nucleotides required on average 84 million years. To establish a string of five nucleotides required on average 2 billion years.” “Routine establishment of specific beneficial strings of two or more nucleotides becomes very problematic.” ↩ ↩ ↩
- 13b.
Page 2: “Chimp and human genomes are minimally 5 % divergent [1], representing about 150 million nucleotide differences.” ↩
- 13c.
Pages 11–12, table 2: Corrected waiting times to establish a specified string: 1 nucleotide 1.53 × 106 years, 2 nucleotides 8.41 × 107, 5 nucleotides 2.31 × 109, 8 nucleotides 1.85 × 1010. ↩
- 13d.
Pages 4–5: “Some may say the string should be able to arise anywhere (genetic context does not matter).” “In all these special circumstances the waiting time will certainly be shorter than this study indicates.” ↩ ↩
- 13e.
Page 25: “works mathematically, and can yield very short waiting times (about 7,000 years).” “biologically it is simply not reasonable to assume that a specific short string can be, by itself, beneficial almost anywhere in the genome, with minimal contextual constraint.” Sanford is describing Behrens and Vingron's model, in which the string may arise in any of 20,000 promoters. ↩
- 13a.
- 14.
Durrett, Rick, and Deena Schmidt. “Waiting for two mutations: with applications to regulatory sequence evolution and the limits of Darwinian evolution.” Genetics 180(3):1501–1509. 2008. A mathematical paper from Cornell on how long a population waits for two specific mutations when the first gives no benefit, applied to gene-control sites in fruit flies and humans and set against Michael Behe's arguments. The article uses its human result as the mainstream counterpart to Sanford's.
- 14a.
Page 1501: “for humans with a much smaller effective population size, this type of change would take >100 million years.” Page 1507: “Multiplying by 25 years per generation gives 216 million years.” “Our previous work has shown that, in humans, a new transcription factor binding site can be created by a single mutation in an average of 60,000 years”. ↩ ↩
- 14a.
- 15.
This site's arithmetic. If k different strings, or k different places, would each serve equally well, a usable string appears about k times as often, so the expected wait falls by about k. The table divides each waiting time by the time available and rounds up, since the wait divided by k must fit inside the window: 84.1 million ÷ 6 million = 14.02, so 15; 2.31 billion ÷ 6 million = 385; 18.5 billion ÷ 6 million = 3,083.3, so 3,084; 216 million ÷ 6 million = 36; and the same by 13 million, which gives 7, 178, 1,424 and 17. It ignores the time to spread the change through the population once it appears, which in Sanford's runs is about 15,000 to 17,000 years and makes no difference at this scale. ↩ ↩
- 16.
Jónsson, Hákon et al. “Parental influence on human germline de novo mutations in 1,548 trios from Iceland.” Nature 549:519–522. Sep 28, 2017. The largest family study of new mutations at the time, by deCODE genetics: 1,548 Icelandic trios sequenced at 35-fold coverage, 108,778 new mutations, 42,961 of them traced to the parent of origin. Mothers' counts rose by 0.37 per year of age and fathers' by 1.51. The article uses its rate per generation and per year.
- 17.
Wang, Richard J. et al. “Human generation times across the past 250,000 years.” Science Advances 9(1):eabm7047. 2023. A genetic study that estimated past human generation lengths from the kinds of mutations found in variants of different ages, using the way fathers' and mothers' ages shift the mix of new mutations. The article uses its long-run average of 26.9 years.
- 17a.
Page 1: “Our analyses of whole-genome data reveal an average generation time of 26.9 years across the past 250,000 years, with fathers consistently older (30.7 years) than mothers (23.2 years).” ↩
- 17a.
- 18.
Scally, Aylwin, and Richard Durbin. “Revising the human mutation rate: implications for understanding human evolution.” Nature Reviews Genetics 13:745–753. Oct 2012. A review by two Sanger Institute geneticists of what the newly measured family mutation rate, about half the old fossil-calibrated value, does to the dates of human evolution, with a proposed revised rate of 0.5 × 10-9 per year. The article uses its statement of the gap and its argument that generation length alone cannot close it.
- 18a.
Page 745: “These measurements reveal a value that is approximately half of that previously derived from fossil calibration, and this has implications for our understanding of demographic events in human evolution and other aspects of population genetics.” ↩ ↩
- 18b.
Page 745: “A much lower value of 10–15 years, which is needed for the de novo and phylogenetic rates to … match, would be shorter than any other living great ape and less than half of the value measured in present-day humans.” ↩
- 18a.
- 19.
Sun, James X. et al. “A direct characterization of human mutation based on microsatellites.” Nature Genetics 44(10):1161–1165. 2012. A study of 2,058 new mutations at short repeated DNA segments in 85,289 Icelanders, from deCODE genetics and Harvard. From these it inferred a single-letter rate of 1.82 × 10-8 per generation, higher than whole-genome family counts, and a human-chimp split of 3.75 to 6.57 million years. The article uses it as an independent direct method that disagrees with the family counts.
- 19a.
Table 2: Nucleotide rate 1.82 × 10-8 per site per generation (1.40–2.28) or 6.76 × 10-10 per year; human-chimpanzee divergence 7.49 million years (5.80–9.77); human-chimpanzee speciation 4.97 million years (3.75–6.57). Section Results: “We hypothesize that the lower mutation rate estimates from the WGS studies might be due to: (i) the limited number of mutations detected in the WGS studies which explains why their confidence intervals overlap ours, (ii) possible underestimation of the false-negative rate in the WGS studies”. ↩ ↩ ↩ ↩
- 19b.
Section Results: “Our upper bound of τHC < 6.57 Mya is lower than the 6.8–7.2 Mya”. “date estimate for Sahelanthropus tchadensis, a fossil interpreted as being on the human lineage after the final separation of human and chimpanzee ancestors”. ↩
- 19a.
- 20.
Porubsky, David et al. “Human de novo mutation rates from a four-generation pedigree reference.” Nature 643(8071):427–436. 2025. A family study that read a four-generation, 28-member family (CEPH 1463) with five short-read and long-read sequencing technologies, assembling more than 95 % of each genome, including centromeres and repeats earlier studies could not read. Its rate of new single-letter mutations matches the older family studies. It matters here because it tests the suggestion that whole-genome counts run low by missing mutations in hard-to-read DNA.
- 20a.
Page 427, abstract: “Here using five complementary short-read and long-read sequencing technologies, we phased and assembled more than 95% of each diploid human genome in a four-generation, twenty-eight-member family (CEPH 1463).” Page 429: “We successfully assayed 91.9% of the autosomal genome (2.66 Gb)”. “Excluding all variants classified as postzygotic, we find that the parental germline contributes 1.17 × 10−8 SNVs per bp per generation (95% confidence interval (CI) = 1.08 × 10−8–1.27 × 10−8). De novo SNVs are significantly enriched in repetitive sequences, as much as 2.8-fold in centromeres”. ↩ ↩
- 20a.
- 21.
Amster, Guy, and Guy Sella. “Life history effects on the molecular clock of autosomes and sex chromosomes.” Proceedings of the National Academy of Sciences 113(6):1588–1593. 2016. A population-genetics model from Columbia University of how the ages at which males reach puberty and breed, and the pace of sperm production, set the yearly mutation rate in each ape lineage. It accounts for the low X-to-autosome ratio and the small rate differences among humans, chimpanzees and gorillas, and puts the human-chimp split as recent as 6.6 million years. It matters here because it shows how the slowdown repair works without the 10- to 15-year generations that generation length alone would need, and which inputs it rests on. Mirror: PMC
- 21a.
Page 1588, abstract: “we focus on the effects that the age of onset of male puberty and rates of spermatogenesis have likely had in hominids (great apes)”. “In so doing, it largely bridges the gap reported between estimates of split times based on fossil and molecular evidence”. “split may have occurred as recently as 6.6 Mya”. “explaining why the generation time has a major influence on yearly substitution rates in mammals but only a subtle one in human pedigrees”. ↩
- 21b.
Page 1588, abstract: “One of the foundational results in molecular evolution is that the rate at which neutral substitutions accumulate on a lineage equals the rate at which mutations arise.” ↩
- 21a.
- 22.
Callaway, Ewen. “DNA mutation clock proves tough to set.” Nature 519:139–140. Mar 12, 2015. A Nature news report on a February 2015 meeting in Leipzig where about a dozen speakers tried to explain why family-counted human mutation rates come out at half the older fossil-calibrated rate. It records how uncertain the leading geneticists themselves judged the clock to be, and why most of them still preferred the slower rate. Mirror: Archive.org
- 22a.
Paragraph 7: “But calculations using the slow clock gave nonsensical results when extended further back in time — positing, for example, that the most recent common ancestor of apes and monkeys could have encountered dinosaurs.” “many researchers have started hedging their bets in papers, presenting multiple dates for evolutionary events depending on whether mutation is assumed to be fast, slow or somewhere in between”. Paragraph 8: “The fact that the clock is so uncertain is very problematic for us”. “It means that the dates we get out of genetics are really quite embarrassingly bad and uncertain.” Paragraph 13: “Even though the human mutation rate is still uncertain and unstable, Reich proposed at the meeting that researchers use the slower value for their work, at least until better data come along.” “This is not the speed of light. This is not physics.” The speaker in paragraph 8 is David Reich of Harvard Medical School. Paragraphs are counted from the one that begins Mathematicians keep refining π. ↩
- 22a.
- 23.
Tomkins, Jeffrey P., and Jerry Bergman. “Evolutionary molecular genetic clocks: a perpetual exercise in futility and failure.” Journal of Creation 29(2):26–35. 2015. The fullest young-earth critique of molecular clocks, by a geneticist at the Institute for Creation Research and a biologist. It argues that fossil calibration is circular, that rates differ between genes and lineages, that rates counted in families exceed fossil-calibrated ones, and that widespread genome function undercuts the neutral theory clocks rest on. It treats clocks in general and does not work the human-chimp numbers.
- 23a.
Page 26, abstract: “Because the molecular clock idea is directly tied to the neutral model theory of evolution, recent discoveries in full codon utility and pervasive genome-wide biochemical functionality negate its foundational premise.” ↩
- 23b.
Page 26: “human evolutionary geneticist David Reich of Harvard stated”. “It means that the dates we get out of genetics are really quite embarrassingly bad and uncertain.” ↩
- 23c.
Page 28: “The most empirical way to measure genetic change is by determining DNA base substitutions observed between generations of pedigree lines. These have been shown to greatly exceed the more speculative and spurious rates achieved from paleontology.” ↩
- 23a.
- 24.
For three species, two gene copies drawn from the population that lived between the older and the younger split fail to find their common ancestor there with a chance of e-t/2N, where t is the interval in generations and N the population size. One third of those that fail still give the species tree's shape; two thirds give one of the two other shapes. So the share of discordant trees is (2/3)e-t/2N. A 30 % share gives t/2N = ln((2/3)/0.30) = 0.80; 39.5 % gives 0.52. With N = 10,000, a 4-million-year interval and 21.75-year generations the share is 0.0068 %. Yoo and colleagues counted a region as ILS when its tree differed from the species tree, so their 39.5 % is a discordant share like Scally's 30 %.8 ↩
- 25.
Jeanson, Nathaniel T. Replacing Darwin: The New Origin of Species. Master Books. 2017. A book by a young-earth creationist biologist with a Harvard doctorate, then at Answers in Genesis, proposing a creation model of species origins based on genetic clocks. Chapters 7 and 8 compare human and chimpanzee mitochondrial and nuclear DNA with evolutionary and young-earth predictions, and conclude that most nuclear differences between species were created. The article uses it as the fullest young-earth treatment of the human-chimp differences.
- 25a.
Page 213: “Traditionally, evolutionists have put the split time between these species at 4.5 to 7 million years ago.” “In short, if the human and chimpanzee lineages diverged 4.5 million years ago, then only around 8.4 million DNA differences would separate them today. Instead, if the human and chimpanzee lineages diverged 7 million years ago, then only around 13.1 million DNA differences would separate them. Today, over 26 million single base pair differences separate the human and chimpanzee nuclear DNA sequences.” The second quotation is from the caption of Figure 8.3. ↩
- 25b.
Pages 214–215: “Remarkably, the authors of the chimpanzee genome paper picked a third option as an explanation for the nuclear DNA data. They changed the timescale. Rather than put the split in the human and chimpanzee lineages at 4.5 to 7 million years ago, they suggested bumping it to 11 to 17 million years ago.” ↩ ↩
- 25c.
Page 221: “If preexisting differences are used to explain the discrepancy between prediction and fact for humans, why not for the population present at the human-chimpanzee split? In fact, why not for any evolutionary split? When can preexisting differences be invoked, and when should they not be invoked? What testable predictions does the hypothesis of preexisting differences make?” ↩
- 25d.
Page 223: “A creationist would respond by saying that these differences were created. For humans, creationists would invoke this at the origin (i.e., creation) of Adam and Eve.” "These differences" are the DNA differences between species that mutation over 4,500 to 6,000 years cannot account for. ↩
- 25e.
Page 227: “As we have observed for the human and chimpanzee nuclear DNA predictions, evolutionary divergence time does not predict mutation rates. The human-chimpanzee divergence time was revised in light of the mutation rates in these species.” ↩ ↩
- 25f.
Page 232: “In short, it appears that mtDNA differences are due to a constant rate of mutation over 6,000 years, whereas the majority — if not vast majority — of nuclear DNA differences are due to preexisting DNA differences that were placed in the ancestors of modern species 6,000 years ago.” ↩
- 25a.
- 26.
Tomkins, Jeffrey P., and Jerry Bergman. “Incomplete lineage sorting and other 'rogue' data fell the tree of life.” Journal of Creation 27(3):84–92. 2013. A review by two young-earth creationist writers arguing that gene trees which disagree with the species tree, along with microRNA genes and taxon-restricted genes, count against common descent and for separately created kinds. It discusses the human, chimpanzee and gorilla case through Ebersberger and colleagues 2007 and Scally and colleagues 2012. The article uses it as the main young-earth treatment of ILS in apes.
- 26a.
Page 84: “ILS across clearly different and unrelated kinds of organisms represents a mosaic of DNA sequence patterns that cannot be explained by common ancestry.” ↩
- 26b.
Page 85: “‘ancient ancestral polymorphisms’ must involve the transfer of ILS-related fragments across skipped taxa in a lineage—a feat which is impossible for non-interfertile taxa”. ↩
- 26c.
Page 86: “The overall outcome, as with previous studies, was that no consensus path of common ancestry between humans and various apes exists, and no coherent model of primate evolution could be achieved.” ↩
- 26a.
- 27.
Replacing Darwin treats human-chimpanzee nuclear and mitochondrial DNA in chapters 7 and 8 and does not mention incomplete lineage sorting, CpG sites, or the sex-chromosome ratio. Tomkins and Bergman 2013 discuss ILS but give no account of its size, its even split, or its drop near genes and on the X. Wood 2006, Carter 2023 and Sanford and colleagues 2015 do not address the CpG excess or the sex-chromosome ratio. ↩ ↩
- 28.
Venn, Oliver et al. “Strong male bias drives germline mutation in chimpanzees.” Science 344(6189):1272–1275. 2014. A family study from Oxford and the Biomedical Primate Research Centre that sequenced a nine-member pedigree of western chimpanzees. It found the same rate per generation as in humans but a much stronger male bias, and it computed the average time back to the common ancestor of human and chimpanzee DNA. The article uses its male bias and its 13-million-year divergence time.
- 28a.
Section Abstract: “Our results indicate a mutation rate of 1.2 × 10−8 per base pair per generation, but a male contribution seven to eight times that of females and a paternal age effect of three mutations per year of father’s age.” ↩
- 28b.
Section Main text, the paragraph on the fitted mutation model: “indicating a mutation rate of 1.2 × 10−8 per base pair per generation and α to be 7.8 (table S10).” ↩
- 28c.
Section Main text, the paragraph on the neutral substitution rate: “the level of divergence is 1.2% (13), implying an average time to the most common ancestor of 13 million years, assuming uniformity of the mutation rate over this time (95% ETPI 11 to 17 million years; table S11).” ↩ ↩
- 28a.
- 29.
Carter, Robert W., and Matthew Powell. “The genetic effects of the population bottleneck associated with the Genesis Flood.” Journal of Creation 30(2):102–111. 2016. A young-earth population model by Robert Carter of Creation Ministries International and Matthew Powell. It follows created diversity from two people through a Flood that leaves one couple, their three sons and the sons' wives, and finds the Flood cut heterozygosity only slightly in moderate to large populations. Along the way it estimates how many new mutations the long-lived patriarchs could have passed on, which is why it is cited here. Mirror: Creation.com
- 29a.
Page 109: “a man 500 years of age would donate approximately 10,000 mutations to a child (the current average is two orders of magnitude less than that)”. “This would mean Noah would only contribute slightly more than 1,000 mutations (40 baseline mutations + 500 years x 2) after age 500. But they also discuss models with an exponential mutational increase over time and only studied men under age 50.” The 10,000 figure is extrapolated from Crow 2000; the straight-line figure uses Kong and colleagues' 2012 estimate of about two extra mutations per year of the father's age. They refers to Kong and colleagues. ↩
- 29a.
- 30.
This site's arithmetic. If fathers pass on α new mutations for each one from mothers, the Y chromosome, always in males, changes 2α/(1+α) times as fast as the chromosomes that spend half their time in each sex, and the X, two thirds of its time in females, 2(2+α)/(3(1+α)) times as fast. Observed: Y 1.9/1.23 = 1.54, X 0.94/1.23 = 0.76. Family values of α: 3.16 from 1,991 paternal and 630 maternal new mutations in Dutch families,34 3.5 in British families,35 3.9 in Icelandic families4 and 7.8 in a chimpanzee family.28 These predict Y ratios of 1.52, 1.56, 1.59 and 1.77, and X ratios of 0.83, 0.81, 0.80 and 0.74. Solving the observed ratios for α gives 3.4 from the Y and 5.8 from the X. ↩
- 31.
Smith, Thomas C. A. et al. “Large scale variation in the rate of germ-line de novo mutation, base composition, divergence and diversity in humans.” PLOS Genetics 14(3):e1007254. 2018. An analysis of about 143,000 new mutations from three family datasets, asking how their density across the genome in 1-million-letter windows relates to human-chimp divergence, recombination and diversity. It found the expected positive correlation with divergence, but weaker than mutation alone predicts. The article uses it as the direct test of the regional fingerprint.
- 31a.
Section Correlation with divergence: “As expected the divergence along the human lineage (since humans split from chimpanzees) is significantly correlated to the rate of DNMs (Francioli, r = 0.20 p<0.001; Wong, r = 0.16, p<0.001; Jonsson, r = 0.31, p<0.001).” “we would expect the correlation between divergence and DNM density to be 0.30, 0.44 and 0.68 for the Francioli, Wong and Jonsson datasets respectively”. ↩ ↩ ↩
- 31a.
- 32.
Behrens, Sarah, and Martin Vingron. “Studying the evolution of promoter sequences: a waiting time problem.” Journal of Computational Biology 17(12):1591–1606. 2010. A computer model of how long a given short binding site takes to appear at random in a promoter, the stretch of DNA that switches a gene on. It gives 83–338 million years for a five-letter site in one promoter and 7,467 years averaged across 20,000 promoters. Its model has no population or selection; the drift is folded into substitution rates taken from a human, chimpanzee and macaque alignment. The article uses it as the case Sanford answers. ↩
- 33.
Bergeron, Lucie A. et al. “The Mutationathon highlights the importance of reaching standardization in estimates of pedigree-based germline mutation rates.” eLife 11:e73577. Jan 12, 2022. A methods comparison in which five laboratories estimated the mutation rate from the same three-generation family of rhesus macaques, each with its own software and filters. Their answers ran from 0.46 to 0.85 × 10-8 per site per generation. It matters here because the faster yearly rates of the great apes rest on a few families, where such choices can move the result. Mirror: PMC
- 33a.
Section Abstract: “We report almost a twofold variation in the final estimated rate among groups using different post-alignment processing, calling, and filtering criteria”. Section Mutationathon, twofold variation in estimated rates from the same trio: The lowest estimate, 0.46 × 10-8 per site per generation, and the highest, 0.85 × 10-8, came from the same sequence data. ↩
- 33a.
- 34.
Francioli, Laurent C. et al. “Genome-wide patterns and properties of de novo mutations in humans.” Nature Genetics 47(7):822–826. 2015. A study of 11,020 new mutations in 250 Dutch families from the Genome of the Netherlands project. The article uses its count of new mutations traced to each parent.
- 34a.
Section Online Methods: “Using GATK ReadBackedPhasing we assigned parental origin to 1,991 paternal and 630 maternal de novo mutations based on phase-informative reads.” ↩
- 34a.
- 35.
Rahbari, Raheleh et al. “Timing, rates and spectra of human germline mutation.” Nature Genetics 48(2):126–133. 2016. A deep-sequencing study of three large British families from the Wellcome Trust Sanger Institute, which also timed when in development new mutations arise. The article uses its ratio of new mutations from fathers to those from mothers.
- 35a.
Section Results: “Whereas all DNMs showed a 3.5:1 ratio of paternal to maternal mutations”. ↩
- 35a.