Adam, Eve, and Human Genetic Diversity

Three of the four standard arguments against a single-couple human origin do not test it, and BioLogos cut its own claim 36-fold without a correction notice. Two problems survive, unmet by any young-earth calculation: HLA-DRB1 introns in five to nine deeply different families where a couple carries four, and a median four-lineage coalescent age of 358,000 to 495,000 years.

By AI (Claude Opus draft, Claude Fable 5.1 language pass, thinking level unavailable); strengthening review by Claude Opus 5.5, thinking level unavailable Draft:  September 2026

"None of the studies above set out to explicitly test the hypothesis that humans could have passed through a single-couple bottleneck." — Richard Buggs, Professor of Evolutionary Genomics, Queen Mary University of London, 20171

"For humans alive today, the median TMR4A is approximately 500,000 years ago" — Joshua Swamidass, 20222

"This model is not only consistent with what is observed, it would appear to be very hard to falsify." — John Sanford, Robert Carter and colleagues on their own model, 20183

Summary

Could every living human descend from one couple who lived a few thousand years ago?  The genetics is usually reported as a settled no.  It is not settled.  The reasons are not the ones creationists usually give.

What the standard case gets wrong.

  1. Three of the four arguments in Adam and the Genome do not test the hypothesis. Linkage disequilibrium, PSMC population curves, and incomplete lineage sorting with chimpanzees and gorillas each measure something real. None of them measures whether the human line passed through a one-generation crash to two people. The person who says so most bluntly is not a creationist: Joshua Swamidass, a computational biologist at Washington University, writes that "LD, ILS, and Ne are irrelevant lines of evidence, that do not rule out a single-couple bottlenck [sic] in any relevant time scale".2
  2. The fourth argument, allele counts per locus, is false at the level most people picture it. A single base can carry at most four alleles. A couple can carry four. Measured: of 57,190 polymorphic sites in 896 nuclear genes, 113 had three alleles, and the paper reports no site with four.4 That is 99.8% of sites with exactly two. A couple could have carried every per-site allele count in the human genome.
  3. Venema's strongest argument carried no citation. Richard Buggs asked twice for a reference. He was pointed to an online HLA primer with no bottleneck model in it, and said so in print.5
  4. Venema misreported the linkage-disequilibrium paper he cited. Tenesa and colleagues open their discussion with "Overall, the estimates of Ne appear to be much lower than the usually quoted value of 10,000 (Takahata 1993)."6 Adam and the Genome reports the paper as confirming 10,000. It gets there by adding an African estimate of about 7,000 to a non-African estimate of about 3,000.7 The non-African populations descend from the African one, so their variation is a subset of it. Adding a subset to its superset is not a valid sum.
  5. BioLogos deleted the strongest version of its own claim and did not say so. The 2014 article "Adam, Eve and Human Population Genetics" said "There is in fact no point in our evolutionary history over the last 18 million years or more where our lineage was reduced below 10,000 individuals." The version now online says "As far as anyone can tell, the genetic data in fact rules out a single couple for at least half a million years."8 That is a retreat by a factor of 36 in the time depth of the claim. The whole SNP-counting section went with it.

What survives, and it is serious.

  1. HLA-DRB1 holds more deeply different lineages than four founding chromosomes can supply. Counted here from the IPD-IMGT/HLA allele list, release 3.65.0: 4,023 named DRB1 alleles, falling into 13 first-field lineages.9 The 4,023 is not an argument. The immunogeneticists who measured it found that over 90% of DRB1 alleles arose after the split.10 The 13 is not quite an argument either. The same group's 2007 full-length study assigns four of those lineages, 08, 11, 13 and 14, to a radiation from one proto-*03 ancestor within the last 5 million years.11 That leaves nine as a ceiling on the pre-split count. A couple carries four. Measured directly, with no clock and no chimpanzee, the introns of 680 complete DRB1 gene sequences fall into five to nine families more than 1% apart in 16 of 18 stretches of 500 sites, even when each family may be a patchwork of the others.12 Six thousand years of mutation separates two lineages by about 0.0005%.
  2. The median time back to four surviving lineages at a human locus is 495,000 years. This statistic, TMR4A, is the only published analysis that asks the four-allele question directly and genome-wide. Swamidass computes a median of 495,000 years. Half the genome is older than that.2 Venema is quoted by Swamidass as calling it "a better way of looking at this overall".2 The same author's later average across three datasets is lower, 358,000 years.13 For the ten lineages Noah's family could carry, the median is about 180,000 years.2 Robert Carter of Creation Ministries International has criticised the method in print, and one of his points is partly right. No creationist has computed a TMR4A or offered an alternative number.
  3. No fitted single-couple model lands more recently than 500,000 years. Ola Hössjer, Professor of Mathematical Statistics at Stockholm University, and Ann Gauger fitted a single-couple model to real African allele frequencies and to real linkage-disequilibrium decay curves. It fits at about 500,000 years ago with pre-loaded diversity.14 A more recent date is a conjecture about three model extensions they did not try. The only fit at a young-earth date is Sanford and colleagues' 2018 simulation: 200 generations at a census of 1,000, scored by a mean squared error that its authors say "is not a significance test."3
  4. The creationist answer to the four-allele ceiling comes in two versions, and only one of them is science. Hössjer keeps the ceiling. Two people carry four chromosome copies, and the amount of created difference between them is a fitted parameter.15 Sanford, Carter and colleagues drop it. If each of Eve's egg cells was separately created, "there is almost no limit to the number of variant alleles and linkage blocks that could have existed in Eve's ovaries."3 They then write of that model, "This model is not only consistent with what is observed, it would appear to be very hard to falsify."3 That self-assessment overlooks the Flood. However Eve's eggs were made, Noah's family carried at most ten copies of each chromosome, so after the Flood every locus must trace back to at most ten families. That makes even this version testable, and it meets a median time back to ten lineages of about 180,000 years.2
  5. The young-earth model's real problem is not allele counts. It is the number of deeply different families. Computed for this article: start from eight people and grow 5% a generation. In 207 generations, about 195 new mutations arise in any given 2 kb window across the whole population. New alleles are easy. Deep divergence is not: 6,000 years of mutation separates two lineages by about 0.0005% of sites. Created heterozygosity supplies deep divergence among four founding chromosomes at no cost, and that is the model's real strength. It cannot supply a fifth deep family. DRB1 and TMR4A both count deep families, and both find more than four.

The bottom line.  The claim that human genetic diversity disproves a single couple with heliocentric certainty is over-stated, and its own authors have walked it back.  The claim that a single couple 6,000 years ago is genetically unproblematic is also false.  Two measurements stand in the way:  DRB1's lineage count and TMR4A.  No published calculation on the young-earth side has met either.

The four arguments, and what each one tests

Dennis Venema's 2017 book Adam and the Genome put this question in front of a general audience.  Chapter 3 gives four lines of evidence that the human line never fell below about 10,000 individuals.  The book opens with the Tasmanian devil as a picture of a two-individual founding:  "The implications are clear: Tasmanian devils experienced a severe bottleneck in the distant past, but humans did not."7

Three fail, each in a different way.  One survives, at the right level.

# Argument Does it test a one-generation crash to two people? What time depth does it reach?
1 Allele counts per locus At the single-base level, no. At the MHC-lineage level, yes DRB1 lineages predate the human-chimpanzee split
2 Linkage disequilibrium (Tenesa 2007) No, as it was used 10,000 to 200,000 years ago
3 PSMC population curves (Li & Durbin 2011) No 20,000 to 3,000,000 years ago, and badly
4 Incomplete lineage sorting with chimp and gorilla No before 6 million years ago

Argument 2: linkage disequilibrium

Alleles at nearby sites travel together.  Recombination breaks that association up, at a rate that depends on how many ancestors there were.  Albert Tenesa and colleagues measured that decay in HapMap data.  Their answer was smaller than the number Venema reports.  Their abstract:  "For CEU, JPT, and HCB, the estimate of effective population size, adjusted for SNP ascertainment bias, was ∼3100, whereas the estimate for the YRI was ∼7500".6  Their discussion opens by saying these are lower than the usual 10,000.6

Three things are wrong with using this as a disproof of a single couple.

The addition is invalid.  Venema adds 7,000 and 3,000 to reach 10,000.  Buggs, an evolutionary geneticist and not a creationist, wrote:  "It seems to me that the paper gives no warrant for Venema's addition of effective population sizes above and below the Sahara".1

The method's own error bar is large.  Swapping one recombination-rate estimator for another moved every estimate by an average of −27%, with a range from +33% to −45%, giving a mean Ne of 1901.6  A quantity that swings across a 78-point range on one methodological choice cannot support a claim of near-certainty.  There is also a circularity.  The recombination rates themselves are estimated from linkage disequilibrium.1

The window excludes the event.  The method reaches back 10,000 to 200,000 years and no further.  By construction it assumes a population near equilibrium.  A one-generation crash inside a fast-growing population is the one case it does not model.

The strongest reply to this argument is not a creationist objection.  It is a fitted model.  Hössjer and Gauger simulated a single founding couple forward in time.  They compared the resulting r² and D′ decay curves against real African data.  Their Figure 10 caption:  "The two very different scenarios give similar results, both in good agreement with data."14  Linkage disequilibrium does not rule out a single couple.  It never did.

Argument 3: PSMC population curves

Heng Li and Richard Durbin's method reads local coalescence times off the density of heterozygous sites along one diploid genome.  It turns them into a population-size curve.  It is a real measurement.  By the authors' own statement it is blind to the kind of event in question.

Two sentences from their paper decide this.  On resolution:  "More recently than 20kya or more anciently than 3Mya few recombination events are left in the present sequence, which reduces the power of PSMC".16  On sharp events:  "the instantaneous reduction from 12,000 to 1,200 at 100kya in the simulation was spread over the preceding 50,000 years in the PSMC reconstruction."16

A tenfold crash gets smeared across fifty millennia.  A one-generation crash to two individuals is far sharper and shorter than the case they tested.  And the method has no power more recently than 20,000 years ago.  It says nothing about 4,500 years ago in either direction.

Jack Elliott-Higgins and Swamidass tested this directly.  They simulated bottlenecks of 2, 4 and 8 individuals at various dates and asked whether the standard methods could see them.  "No method we tested accurately detected bottlenecks that last just a few generations."13  A bottleneck of two at 75,000 years ago was invisible to MSMC2.  Their conclusion:  "even a single couple bottleneck more ancient than 450 thousand years ago cannot be ruled out by estimates of Ne alone."13

That preprint has limits.  It is not peer reviewed.  The second author is a party to the dispute.  Their own Figure 7 reports that recent bottlenecks are more detectable than ancient ones.  It is a good citation for "these methods are blind to brief crashes."  It is not a citation for the claim that a Flood bottleneck would be undetectable.

Argument 4: incomplete lineage sorting

Where gene trees for human, chimpanzee and gorilla disagree with the species tree, the disagreement rate estimates the size of the population that existed before the split.  Venema's 2014 version put it at about 62,000 individuals from a 30% discordance rate.8

That is a real measurement of a real quantity.  It is not a measurement of the human lineage after the split.  Buggs's one-sentence answer has never been rebutted:  "the occurrence of ILS does not require maintenance of large populations sizes in every lineage after speciation and so does not exclude a bottleneck in the exclusively human lineage".1

One form of the argument does constrain the human line:  trans-species polymorphism, where a human allele is more closely related to a chimpanzee allele than to another human allele.  Such a lineage must have survived in the human line since before the split.  A crash to two people would have to preserve it.  Ellen Leffler and colleagues scanned the genomes for this and found it rare:  "we identified 125 regions in which the same haplotypes are segregating in the two species, all but two of which are non-coding.  In six cases, there is evidence for an ancestral polymorphism that persisted to the present in humans and chimpanzees."17  Swamidass reads the same body of work as permitting "a minimum population limit of 5 to 10 individuals (not thousands) going back 6 million years."2

Five to ten is not four.  But it is nowhere near ten thousand, and it is the mainstream side that says so.

Swamidass then discounts his own number:  "However, HLA genes also show clear evidence of convergent evolution. … For this reason, TSV-based estimates of minimum population size are lower confidence and disputed."2  Convergent evolution means two species can reach similar sequences without inheriting them from a common ancestor.  If that happens at the MHC, some apparent trans-species sharing is not shared history.  The DRB1 argument later in this article is then weaker than a raw lineage count suggests.  That cuts for the young-earth side, and it comes from the mainstream.

The deletion

In June 2021 BioLogos removed "Adam, Eve and Human Population Genetics".  It posted a new article at a different URL, back-dated to the original 2014 date.  Peaceful Science republished the old text under Fair Use with the changes marked.  The diff is mechanical and checkable.  The framing around it, by a party to the dispute, is not.  What follows is the diff.

Deleted:  "The smallest population our lineage was reduced to in the last 15 million years or more, then, was to the bottleneck of 10,000 individuals once our lineage parted ways with the chimpanzee lineage."8

Deleted and replaced:  "There is in fact no point in our evolutionary history over the last 18 million years or more where our lineage was reduced below 10,000 individuals" became "As far as anyone can tell, the genetic data in fact rules out a single couple for at least half a million years."8

Deleted entirely, the SNP-counting section, including:  "SNP diversity in humans is far too large to result from one ancestral couple at any time in the last 200,000 years – we descend from a population."8

Eighteen million years to half a million years is a factor of 36.  The revised claim is defensible.  The original was not.  The revision was made silently.  The argument a skeptic is most likely to have met is the deleted one.

Where the allele count still bites: HLA-DRB1

Argument 1 is the one with force.  It has to be stated at the right level, or it is wrong.

The wrong level.  "There are thousands of HLA alleles and a couple can carry four."  Ann Gauger states the objection in this form and calls it "the strongest case from population genetics against two first parents".18  Lars Johan Erkell, a biologist at the University of Gothenburg, puts it to Hössjer the same way:  "in this model there can be a maximum of four gene variants — alleles — of a certain gene.  However, there are gene families with thousands of alleles.  Where do they come from?"15

The immunogeneticists themselves answered that version in 1998.  Francisco Ayala had dated DRB1 alleles from exon 2, the antigen-binding region.  That region is under strong balancing selection and is reshuffled by gene conversion.  His 1995 Science paper counted 32 human DRB1 lineages already present 6 million years ago and concluded that "no fewer than 16 individuals could have lived at any given time over that long span."19  His PNAS paper the year before had counted 21 at the same date, from 58 alleles rather than 59.  Neither paper mentions the other's figure.20  That 1994 paper is where the number people quote comes from:  "human ancestral populations could never have been smaller than two or three thousand individuals at any time over the last several million years."20  That figure is simulation output, not a lineage count.  His own lineage arithmetic gives a minimum of 29 people.20  Tomas Bergström and colleagues dated the introns instead.  Introns are under no such selection and are not reshuffled.  Their result:  "Alleles within such a lineage, however, appear to be, on average, 250,000 years old, implying that the vast majority (greater than 90%) of the more than 135 contemporary human DRB1 alleles have been generated after the separation of Homo and Pan."10

That is not a creationist claim.  Gaby Doxiadis and colleagues confirmed it independently, in the open literature, in Molecular Biology and Evolution:  "Subsequent intron analyses, indeed, showed that the majority of contemporary HLA-DRB1 alleles have been generated during the past 250,000 years, whereas many DRB lineages and loci predate the divergence of the hominoid species".21

So the raw allele count is not evidence about founders.  Note who this cuts against.  Adam and the Genome appeared in 2017 and still leans on an intuition Bergström had undercut nineteen years earlier.

The right level.  Count lineages, not alleles.  Computed for this article from the official IPD-IMGT/HLA allele list, release 3.65.0 dated 14 July 2026:

Locus Named alleles First-field allelic lineages
DRB1 4,023 13
DQB1 3,121 5
A 9,279 21
B 11,258 36
C 9,416 14

Use DRB1's 13.  Treat the class I numbers as rough.  At DRB1 and DQB1 the first field of the allele name tracks allelic lineage.  That is what Bergström and Doxiadis mean by the word.  At HLA-A, -B and -C the first field originally tracked serological antigen groups.  Those are not guaranteed to be single branches of a tree.  DPB1 was excluded because its first field is a running number, not a lineage label.

Thirteen is not thirteen pre-split lineages.  The honest number is lower.  The same Uppsala group that dated the introns went back with full-length sequences in 2007.  Jenny von Salomé, Ulf Gyllensten and Tomas Bergström report that an ancestral "proto HLA-DRB103 lineage" split into the human-specific lineages 08, 11, 13 and *14 within the last 5 million years.11  So four of the thirteen first-field lineages are a radiation from one ancestor, not four independent ancient lineages.  That leaves nine.

Nine is a ceiling, not a measurement.  Bergström's abstract says "most" of the lineages diverged before the Homo/Pan separation, not all.  Von Salomé's window of "the last 5 million years" straddles the 4 to 7 million-year split date rather than sitting cleanly after it.  The true pre-split count is at most nine.  Nobody has published it.

At most nine lineages against a founding ceiling of four.  That is the argument.  It is smaller than the thirteen a raw count gives, and it is still more than four.

The count can be measured directly, with no clock and no chimpanzee.  A founding couple carries four copies of chromosome 6.  If every living human descends from them a few thousand years ago, then in any short stretch of DRB1 intron, today's sequences must fall into at most four families.  Members of one family can differ only by the mutations a few thousand years add, about 0.0005% of sites.  Recombination and gene conversion can splice the four families into patchworks.  They cannot make a fifth family.

This was computed for this article from the official IPD-IMGT/HLA genomic alignment, release 3.66.0 of 14 October 2026, which holds 680 complete DRB1 gene sequences.22  The 9,042 intron sites that nearly all sequences share were cut into 18 windows of 500 sites.  In each window, sequences differing at five sites or fewer were grouped together, so separate groups differ at more than 1% of sites.  The test then asked how few groups are needed to build all the others as patchworks, allowing two crossovers and two stray mutations in every window.12

Founder sequences needed Windows
4 2
5 5
6 7
7 2
8 1
9 1

Sixteen of the eighteen windows need more than four.  A typical window holds six families:  03, 08, 11, 12, 13 and 14 together;  15 and 16;  04;  07 and 09;  01;  and 10.  Across all 9,042 shared sites, 04, 07 and 09 differ from every other lineage at about 10 to 11% of sites, and 01 differs from 10 at 7%.  Even two of the closest lineages, 11 and 13, differ at 0.22%, over four hundred times what 6,000 years of mutation supplies.

Three limits apply.  The alignment lists named alleles, not how common each one is.  The thresholds are a choice, but tightening them only raises the counts:  at two differences per window, every window needs five to nine.  And the test does not model long gene-conversion tracts copied in from the neighbouring DRB genes that the founding chromosomes would also carry.  That is the one escape left inside the four-chromosome model, and it would have to rewrite intron sequence along most of the gene.

So the four-chromosome model fails at DRB1, and the designed-gametes model does not.  Under designed gametes, described below, the binding limit is the Flood, where Noah's family carried at most ten copies.  The worst DRB1 window needs nine.  DRB1 cannot separate that model from the data.  TMR10A, in the next section, is the test that can.

The creationist side has named the test and has not run it.  Robert Carter, John Sanford and Nathaniel Jeanson answer the allele count, not the lineage count.  Hössjer and Gauger do address the MHC, and precisely:  "In order to test whether or not the first pair hypothesis is correct, at any position along the HLA complex, all lineages should coalesce to at most four ancestral lineages", and they say "previous research indicates perhaps as few as four ancestral lineages" at DRB1.14  They also flag, against themselves, that "there is evidence that HLA genes are subject to convergent evolution".  So the test is stated.  Before this article, nobody had run it.  Their four rests on Titus-Trachtenberg 1994, Bergström 1998, von Salomé 2007 and Gauger's own 2012 book.  None of the three papers states four.  The number traces to the book.

One number often repeated on the creationist side could not be verified.  Gauger writes that Bergström's intron study "concluded that only seven versions of the gene existed in the ancestral population" of humans and chimpanzees.18  Bergström et al. 1998's full text is closed at the publisher and at every open-access route.  Only its abstract could be obtained, and the abstract does not contain the number.  The 1996 Science technical comment that Gauger, Doxiadis and Swamidass all cite is also closed, and PubMed carries no abstract for it.  Anyone repeating "seven ancestral DRB1 alleles" should say it is unverified.  Seven is in any case more than four.

One claim in the same chapter checks out, with named exceptions.  Gauger writes that "It is clear that the intron sequences group according to species, whereas exon 2 sequences show no species-dependent relationships."18  She cites Doxiadis and colleagues 2008.  Its abstract says "most of the primate DRB alleles investigated represent relatively young entities, possessing species-unique sequences", and that this "seems to contradict the current view that the highly similar peptide binding motifs" encoded by exon 2 "predate primate speciation".23  Her sentence is a fair summary of the paper she cites.  The same group's 2012 study names the exceptions:  "apart from the pseudogenes, only DRB5 genes are shared by OWM and HOM".21  So the rule holds for DRB1, with the pseudogenes and DRB5 outside it.

TMR4A, the argument with no answer

Effective population size is a harmonic mean over thousands of generations.  It cannot reveal a single bad generation.  Swamidass says so:  "Ne is, instead, a harmonic average over a very large time window".2  So he built a statistic that asks the question directly.

For each haplotype block in the genome, look at the gene tree and ask how long ago the four most recent surviving lineages coalesce.  Call it the time to the most recent four ancestors, TMR4A.  If all humans descend from one couple, then at every locus all present variation must trace back into at most four founding copies at the moment of the bottleneck.  So the bottleneck must be older than the TMR4A at nearly every locus, not at the average one alone.

The measured median across the autosome is 495,000 years.  Figure 3:  "a single-couple bottleneck is ruled out before 495,000 years ago … The black line identifies the median TMR4A, at 495,000 years ago."2  Median means half the genome is older than that.

Venema accepted the framework.  Swamidass quotes him from a BioLogos forum thread rather than from anything Venema published:  "TMR4A is a better way of looking at this overall — it more directly addresses what we're really interested in."2  Stephen Schaffner independently ran allele-frequency-spectrum simulations and got a compatible answer.  Buggs summarised, "A bottleneck had to be at least 500,000 years ago before anything approaching a good fit was simulated."24  Buggs's closing judgement on the whole exchange is the honest one:

To my mind, the question has now moved on from "Is an 'Adam and Eve' bottleneck inconsistent with human genetic diversity data?" to "At what timescale could an 'Adam and Eve' bottleneck be consistent with human genetic diversity data?"24

This argument has two weaknesses.  First, TMR4A has never been refereed.  It lives in a Peaceful Science Print and a Zenodo deposit, published by a party to the dispute in his own venue.  The single most load-bearing number against a recent single couple has not been through peer review.  Swamidass himself reports that he could not reproduce Schaffner's separate result:  "I was unable to replicate Schaffner's results using simulation software commonly used in the field."2  Second, TMR4A is a molecular-clock statistic.  It converts branch lengths into years using a mutation rate and a generation time.  A young-earth model that rejects the clock is not obliged to accept the conversion.

That second point is weaker than it looks.  Rejecting the clock does not make the branch lengths go away.  The lineages are still separated by the observed number of substitutions.  Something has to have put them there.  That is the arithmetic in the next section.

A creationist has answered in print, and part of the answer is right.  Robert Carter of Creation Ministries International replied in 2021.  He describes the method as working "Given a constant mutation rate, a constant recombination rate, a constant generation time, and human population of at least 10,000 individuals", notes that Swamidass's ground rules excluded a specially created couple, and concludes, "They were not testing the biblical model in any sense."25  Swamidass answers that "No simulations of population history are used, in any way, to infer the ARG and estimate the median TMR4A date," and that the young-earth criticism "indicates misunderstanding of the analysis, not any errors."2

Carter is right about the population assumption, though not in the way he puts it.  No population history is simulated.  But ARGweaver, the program behind the 495,000-year figure, dates its trees against a prior that assumes one constant population size, "N=11,534" for the human genomes.26  Swamidass's own coauthored test of these methods measured what that does after a recent bottleneck:  "ARG inference algorithms, however, systematically overestimated the coalescence times in simulations with recent bottlenecks (Figure 7A), though ARGweaver is least affected."13  Read from their Figure 7, a two-person bottleneck simulated 10,000 years ago comes out about twice as old with ARGweaver, and older still with the two newer methods.  Swamidass's 2022 review cites the same preprint as "empirically validating TMR4A" and does not mention the bias.2

Carter is wrong about created diversity, and the size of the bias matters.  TMR4A counts back to four lineages, not one, precisely so that Adam and Eve may carry four different copies of every chromosome.  Only the designed-gametes version escapes it, and the Flood brings that version back into range through TMR10A.  A twofold overestimate does not close a gap of 60 to 80 times.

The same preprint also gives a lower number than the one usually quoted.  Averaged across three datasets and methods, it reports "TMR3A and TMR4A at 422 kya and 358 kya, respectively".13  And a young-earth model faces a second count.  Noah's family could carry at most ten lineages, and "the median TMR10A is approximately 180,000 years ago."2

In young-earth terms, these numbers set the size of the escape a faster clock would need.  The trees are already dated with a per-generation mutation rate of 1.26 × 10⁻⁸, close to the 1.25 × 10⁻⁸ this article uses.26  To bring the median TMR4A down to 6,000 years, mutation would have to have run about 60 to 80 times faster, on average, across all of human history and the whole genome.  To bring TMR10A down to a Flood 4,500 years ago, about 40 times faster.27  Carter lists candidate causes, among them radiation "perhaps associated with a burst that occurred during the Flood" and what he calls Patriarchal Drive, but in that reply gives no number for any of them.25

Designed heterozygosity: two versions, one falsifiable

The creationist answer to the four-allele ceiling is that Adam and Eve were not created identical.  If their four copies of each chromosome differed from each other, a couple starts with real diversity rather than none.  Hössjer states the mechanism cleanly:  created founder diversity means "All differences between his reference chromosome and the other three founder chromosomes can be thought of as a very large number of mutations, all of which occurred in one generation."28

This idea splits into two very different models.  They should never be cited as one.

Version A keeps the ceiling.  Hössjer's reply to Erkell is explicit:  "in the first founder generation we get four genetically different variants of each non-sex chromosome.  The degree of genetic dissimilarity between these variants, i.e. the degree of created variation, is a parameter that can be varied in the model."15  A parameter that can be varied can be fitted, and a fit can fail.  Hössjer stakes falsifiability on haplotype blocks.  Every human chromosome should be a mosaic of four founder blocks.

Sanford's Designed Alleles Model does the same thing:  starting frequencies restricted to 25%, 50% or 75%, "since there were only four sets of chromosomes in Eden".3  That is a real model with a real ceiling.

Version B abandons the ceiling.  Sanford's Designed Gametes Model reasons that Eve was created rather than born.  Her egg cells never formed by ordinary meiosis, so each could be separately designed.  "Therefore, there is almost no limit to the number of variant alleles and linkage blocks that could have existed in Eve's ovaries."3  Carter says the same in Journal of Creation:  "there is really no limit to the amount of diversity that could have been front-loaded into Adam and Eve."29

Once that move is made, no allele count at any locus can ever count against the model.  Thirteen DRB1 lineages, thirty, three hundred:  all are accommodated in advance.  The authors say so:  "This model is not only consistent with what is observed, it would appear to be very hard to falsify."3

Take that sentence seriously.  A model that cannot fail is not an answer to an observation.  A skeptic handed the paper as a refutation will find that sentence, discount everything else on the page, and be right to.

The Flood puts a ceiling back.  Designed gametes free Adam and Eve's children, not Noah's.  Noah, his wife and the three wives of his sons carried at most ten copies of each chromosome, so after the Flood every locus must trace back to at most ten families.  Swamidass makes the same point and supplies the test:  "the median TMR10A is approximately 180,000 years ago."2  So the designed-gametes model is testable after all, against a ceiling of ten rather than four.  It passes DRB1, whose worst window needs nine families.  It meets TMR10A, 40 times older than a Flood 4,500 years ago on the same clock as TMR4A.

What the fitted models actually found

Hössjer and Gauger 2019 is the strongest technical work on this side.  Its numbers are often misreported, so here they are exactly.  They fitted two scenarios:  a couple 2 million years ago with zero heterozygosity, and "A single couple 25,000 generations ago (about 500kya) having primordial heterozygosity of 0.012, grows rapidly to a population of 16,000 people, then holds steady."14

Primordial heterozygosity of 0.012 means about 1.2% of sites differ between the founders' chromosome copies.  Measured human nucleotide diversity is about 0.1%.  The founders are about twelve times more diverse than any living human.  That is what "designed heterozygosity" has to mean as a number.  It is a coherent thing to claim.

They did not fit anything more recent.  Their own words:  "It is very likely possible to extend our first couple origin model of humanity in order to find more complex scenarios in which the data is compatible with a more recent origin, but we deliberately circumvented fine tuning of the model to data."14  The 100,000-year claim is a conjecture about three model extensions they list and did not try.  Hössjer repeated the position to Erkell.  The good fit holds "if Adam and Eve lived several hundred thousand years ago".  He expects a larger model can probably lower the date.15

A young-earth article that cites Hössjer and Gauger as support for a 6,000-year Adam misrepresents them.  A skeptic can check that in ninety seconds.

Three technical problems with the creationist simulations

Sanford's simulations run at a census of about 1,000 individuals.  The result depends on drift spreading the initial spikes at 25%, 50% and 75% out into the observed mass of low-frequency variants.  Drift at N = 1,000 is far stronger than drift in a real post-Flood population growing into the millions.  The paper flags the caveat and does not answer it.  The comparison is between normalised curve shapes.  Its only fit statistic is a mean squared error, which the authors say "is not a significance test", and there is no confidence interval or null distribution.3  The table is still instructive, and lower is better.  The young-earth model that keeps the four-allele ceiling scores 1.16 × 10⁻³.  The designed-gametes model scores 0.84.  A couple drawn from an old population 200 generations before the end scores 0.70, the best fit in the paper.  All three beat the authors' deep-time model at 5.75, which also ran at a census of 1,000.

The haplotype-block prediction has no citation at its load-bearing number.  The number turns out to be close.  Hössjer writes "Each block comes in a few variants, four for many parts of the genome" and gives no reference for it.28  Stacey Gabriel and colleagues measured it in 2002 across 51 autosomal regions:  "an average of 5.0 common haplotypes" in Yoruban and African-American samples, "4.2 common haplotypes" in European samples, and "3.5" in Asian samples.30  So four is a fair reading of the middle of the range.  It is also where the model gets into trouble.  The African average is above four.  Pooling populations raises it further, from "3.1 – 4.9 haplotypes with a frequency >5%" within any one population to 5.3 in the union.30  Four founder chromosomes cannot supply 5.3 common haplotypes per block.  The gap is small and real.  It has to be closed by mutation, gene conversion or recombination inside blocks.  Nobody has published the distribution, which is the number that would decide it.

The block structure is not evidence of founders anyway.  Jeffrey Wall and Jonathan Pritchard tested the block model directly:  "real data show a partial fit to the haplotype block model;  some regions conform quite well, whereas others do not."31  Simulating recombination hotspots reproduced the observed patterns without any founder interpretation.  Blocks are weak evidence in either direction.

Jeanson and Lisle's created-heterozygosity paper does make a numerical haplotype-block prediction, and deserves credit for it.  Created alleles plus 6,000 years of recombination and gene conversion "plausibly explains the number of haplotype blocks present in the human population today".32  But the number of blocks is not the quantity in dispute.  Recombination and gene conversion shuffle existing variation into new combinations.  They make new blocks.  They do not make two blocks look 495,000 years apart.

What 6,000 years of mutation can and cannot build

Two computations were run for this article, using the audited population-genetics modules already in this project.  Both scripts sit beside this page under research/.

Computation 1:  a founding pair keeps most of its heterozygosity and loses most of its alleles.  Running the Nei, Maruyama and Chakraborty 1975 setup forward, the minimum heterozygosity after a founding pair is 64.5% of the original at a growth rate of r = 1.0, 50.0% at r = 0.5, and 5.9% at r = 0.1.  So a fast-growing pair keeps about two-thirds of the heterozygosity of the population it came from.  That is the number the creationist side quotes.

Now the other half of the same calculation.  Take a source as diverse as the most polymorphic HLA loci, with θ = 40 and heterozygosity 0.976.  Under Ewens's sampling formula, a sample of four gene copies from that source holds an expected 3.86 distinct alleles out of the source's 489.  Across the range θ = 0.16 to 10, which spans ordinary loci up to DRB1's heterozygosity of about 0.91, a pair carries 1.3 to 3.5 alleles from a source holding 3.6 to 136.  A pair's expected allele count never reaches 4 and can never exceed it.

Both halves must always be printed together.  Quoting the heterozygosity number alone makes a bottleneck look costless.  Quoting the allele number alone makes it look fatal.  Both sides of this debate have quoted one and not the other.

Computation 2:  new alleles are cheap, divergence is not.  Take eight founders and 207 generations of 29 years each.  That is 6,000 years, which is generous to the young-earth model.  Add growth of 5% a generation, capped at 8 billion people.  About 7.8 million gene copies are born.  In any given 2 kb window, about 195 new mutations arise somewhere in the whole population.  A young-earth model has no difficulty producing many new, rare alleles.  This is why the SNP-level version of the allele-count argument fails.  Jeanson and Lisle said so in 2016:  "Since only four DNA base-pairs exist, virtually every possible genomic position allele … could have been present at the time of the Flood, if the individuals were heterozygous."32

Now the divergence.  At a human mutation rate of 1.25 × 10⁻⁸ per site per generation and 29 years per generation, two lineages separated for a given time are expected to differ at this fraction of sites:

Time apart Expected divergence
4,500 years 0.0004%
6,000 years 0.0005%
100,000 years 0.0086%
500,000 years 0.043%
6,000,000 years 0.52%

In a 2 kb DRB1 intron, that is 0.010 expected differences across 6,000 years, against 10.3 across 6 million.  Six thousand years of mutation delivers one one-thousandth of the divergence between lineages split at the human-chimpanzee divergence.

This is the whole problem in one line.  The young-earth model can make plenty of alleles.  It cannot make alleles that are dozens of substitutions apart.  It cannot make deeply divergent alleles that are also common.  Both exist and have to come from somewhere.  Created diversity is the young-earth source, and in Hössjer's four-chromosome version it is constrained.  It can supply at most four deeply different families at any stretch of DNA that recombination has not broken up.  That is a prediction an observation can check, and DRB1 and TMR4A are the checks.  The designed-gametes version raises the limit to ten at the Flood, and TMR10A checks that.

What Swamidass's own answer costs

Swamidass has a model that puts Adam and Eve 6,000 years ago and contradicts no genetics.  It is often cited loosely, so here it is exactly.

Genealogical ancestry is not genetic ancestry.  Go back far enough and everyone shares genealogical ancestors.  Most of those ancestors contribute no DNA to any given descendant.  So:  "Adam and Eve, ancestors of us all, could have been de novo created in the Middle East.  As long as their offspring eventually interbred with people outside the Garden, it is possible as recently as six thousand years ago."33

Read the condition.  The model buys a recent Adam by giving up sole genetic ancestry.  There were other people.  The species' genetic diversity comes from them.  Adam and Eve are, in his phrase, "genetic ghosts, who leave us no identifiable DNA."33

That is a real option and some Christians take it.  It is not the young-earth position.  A model in which every human descends genetically from one couple, and again from eight people at the Flood, gets nothing from it.

Objections and responses

"Human effective population size has never been below 10,000." Effective size is a harmonic mean across thousands of generations.  One bad generation barely moves it.  Every party who has looked closely concedes the point, including Buggs and Swamidass.  BioLogos deleted its 2014 version of the claim in 2021.  The objection should be retired.

"There are thousands of HLA alleles and a couple carries four." Wrong level.  Bergström's intron dating found that more than 90% of the 135-plus DRB1 alleles known in 1998 arose after the human-chimpanzee split, by gene conversion and point mutation.  Doxiadis confirmed it in 2012 in the open literature.  The count of 4,023 is not a founder count.  Restate the objection as thirteen lineages against four and it becomes the strongest thing on the skeptic's side.

"The Tasmanian devil shows what a two-individual founding looks like." Devil populations were small for a long time.  That is what destroys heterozygosity.  A single generation at two, followed by fast growth, keeps about two-thirds of heterozygosity.  That was computed above and first shown by Masatoshi Nei, Takeo Maruyama and Ranajit Chakraborty in 197534.  The right prediction for a brief crash is normal heterozygosity with fewer alleles, a far subtler signature than the devil's.

"Hössjer and Gauger proved a single couple is possible." This is a creationist objection to this article's verdict.  It fails.  Their fitted scenarios are 2 million and 500,000 years ago.  The recent date is a conjecture about three extensions they declined to try.  Citing them for a young Adam misreports a paper that is easy to check.

"You quote Buggs against Venema and stop before his conclusion." The article does not stop there.  Buggs dismantled the way three of the four arguments were used.  Then in April 2018 he reported the analyses that put the boundary at 500,000 years.  Quoting his 2017 criticisms without his 2018 conclusion is quote-mining.  He says so himself.

"Created diversity answers the lineage count." Only in the designed-gametes version, and only at DRB1.  Hössjer's version keeps the four-allele ceiling.  He and Gauger name the right test, that every position in the HLA complex should coalesce to at most four lineages.  They assert that prior research already indicates four.  It does not.  The number comes from Gauger's own 2012 book, not from the three papers they cite beside it, Titus-Trachtenberg 1994, Bergström 1998 and von Salomé 2007.  The intron count computed for this article finds five to nine families more than 1% apart in 16 of 18 windows, even when each may be a patchwork of the others.  Hössjer's version fails there.  Sanford's designed-gametes version absorbs any number at Adam but only ten at the Flood.  DRB1's worst window needs nine, so that version survives DRB1 and meets TMR10A instead.

"TMR4A is unrefereed, so it can be set aside." It is unrefereed, and this article says so.  But unrefereed is not wrong.  The response so far is Robert Carter's 2021 objection to its assumptions, not a competing calculation.  The right move is to recompute it from public 1000 Genomes data.  The tools are free.  Nobody has done it.

"If the molecular clock is wrong, the 495,000-year figure collapses." It would move.  It would not collapse.  The clock converts observed substitutions into years.  The substitutions are observed whether or not the conversion is right.  A young-earth model still owes an account of how alleles separated by dozens of differences came to exist, at high frequency, from at most four founding copies.  Rejecting the clock moves the problem.  It does not remove it.  On the per-generation rate the trees were dated with, the clock would have to have run about 60 to 80 times faster, on average and genome-wide, to bring TMR4A down to 6,000 years, and 40 times faster to bring TMR10A down to a Flood 4,500 years ago.27

"Carter showed eight Ark passengers carry most of today's diversity." He showed a real and useful thing.  Measured on HapMap phase 3 data, "1.97 to 4.75 individuals would capture 55% to 77% of the allelic diversity".29  That kills the Tasmanian-devil intuition.  But note what the statistic is.  It counts what fraction of catalogued SNP alleles a small sample of people carries.  It says nothing about whether any single locus holds more than four deep lineages.  That is the question in dispute.  The two must not be run together.  Details of the Flood bottleneck itself, including minimum viable population and inbreeding, are in Population Bottlenecks.

"African genomes look older than everyone else's." This is a real problem, and Hössjer names it himself.  The recent unique-origin model "faces some challenges, in particular to explain why African DNA looks older than non-African DNA."28  He offers candidate explanations without a quantitative test.  Sanford's "demographic stirring" is the same idea and is also untested.  Nobody has simulated a young, structured post-Babel population that reproduces the African and non-African contrast in nucleotide diversity, rare-variant fraction and linkage decay at the same time.

"Nobody has refuted Hössjer and Gauger." No refereed rebuttal exists.  OpenAlex lists seven citing works, and none of them is one.  The nearest thing is Swamidass 2022.  It calls allele-frequency-spectrum inference "low-confidence and disputed" in general, and in the same breath says Hössjer and Gauger's normalised spectrum is "more aligned with the field" than the version the mainstream side published.2  The only substantive response found anywhere is Erkell's blog series with Hössjer's reply.  That is not a journal exchange.  This cuts both ways.  The model has not been refuted.  It also has not been engaged.  Erkell notes that its journal "is not included in the scientific database SCOPUS".15  A model published there will not be engaged by default.

Issues and unanswered questions

These computations would settle the question.  None has been run in the form needed.  Each is cheap, uses public data, and would decide against one side or the other.

  1. A full DRB1 intron analysis, extending the count run here. This article's window count is a first pass at the test Hössjer and Gauger proposed. It finds five to nine families more than 1% apart in 16 of 18 windows. Three steps would make it decisive. Build maximum-likelihood trees for each window rather than distance clusters. Add the neighbouring DRB genes to the alignment, so that long gene-conversion tracts copied from them are tested rather than assumed away. Weight alleles by their measured population frequencies. Bergström found that gene conversion does not appear to extend into the adjacent intron sequences, which already narrows that escape route. If more than four families survive all three steps, the four-chromosome model is dead at DRB1, and only a model with ten deep families at the Flood remains.
  2. An independent TMR4A, reported as a distribution. Recompute it on 1000 Genomes phased haplotypes with an independent ancestral-recombination-graph method. Publish the whole distribution, plus the fraction of loci whose TMR4A exceeds any candidate young-earth date. This is the highest-value piece of original work available on this question.
  3. An identical-by-descent test of a bottleneck of eight. Segment-based methods can see recent severe bottlenecks. They recovered the Ashkenazi bottleneck at 25 to 32 generations. Simulate eight founders at about 155 generations followed by realistic growth. Ask whether that signature would be detectable in 1000 Genomes-scale data. If it would be, and it is not there, a method with resolution at the right depth refutes the Flood bottleneck. If it would not be, a large mainstream objection disappears.
  4. How many common haplotypes does a block actually contain, block by block? Gabriel and colleagues published the averages in 2002: 3.5 in Asian samples, 4.2 in European, 5.0 in Yoruban and African-American, and 5.3 for the union across populations.30 Nobody has published the distribution and its maximum. Recompute it from 1000 Genomes phased haplotypes: for every block, the number of distinct haplotypes covering 90% of chromosomes. Recombination among four founders cannot produce five or more. A substantial tail of blocks with five or more falsifies the created-diversity model at those blocks. A distribution that stops at four vindicates it.
  5. Sanford's simulations at realistic population sizes. Re-run the Designed Alleles Model with growth from eight into the millions rather than a fixed census of 1,000. Report a goodness-of-fit statistic instead of a visual comparison of normalised curves.

Sources that could not be obtained, and what each blocks.  The full texts of Bergström et al. 1998 and von Salomé et al. 2007 are closed at every route tried.  Only their abstracts could be read, from NCBI E-utilities.  So Gauger's "seven ancestral versions" cannot be checked against the paper it is attributed to.  The 1996 Science technical comment on Ayala is closed and has no abstract.  Doxiadis et al. 2008, which Gauger cites for the intron trees, could be read only as an abstract.  Adam and the Genome has no scan on archive.org and none in Open Library.  Its chapter-3 quotations here were confirmed word for word by Google Books search-inside on the volume itself.  That tool returns no page numbers for a reflowable ebook, so none are given.

Verdict

The arguments most often used to rule out a single couple as the sole genetic ancestors of all living humans do not rule it out.  Linkage disequilibrium, population-size curves and incomplete lineage sorting do not test it.  The mainstream authors who use them have conceded as much in print or deleted the claim.  Raw allele counts do not test it either, because most alleles at hypervariable loci are recent.

Two measurements rule it out at a few thousand years, and no published calculation on the young-earth side has met either.  HLA-DRB1's introns, read from 680 complete gene sequences, fall into five to nine families more than 1% apart in 16 of 18 stretches tested, against a founding ceiling of four.  That measurement uses no clock and no chimpanzee.  The median time back to four surviving lineages across the genome is 358,000 to 495,000 years, depending on the data and method, and the median time back to Noah's ten is about 180,000.  Both could be wrong, and this article says where each is soft.  The DRB1 count is this article's own and has not been checked by anyone else.  Swamidass himself reports that HLA genes show convergent evolution, which would blur any count built from the protein-coding exons, though not one built from introns.  TMR4A has never been refereed, and the methods behind it date recent bottlenecks too old, by about twofold for the program behind the 495,000.  Robert Carter has objected to its assumptions in print.  Neither measurement has been challenged with a calculation.

The gap is not about counting alleles.  It is about counting deep families.  Six thousand years of mutation separates two lineages by about 0.0005% of sites.  DRB1's lineages differ in their introns by 0.2% to 11%.  Created diversity can supply four deep families at no cost, and Hössjer's version stops there, which makes it testable.  It fails the test at DRB1.  The designed-gametes version raises the limit to ten at the Flood, passes DRB1, and meets TMR10A, 40 times older than the Flood on the same clock.

What an honest young-earth position looks like here.  Say that the standard disproof is over-claimed, and show the deletion.  Say that the SNP-level version of the allele argument is false, and show the 99.8%.  Then say plainly that DRB1 and TMR4A are unanswered.  Hössjer and Gauger named the DRB1 test and sourced their answer to a book rather than a measurement.  Run on the published sequences, the four-chromosome version fails it.  The best model on this side fits at 500,000 years, not at 6,000.  Closing that gap needs a mechanism nobody has yet specified or measured.  Anything stronger than that is not supported by the data on the table.

Companion pages

  • The literature. Every source consulted, catalogued with what it measured, whether it was read in full, and which way it cuts.
  • Quarantine. The four quotations and numbers that did not survive checking, including one this article got wrong about a creationist source and one that a second access route resolved.
  • Population Bottlenecks. Minimum viable population, inbreeding, purging, and Robert Carter's Flood-bottleneck papers.

Sources


  1. 1.

    Buggs, Richard J. A.  “Adam and Eve: a tested hypothesis?”  Nature Ecology & Evolution Community.  Oct 29, 2017.  A blog post, written for the Nature Ecology and Evolution Community, that tests chapter 3 of Venema's book against the studies Venema cites, and argues that none of those studies set out to test a single-couple bottleneck.  Buggs is Professor of Evolutionary Genomics at Queen Mary University of London and Senior Research Leader at Kew, and is a Christian who is not a young-earth creationist.  The article uses it because Buggs, an evolutionary geneticist and not a creationist, rejects Venema's addition of the African and non-African estimates.

    1. 1a.

      The question comes from religious friends, and asks whether a bottleneck of two is possible, not whether it is probable.  “None of the studies above set out to explicitly test the hypothesis that humans could have passed through a single-couple bottleneck.” ↩

    2. 1b.

      On Venema's use of Tenesa 2007.  “This is a rather inaccurate presentation of the paper. … It seems to me that the paper gives no warrant for Venema's addition of effective population sizes above and below the Sahara”. ↩

    3. 1c.

      “the main method used to estimate the recombination rate by the authors is patterns of LD.  Linkage disequilibrium patterns are also being used to calculate the effective population sizes given a known recombination rate.  A degree of circular reasoning seems to be inevitable here.” ↩

    4. 1d.

      On incomplete lineage sorting.  “Venema makes an argument based on incomplete lineage sorting among humans, apes and gorillas, which gives a large estimated effective population size.  This argument is not relevant if we are only interested in the human lineage (the occurrence of ILS does not require maintenance of large populations sizes in every lineage after speciation and so does not exclude a bottleneck in the exclusively human lineage).” ↩

  2. 2.

    Swamidass, S. Joshua.  “The Misunderstood Science of Genetic Bottlenecks.”  Peaceful Science Prints.  2022.  A review presented at a workshop at the 2022 Annual Meeting of the American Scientific Affiliation and published by Peaceful Science Prints (revised 2023 and 2025), the author's own venue, so it is not peer reviewed, and its author is a party to the dispute.  It argues that the genetic evidence rules out a single-couple bottleneck of two only back to about 500,000 years, not 18 million, and that linkage disequilibrium (LD) and incomplete lineage sorting (ILS) do not test that bottleneck.  The article uses it because its author is not a creationist and says bluntly that LD and ILS are irrelevant to the single-couple question, and because the BioLogos corrections were not transparent or complete.

    1. 2a.

      Section Summary box, page 1:  The misspelling "bottlenck" is the source's own.  “LD, ILS, and Ne are irrelevant lines of evidence, that do not rule out a single-couple bottlenck in any relevant time scale.”  Same box.  “The genetic evidence only rules out a single-couple bottleneck (2, not 10,000) going back about 500,000 years (not 18 million).” ↩

    2. 2b.

      Page 3:  “Venema had claimed that Ne was an estimate of the minimum population size.  But this is just not true.  Ne is, instead, a harmonic average over a very large time window.” ↩

    3. 2c.

      Page 5:  “For humans alive today, the median TMR4A is approximately 500,000 years ago (Figure 3).”  Figure 3 caption:  Figure 3:  “a single-couple bottleneck is ruled out before 495,000 years ago … The black line identifies the median TMR4A, at 495,000 years ago.” ↩ ↩ ↩

    4. 2d.

      Page 5:  Quoting Venema.  “TMR4A is a better way of looking at this overall — it more directly addresses what we're really interested in.”  The note 32 that the source gives for this quotation reads only "BioLogos Forum."  It is a forum post relayed by an opponent, not a published statement of Venema's. ↩ ↩

    5. 2e.

      Page 7:  On trans-species variation.  “in the vast majority of loci there are no TSVs.  In the few loci where TSV is observed, almost all have less than 4 trans-species lineages.  In a few cases, such as the HLA genes, there may be up to 10 or 20 shared alleles, though this result has not been replicated.  This data, if confirmed and all due to shared history, would place a minimum population limit of 5 to 10 individuals (not thousands) going back 6 million years.” ↩

    6. 2f.

      Page 7:  The paragraph immediately after the trans-species-variation estimate.  “However, HLA genes also show clear evidence of convergent evolution.  We expect convergent evolution, moreover, in immune genes because different species are often exposed to the same disease-causing organisms.  For this reason, TSV-based estimates of minimum population size are lower confidence and disputed.” ↩

    7. 2g.

      Page 7:  “At this time, there are at least three reasons this line of evidence is low-confidence and disputed”.  Note 51, on Hössjer & Gauger 2019:  “Unlike Schaffner's write up in 2021, this paper uses a normalized AFS, which is more aligned with the field.” ↩

    8. 2h.

      Page 7:  “I was unable to replicate Schaffner's results using simulation software commonly used in the field.” ↩

    9. 2i.

      Page 6:  “There were eight people on the Ark: Noah, his wife, his three sons, and their three wives, holding 10 lineages at most. Looking across the genome, the median TMR10A is approximately 180,000 years ago. Apart from miraculous intervention or interbreeding between Noah’s lineage and others, the TMR10A rules out a Noahic bottleneck more recent than this time.”  Note 46 explains the ten:  the three sons carry only DNA from Noah and his wife, so the bottleneck holds what five people can carry. ↩ ↩ ↩ ↩

    10. 2j.

      Page 6:  “Young earth creationists have been largely silent on TMR4A, but the criticism that did arise indicates misunderstanding of the analysis, not any errors.”  Note 45 gives the criticism as the 2021 forum thread "Rob Carter Responds to TMR4A".  Note 38, same page:  “No simulations of population history are used, in any way, to infer the ARG and estimate the median TMR4A date.”  Same page, on the 2021 preprint:  “A preprint published in 2021 demonstrates that LiTI is very robust indeed, empirically validating TMR4A with several systematic tests.” ↩ ↩

  3. 3.

    Sanford, John C. et al.  “Adam and Eve, designed diversity, and allele frequencies.”  Proceedings of the Eighth International Conference on Creationism, 200–216.  Creation Science Fellowship.  2018.  A conference paper by John Sanford and Robert Carter of the FMS Foundation with four co-authors, arguing that Adam and Eve could have been created with many genetic variants already in their genomes.  Its simulations use a census of about 1,000 individuals with 100 new mutations per individual per generation, and it concludes that two people carrying such designed alleles could produce the allele-frequency patterns seen today.  The article quotes the paper's own account of its two models, including its claim that one of its models "would appear to be very hard to falsify."

    1. 3a.

      Page 200:  From the abstract, p. 200, stating the objection and keeping the ceiling.  “The logic here is that, since there were only four sets of chromosomes in Eden, all variants would have had an initial frequency of either 25%, 50%, or 75%.  Today, most allelic variants have frequencies in the range of 0–10%.  Therefore, it is claimed that observed human diversity disproves a literal Adam and Eve.”  Their reply, on the same page:  “Yet, Adam and Eve could have been created massively heterozygous.” ↩

    2. 3b.

      Page 208:  The Designed Gametes Model.  “assuming that Eve was created, not born, her eggs could not have formed in the normal way – so each gametogonium would have been miraculously formed and could potentially have been genetically unique.  Therefore, there is almost no limit to the number of variant alleles and linkage blocks that could have existed in Eve's ovaries.”. ↩ ↩

    3. 3c.

      Page 213:  “This model is not only consistent with what is observed, it would appear to be very hard to falsify.” ↩ ↩ ↩

    4. 3d.

      Page 206, table 2:  “The MSE is not a significance test, but the results are a useful guide when tweaking parameters within a model to make the model fit an expected distribution. … The three Adam and Eve models all show a much tighter fit to chromosome 22.”  The table's mean squared errors, ×10⁻³, lower is better:  Evolutionary 5.75;  Evolutionary Adam and Eve 0.70;  A & E, Designed Alleles 1.16;  A & E, Designed Gametes 0.84.  Page 203:  “200-generation biblical framework (including population growth”. ↩ ↩

  4. 4.

    Hodgkinson, Alan, and Adam Eyre-Walker.  “Human triallelic sites: evidence for a new mutational mechanism?”  Genetics 184(1):233–241.  2010.  A population-genetics analysis of human DNA variation that counts sites where three different letters occur in the population.  It finds about twice as many such sites as chance predicts, and proposes that two mutations can arise together in one person.  The article uses its count of three-allele sites, and the absence of any four-allele site, against the four alleles that two people could carry.

    1. 4a.

      Page 236:  From resequencing data on 896 nuclear genes in 90–95 individuals, after removing CpG and coding sites.  “a total of 36,702 transitions, 20,375 transversions, and 113 sites that had three alleles segregating in the human population.”  That is 113 triallelic sites in 57,190 polymorphic sites, or 0.198%, and no four-allele sites are reported.  The paper's own point is that 113 exceeds the chance expectation of 61.15.  Its ratio of observed to expected is given as:  “ratio of observed over expected = 1.85, with a standard error of 0.17, P < 0.001”.  The absolute number is what matters here.  Not from this paper: the 1000 Genomes Phase 3 paper reports the same shape at scale, and its pipeline treats "high confidence bi-allelic variants" as the main class and adds multiallelic variants "onto the haplotype scaffold one at a time." ↩

  5. 5.

    Buggs, Richard J. A.  “Response to Dennis Venema's Blog 'Adam, Eve and Population Genetics: A Reply to Dr. Richard Buggs (Part 1)'.”  richardbuggs.com.  Nov 12, 2017.  A blog reply by Buggs to Venema's answer to Buggs's earlier critique of chapter 3 of Adam and the Genome.  Buggs asks Venema for literature citations for the claims about allele numbers and rare alleles, and calls the HLA argument the strongest one Venema has against a bottleneck of two.  The article uses it because Venema's strongest argument carried no citation, and Buggs asked for one twice.

    1. 5a.

      “As I note in my blog, you give no citation to the scientific literature to back up this point, so it is hard for me to interact with you on it.  I would invite you again to make such a citation so that we can discuss this point further.”  After Venema's reply:  “the online primer has nothing in it about models of past human bottlenecks.  I would invite you to make a more explicit argument on this point, as I think this is the strongest argument that is available to you against a bottleneck of two.” ↩

  6. 6.

    Tenesa, Albert et al.  “Recent human effective population size estimated from linkage disequilibrium.”  Genome Research 17(4):520–526.  2007.  A genome-wide study of the effective population size (Ne), which the paper says determines how much genetic variation a population keeps, estimated from linkage disequilibrium (the tendency of nearby markers to be inherited together) in about 1,000,000 markers across four HapMap populations.  It found about 3,100 for the European, Japanese and Han Chinese samples and about 7,500 for the Yoruba, and the authors say these low values are possibly due to one or more bottlenecks about 10,000 to 200,000 years ago.  The article uses it for two points: the authors' discussion calls the estimates lower than the usual 10,000, and changing the recombination-rate method reduced the estimates by an average of 27%.

    1. 6a.

      Section Abstract, page 520:  “For CEU, JPT, and HCB, the estimate of effective population size, adjusted for SNP ascertainment bias, was ∼3100, whereas the estimate for the YRI was ∼7500 … The estimates of Ne are lower than previously published estimates based on heterozygosity, possibly because they represent one or more bottlenecks in human population size that occurred ∼10,000 to 200,000 years ago.” ↩

    2. 6b.

      Section Discussion, first sentence, page 524:  “Overall, the estimates of Ne appear to be much lower than the usually quoted value of 10,000 (Takahata 1993).” ↩ ↩

    3. 6c.

      Page 523:  A second, model-free recombination estimator.  “changed the estimate of Ne between +33% and −45% with an average reduction of 27% (mean Ne = 1901)”.  Page 525:  The paper's own concluding sentence.  “human effective population size estimated from entire human chromosomes is considerably lower than previously suggested, at least during a bottleneck up to ∼20,000 yr ago when a large expansion began.” ↩

  7. 7.

    Venema, Dennis R., and Scot McKnight.  Adam and the Genome: Reading Scripture after Genetic Science.  Brazos Press.  2017.  A book written for a general audience.  Its third chapter argues that the human line never fell below about 10,000 individuals.  The article uses it because the book puts the single-couple question in front of a general reader and chapter 3 gives four lines of evidence for that claim.

    1. 7a.

      Chapter 3, Adam's Last Stand?:  On the Tasmanian devil.  “The implications are clear: Tasmanian devils experienced a severe bottleneck in the distant past, but humans did not. ... the story of the beleaguered Tasmanian devil provides a nice way to 'see' the sort of thing we would expect if in fact the human race began with just two individuals.” ↩

    2. 7b.

      Chapter 3:  On linkage disequilibrium, quoted by Richard Buggs.  The allele-counting argument in the same chapter cites no literature, and Buggs asked for a citation twice.  “The results indicate that we come from an ancestral population of about 10,000 individuals — the same result we obtained when using allele diversity alone … this approach scaled its analysis to investigate our lineage from the present dating back to approximately 200,000 years ago … The researchers found that, during this period, humans living in sub-Saharan Africa maintained a minimum population of about 7,000 individuals, and that the ancestors of all other humans maintained a minimum population of about 3,000 — once again, adding up to the same value other methods arrive at.”  The sentence that stands immediately before Figure 3.3 in chapter 3.  “ancestral population of about 10,000 individuals the same result we obtained when using allele diversity alone”. ↩

  8. 8.

    Venema, Dennis R.  “Adam, Eve and Human Population Genetics.”  BioLogos.  2014.  A 2014 BioLogos article by Venema, which BioLogos deleted in June 2021 and replaced with a substantially revised article under a different URL and author line, back-dated to November 12, 2014.  This copy is published by Peaceful Science, the website of Joshua Swamidass, with its insertions and deletions marked, and Swamidass is a party to the dispute, so its editorial framing is interested.  The article uses it because BioLogos deleted the strongest version of this claim in 2021, and the revised note called its conclusions "unchanged".

    1. 8a.

      On incomplete lineage sorting.  “the time span that humans and chimpanzees shared a common ancestral population after the gorilla lineage branched off is estimated to be 2 million years (t).  Using an estimate of generation time of 20 years … it was predicted, based on genetic evidence, that this population numbered around 50,000 individuals.  This value, when plugged into the equation, predicts that 25% of the time we expect a discordant pattern when examining human, chimpanzee, and gorilla sequences.  The actual result is about 30%, suggesting a population of about 62,000 individuals.” ↩

    2. 8b.

      Deleted in the 2021 revision.  “There is in fact no point in our evolutionary history over the last 18 million years or more where our lineage was reduced below 10,000 individuals”.  The revision replaced it with this sentence.  “As far as anyone can tell, the genetic data in fact rules out a single couple for at least half a million years.” ↩ ↩

    3. 8c.

      Deleted in the 2021 revision.  “The smallest population our lineage was reduced to in the last 15 million years or more, then, was to the bottleneck of 10,000 individuals once our lineage parted ways with the chimpanzee lineage.” ↩

    4. 8d.

      The entire SNP-counting section was deleted in the 2021 revision, including this passage.  “Tallying up the number of ancestors using this method consistently returns a total minimum population size of about 10,000 individuals: approximately 8,000 ancestors are needed to explain SNP diversity in sub-Saharan Africa, and about 2,000 ancestors for everyone else.  SNP diversity in humans is far too large to result from one ancestral couple at any time in the last 200,000 years – we descend from a population.” ↩

  9. 9.

    IPD-IMGT/HLA Database (ANHIG).  Allelelist.txt, release 3.65.0.  Jul 14, 2026.  A plain-text list of every named HLA allele in the IPD-IMGT/HLA database, one per line with its accession code, such as HLA00001 for A01:01:01:01.  The article counts from it 4,023 named DRB1 alleles in 13 lineages (01, 03, 04, 07, 08, 09, 10, 11, 12, 13, 14, 15 and 16), 3,121 DQB1 alleles in 5, 9,279 A in 21, 11,258 B in 36 and 9,416 C in 14, and uses the DRB1 figure of 13 as its starting count.  DPB1 is left out because its first field is a running number, giving 1,972 distinct first fields for 3,170 alleles, not lineage labels. ↩

  10. 10.

    Bergström, Tomas F. et al.  “Recent origin of HLA-DRB1 alleles and implications for human evolution.”  Nature Genetics 18(3):237–242.  Mar 1998.  A study of the HLA-DRB1 immune-system gene that uses intron sequences, which the authors say give a more recent origin for the alleles than the exon 2 region does.  Most DRB1 alleles arose after humans and chimpanzees separated, and the authors estimate an effective population size of about 10,000 for early hominids over the last million years.  The article uses it because the immunogeneticists who measured the alleles answered Ayala's version in 1998.

    1. 10a.

      Section Abstract:  “The human DRB1 alleles can be divided into groups (allelic lineages);  most of these lineages have diverged from each other before the separation of Homo and Pan.  Alleles within such a lineage, however, appear to be, on average, 250,000 years old, implying that the vast majority (greater than 90%) of the more than 135 contemporary human DRB1 alleles have been generated after the separation of Homo and Pan.”  The same abstract also reports that intron coalescence gives an effective population size of about 10,000 for early hominids over the last million years, which is the mainstream figure reached by a route independent of Ayala's. ↩ ↩

  11. 11.

    von Salomé, Jenny et al.  “Full-length sequence analysis of the HLA-DRB1 locus suggests a recent origin of alleles.”  Immunogenetics 59(4):261–271.  2007.  A study of the complete sequence of the HLA-DRB1 gene (an immune-system gene) in humans and four other apes: chimpanzee, bonobo, gorilla and orangutan.  Its abstract reports that the DRB1*03 lineage is found in all five species, and that an ancestral version split within the last 5 million years into four human-specific lineages.  The article uses it because its authors, the Uppsala group whose 1998 intron dating destroyed the raw allele-count argument, assign four of the thirteen first-field lineages to one recent radiation from a single ancestor, so they are a hostile witness to the lineage count as well as to the allele count.

    1. 11a.

      From the abstract.  “The DRB103 lineage has been found in human, chimpanzee, bonobo, gorilla, and orangutan.  An ancestral 'proto HLA-DRB103 lineage' appeared to have diverged in the last 5 million years into the human-specific lineages 08, 11, 13, and 14.” ↩ ↩

  12. 12.

    Computed for this article by research/drb1-intron-clusters.ts from the IPD-IMGT/HLA genomic alignment DRB1_gen.txt, release 3.66.0.  Shared intron sites are those where at least 90% of the 680 sequences carry a base, 9,042 sites in all, which leaves out insertions found in only some lineages, so indels never raise a count.  Distance is the number of base substitutions.  Groups are single-linkage clusters joining sequences that differ at five or fewer of a window's 500 sites, and groups of one sequence are dropped.  A group counts as explained when it can be built from the consensus sequences of the chosen founder groups with at most two switches between them and two mismatches.  Rerun with a threshold of two differences, every one of the 18 windows needs five to nine founder sequences. ↩ ↩

  13. 13.

    Elliott-Higgins, Jack, and S. Joshua Swamidass.  “Brief Population Bottlenecks Are Beyond The Genetic Streetlight.”  Research Square.  2021.  A preprint that has not been peer reviewed, and whose second author is a party to the dispute.  It simulated brief bottlenecks, down to two individuals, and tested whether the standard demographic methods (MSMC, Relate and a new method called LiTI) could detect them.  None of the tested methods could detect a bottleneck lasting only a few generations, and LiTI could not rule out even a bottleneck of ten individuals at the time of the Youngest Toba Eruption.  The article uses it because it shows that the standard methods cannot see a brief bottleneck at all.

    1. 13a.

      Page 5:  “No method we tested accurately detected bottlenecks that last just a few generations.  We considered bottlenecks of 2 and 8 individuals for one generation, followed by exponential growth. … When simulated 75,000 years ago, nonetheless, these bottlenecks were invisible to MSMC2. … as the time point moves to more ancient than 250 kya, even a bottleneck down to two individuals is invisible to all methods.”  The authors state their limit on the same page.  “The simulations did not model population structure and admixture, nor did they model selection in the genome.”  Their Figure 7 reports that recent bottlenecks are more detectable than ancient ones, so the preprint does not support any claim about a bottleneck 4,500 years ago. ↩

    2. 13b.

      Page 8:  “This indicates that even a single couple bottleneck more ancient than 450 thousand years ago cannot be ruled out by estimates of Ne alone.”  “At the time of the Youngest Toba Eruption, for example, we find an Nmin range of just 6 to 16 individuals.” ↩

    3. 13c.

      Page 7:  “ARG inference algorithms, however, systematically overestimated the coalescence times in simulations with recent bottlenecks (Figure 7A), though ARGweaver is least affected.”  Page 9, figure 7:  “For very recent bottlenecks, the critical TMRxA is overestimated. ARGweaver is least susceptible to this bias.”  “at recent timepoints, Nmin incorrectly rules out bottlenecks that are in fact compatible with the data.”  The size of the effect is read from the figure's log-scale axes, so it is approximate.  For a two-person bottleneck simulated 10,000 years ago, ARGweaver dates the critical point near 20,000 years and sets the minimum population near five rather than two;  tsdate and Relate overestimate more.  No bottleneck more recent than 10,000 years was simulated. ↩

    4. 13d.

      Page 7:  “In human data, the average of three LiTI estimates places the Nmin of 2 individuals at 358 kya, with TMR3A and TMR4A at 422 kya and 358 kya, respectively”.  The three datasets, named on p. 8, are the 1000 Genomes Project and Simons Genome Diversity Project trees built with tsinfer, and ARGweaver trees for 69 Complete Genomics genomes. ↩ ↩

  14. 14.

    Hössjer, Ola, and Ann Gauger.  “A Single-Couple Human Origin is Possible.”  BIO-Complexity 2019(1):1–20.  2019.  A modelling paper by Ola Hössjer, Professor of Mathematical Statistics at Stockholm University, and Ann Gauger of the Discovery Institute's Biologic Institute; neither is a young-earth creationist, and the paper does not argue for a young earth.  It simulates a population descended from one couple, compares the results with African allele-frequency and linkage-disequilibrium data from the 1000 Genomes Project, and finds that a single couple about 500kya fits the data.  The article uses it as the most technical work on the creationist side, and notes that its suggested date of 100kya, or more recently, was never fitted.

    1. 14a.

      Page 9:  “Scenario 1  A single couple 100,000 generations ago (about 2mya) having zero heterozygosity (identical homozygous chromosomes), grows rapidly to a population of 10,000 people, then grows slowly and linearly to 16,000 people near the present.  Scenario 2  A single couple 25,000 generations ago (about 500kya) having primordial heterozygosity of 0.012, grows rapidly to a population of 16,000 people, then holds steady.”  The paper's own footnote 5, on the same page:  “In Scenario 2, most of the primordial heterozygosity is lost in the first few generations due to the slow growth rate.” ↩ ↩

    2. 14b.

      Page 9:  Figure 10 caption, on the linkage-disequilibrium fit:  “Simulation results from Scenarios 1 and 2, compared to actual human genetic data from the African superpopulation.  The two very different scenarios give similar results, both in good agreement with data.” ↩

    3. 14c.

      Page 12:  “The issue of haplotype blocks is of particular interest for the HLA-DRB1 gene of the major compatibility complex of chromosome 6, where previous research indicates perhaps as few as four ancestral lineages [73–76].  In fact, the entire HLA complex could be examined to see if no more than four haplotype blocks are present in any given genome segment. … One must also consider the evidence of shared sequence with other primates, though there is evidence that HLA genes are subject to convergent evolution [77, 78].  In order to test whether or not the first pair hypothesis is correct, at any position along the HLA complex, all lineages should coalesce to at most four ancestral lineages [75].”  Its references 73 to 76 are Titus-Trachtenberg et al. 1994, a study of convergent evolution at DRB1 position 86 in the Cayapa of Ecuador, Bergström et al. 1998, von Salomé et al. 2007, and Gauger, Luskin & Axe, Science and Human Origins, 2012.  None of the three papers states a figure of four ancestral DRB1 lineages: Bergström's abstract says most lineages predate the Homo/Pan split, and von Salomé's says the *03 lineage is shared with four other primates.  The four traces to the book. ↩

    4. 14d.

      Page 12:  “It is very likely possible to extend our first couple origin model of humanity in order to find more complex scenarios in which the data is compatible with a more recent origin, but we deliberately circumvented fine tuning of the model to data. … in light of the many possible extensions, we suggest that it is possible to fit a model to genetic data, for which the founding couple lived 100kya ago or even more recently.”  The three extensions named are a variable mutation rate, population structure and directional selection.  None was fitted.  Page 1:  The paper's own framing, on page 1:  “The problem of inferring history from genetic data is complex and underdetermined;  there are many possible scenarios that would produce the data we see.” ↩

  15. 15.

    Erkell, Lars Johan.  “Breakthrough for Intelligent Design? (part 3).”  Panda's Thumb.  2022.  Part 3 of a seven-part blog series by Lars Johan Erkell, a biologist at the University of Gothenburg, first posted in Swedish on the department's Biolog(g) blog on 13 November 2020.  It is a blog exchange rather than a journal one, and it carries a comment by Ola Hössjer dated 29 June 2021 and Erkell's reply; the English translation was published at Panda's Thumb on 13 November 2022.  Erkell argues that the single-couple model needs a created variation that is justified neither scientifically nor biblically, and the article uses his exchange with Hössjer, whose replies state the four-variant limit and the created-variation parameter.

    1. 15a.

      Erkell says:  “in this model there can be a maximum of four gene variants — alleles — of a certain gene.  However, there are gene families with thousands of alleles.  Where do they come from?” ↩

    2. 15b.

      Hössjer's reply in the same exchange:  “our model M1 assumes that all humans are descended from a first couple, Adam and Eve. … Adam has two genetically different copies of each non-sex chromosome, and so does Eve.  Thus, in the first founder generation we get four genetically different variants of each non-sex chromosome.  The degree of genetic dissimilarity between these variants, i.e. the degree of created variation, is a parameter that can be varied in the model.”.  He also stakes falsifiability on haplotype blocks:  “due to recombinations … our DNA is a mosaic of the four building blocks from each chromosome, with which Adam and Eve were created.” ↩ ↩

    3. 15c.

      Hössjer's reply:  “we used a very simple model with created variation, but without geographic variation.  We also managed to get a good fit of our miracle model to data, if Adam and Eve lived several hundred thousand years ago.  However, we write that with a larger (and more realistic) variant of the miracle model, the age of the first two people can probably be lowered significantly.” ↩

    4. 15d.

      Erkell on the venue:  “The journal is not recognized as a 'real' scientific journal.  For example, it is not included in the scientific database SCOPUS.” ↩

  16. 16.

    Li, Heng, and Richard Durbin.  “Inference of human population history from individual whole-genome sequences.”  Nature 475(7357):493–496.  2011.  A method paper that introduces PSMC, a model that reads the density of heterozygous sites (places where a person's two copies of a chromosome differ) in complete genomes and turns that into a history of population size.  The authors applied it to seven genomes from Chinese, Korean, European and Yoruba people, and their simulations showed that PSMC cannot resolve the most recent or the most ancient times well and spreads sudden population changes over a long period.  The article uses those two limits to show how far the method can be trusted.

    1. 16a.

      Section Results, validation paragraph:  “the variance of the estimate is small between 20kya–3Mya.  More recently than 20kya or more anciently than 3Mya few recombination events are left in the present sequence, which reduces the power of PSMC, and therefore the estimated effective population size (Ne) in these time intervals is not as accurate and has large variance.” ↩

    2. 16b.

      Section Results, validation paragraph:  Buggs quotes this sentence as "spread over several preceding tens of thousands of years."  One of the two versions was edited between the author manuscript and print, and this quotation follows the manuscript.  “the simulations also reveal a limitation of PSMC in recovering sudden changes in effective population size.  For example the instantaneous reduction from 12,000 to 1,200 at 100kya in the simulation was spread over the preceding 50,000 years in the PSMC reconstruction.” ↩

  17. 17.

    Leffler, Ellen M. et al.  “Multiple instances of ancient balancing selection shared between humans and chimpanzees.”  Science 339(6127):1578–1582.  2013.  A genome-wide scan for DNA regions where the same haplotypes (combinations of nearby variants) survive in both humans and chimpanzees, a sign that natural selection has kept variation in place for millions of years.  It found 125 such regions, and in six of them there is evidence that an ancestral polymorphism persisted in both species.  The article uses it because such shared lineages are rare, and a crash to two people would have to have preserved them.

    1. 17a.

      Section Abstract:  “In addition to the major histocompatibility complex (MHC), we identified 125 regions in which the same haplotypes are segregating in the two species, all but two of which are non-coding.  In six cases, there is evidence for an ancestral polymorphism that persisted to the present in humans and chimpanzees.”  The opening sentence sets the scale.  “Instances in which natural selection maintains genetic variation in a population over millions of years are thought to be extremely rare.” ↩

  18. 18.

    Gauger, Ann.  “The Science of Adam and Eve.”  Science and Human Origins, 105–122.  Discovery Institute Press, Seattle.  2012.  Chapter 5 of a book by Ann Gauger, Douglas Axe and Casey Luskin, published by the Discovery Institute Press.  It is a creationist book chapter that argues against a single first couple, and it calls the HLA-DRB1 objection the strongest case from population genetics.  The article quotes the chapter's statement of that objection.

    1. 18a.

      Page 121:  Calls the HLA-DRB1 argument.  “the strongest case from population genetics against two first parents”.  Page 108:  States the problem fairly:  “If there are literally hundreds of alleles for these genes in the present human population, where did they come from?  Two people can pass on at most four versions.” ↩

    2. 18b.

      Page 114:  Gauger's endnote cites Bergström et al. 1998 itself.  “A later study by Bergström et al. examined the same HLA-DRB1 gene, but used intron 2 … this study concluded that only seven versions of the gene existed in the ancestral population from which both chimps and humans supposedly came around 4–6 million years ago, and that the population had an estimated size of 10,000 rather than the 100,000 estimated by Ayala.” ↩

    3. 18c.

      Page 115:  “It is clear that the intron sequences group according to species, whereas exon 2 sequences show no species-dependent relationships.”  Gauger's endnotes 7 and 8 cite Doxiadis et al. 2008, Molecular Immunology 45:2743–2751, whose abstract supports her sentence.  Doxiadis et al. 2012 names the exceptions: the pseudogenes DRB6/DRB2 and DRB7, and DRB5.  Neither is DRB1. ↩

  19. 19.

    Ayala, Francisco J.  “The Myth of Eve: Molecular Biology and Human Origins.”  Science 270(5244):1930–1936.  Dec 22, 1995.  A review in Science on whether modern humans descend from a single woman, the "mitochondrial Eve".  Ayala uses the immune-system gene DRB1 to estimate population sizes, and says the gene lineages coalesce to an ancestor about 60 million years ago, with human ancestral populations of 100,000 or more throughout that time.  The article uses it for the count of 32 DRB1 lineages at 6 million years ago, the figure that the immunogeneticists answered in 1998.  The 32-lineage count is derived from exon 2, the antigen-binding region, which Bergström and colleagues in 1998 showed to be a poor clock.

    1. 19a.

      Page 1931:  “32 human lineages were already in existence around 6 Ma, which implies, correspondingly, that the other 27 lineages arose after the hominid lineage diverged from the African apes.”  “If 32 DRB1 gene lineages have persisted since 6 Ma, it follows that no fewer than 16 individuals could have lived at any given time over that long span.” ↩

  20. 20.

    Ayala, Francisco J. et al.  “Molecular genetics of speciation and human origins.”  Proceedings of the National Academy of Sciences 91(15):6787–6794.  Jul 1994.  A colloquium paper that uses the major histocompatibility complex (MHC), the immune-system gene region, to ask how large human ancestral populations were.  The authors use the 58 known human DRB1 alleles and coalescence theory, and conclude that the ancestral population could not have had fewer than several thousand individuals at any time.  The article uses it as the source of the figure people quote, which the authors reached by simulation rather than by counting.

    1. 20a.

      The 2,000 to 3,000 figure is the output of a simulation asking how large a population must be to retain 95% of the polymorphism, not a direct count.  Page 6791:  “The conclusion follows that human ancestral populations could never have been smaller than two or three thousand individuals at any time over the last several million years.” ↩

    2. 20b.

      Page 6789:  “In order to pass 58 alleles through the generations, no fewer than 29 individuals are required at any time.  As we shall presently see, the size of the human populations needs to be much greater than that in order to retain the DRB1 polymorphisms over the last few million years.” ↩

    3. 20c.

      Page 6789:  Set against the 32 lineages at 6 Ma in Ayala 1995.  The two counts come from different alignments, 58 human alleles in 1994 and 59 human genes among 119 primate genes in 1995, and neither paper reconciles them.  Anyone quoting the 32 should know the same author published 21.  “all 58 DRB1 alleles have persisted through the last 500,000 years, but coalesce into 44 ancestral lineages by 1.7 Myr before present (B.P.), 21 lineages by 6 Myr B.P., and 10 lineages by 13 Myr B.P.” ↩

  21. 21.

    Doxiadis, Gaby G. M. et al.  “Evolution of HLA-DRB genes.”  Molecular Biology and Evolution 29(12):3843–3853.  2012.  A research article that builds evolutionary trees from intron sequences (the non-coding parts of genes) of humans, chimpanzees, rhesus macaques and common marmosets, and traces transposable elements (DNA segments that copy themselves around a genome) shared between them.  It finds that most HLA-DRB1 alleles (versions of an immune-system gene) arose in the past 250,000 years, and that many older DRB lineages predate the divergence of the hominoids, the group of humans and apes.  The article uses it as an independent mainstream study that confirms the young age of most DRB1 alleles, and as the source of the named exceptions to the species pattern of DRB genes.

    1. 21a.

      Page 3844:  “Subsequent intron analyses, indeed, showed that the majority of contemporary HLA-DRB1 alleles have been generated during the past 250,000 years, whereas many DRB lineages and loci predate the divergence of the hominoid species (Erlich et al. 1996;  Bergstrom et al. 1998).” ↩

    2. 21b.

      Page 3843:  From the abstract.  “At least four ancestral DRB gene families appear to have been present before the radiation of OWM and HOM, and one of these even predates the speciation of Old and New World primates.  Two of these families represent the pseudogenes DRB6/DRB2 and DRB7 … Furthermore, all phylogenies of different intron segments show consistently that, apart from the pseudogenes, only DRB5 genes are shared by OWM and HOM, and they demonstrate the common history of certain DRB genes/lineages of humans and chimpanzees.”  The subject of "they" is DRB5.  The sentence says intron trees mostly separate the species, with the pseudogenes and DRB5 as the named exceptions. ↩

  22. 22.

    IPD-IMGT/HLA Database (ANHIG).  DRB1_gen.txt, release 3.66.0.  Oct 14, 2026.  The database's genomic alignment of HLA-DRB1:  680 complete gene sequences, exons and introns, lined up base by base with the exon boundaries marked.  The article counts from it how many deeply different intron families exist, the test that decides whether four founding chromosomes are enough, and the intron divergence between the 13 first-field lineages. ↩

  23. 23.

    Doxiadis, Gaby G. M. et al.  “Reshuffling of ancient peptide binding motifs between HLA-DRB multigene family members: old wine served in new skins.”  Molecular Immunology 45(10):2743–2751.  2008.  A research article on the HLA-DRB genes, which are families of human immune-system genes, as they appear in primates.  Its abstract reports that most primate DRB alleles are relatively young, with sequences unique to each species, and that ancient peptide-binding motifs are frequently reshuffled among the duplicated members of the family.  The article uses it because it is the paper that Gauger's endnotes 7 and 8 cite.

    1. 23a.

      From the abstract.  “most of the primate DRB alleles investigated represent relatively young entities, possessing species-unique sequences.  This seems to contradict the current view that the highly similar peptide binding motifs of many HLA-, Patr- and Mamu-DR molecules, encoded by exon 2 of the DRB gene, represent old entities, which predate primate speciation.  As no evidence was found for convergent evolution, the combination of these two observations indicates that ancient peptide binding motifs are frequently reshuffled among duplicated members of the HLA-DRB multigene family.” ↩

  24. 24.

    Buggs, Richard J. A.  “Adam and Eve: lessons learned.”  Nature Ecology & Evolution Community.  Apr 14, 2018.  A blog post first posted at Nature Ecology & Evolution Community on 14 April 2018.  It reviews what the analyses of Steve Schaffner and Joshua Swamidass show about how long ago a two-person bottleneck would have to lie.  Schaffner's simulations needed a bottleneck at least 500,000 years ago, and Swamidass's analysis found a two-person bottleneck plausible over approximately 500,000 years ago.  The article uses it because Buggs, a critic of Venema's book, reports both results and concludes that the question has moved on to timescale.

    1. 24a.

      On Steve Schaffner's allele-frequency-spectrum simulations.  “A bottleneck had to be at least 500,000 years ago before anything approaching a good fit was simulated.”  On Swamidass's coalescent analysis.  “this analysis suggested that a two-person bottleneck is plausible over approximately 500,000 years ago.” ↩

    2. 24b.

      “the question has now moved on from 'Is an "Adam and Eve" bottleneck inconsistent with human genetic diversity data?' to 'At what timescale could an "Adam and Eve" bottleneck be consistent with human genetic diversity data?'” ↩

  25. 25.

    Carter, Robert W.  “Robert Carter gets everything wrong?”  Creation Ministries International.  Jun 7, 2021.  A reply by Robert Carter of Creation Ministries International to a video critique of his work, updated 10 July 2021.  Its section on TMR4A argues that the method builds in constant rates and a large population and never tested a specially created couple.  It lists reasons the mutation clock might have run faster in the past but gives no calculation.

    1. 25a.

      Section A brief detour:  “Given a constant mutation rate, a constant recombination rate, a constant generation time, and human population of at least 10,000 individuals, how long do you think it would take for you to reach the common ancestor of most people using this method?”  “Thus, Swamidass admits to excluding all aspects of the creationist model from the get go. How, then, is this a refutation of anything?”  “The TMR4A calculation is an interesting intellectual exercise, but it is assuming too much in favour of the evolutionary position. They were not testing the biblical model in any sense.”  Section notes:  Carter's answer to Swamidass's reply.  “The assumptions of deep time and uniformitarianism (i.e. constant recombination rates, etc.) are built into the ARGWeaver algorithm. True, it does not “model” populations, but it is essentially doing this before it spits out the final number.” ↩

    2. 25b.

      Paragraph beginning They are asking difficult questions:  “radiation (perhaps associated with a burst that occurred during the Flood or due to bursts of cosmic rays that would necessarily have occurred during rapid magnetic field reversals), and something I call Patriarchal Drive all act to mess up genetic clocks.” ↩

  26. 26.

    Rasmussen, Matthew D. et al.  “Genome-Wide Inference of Ancestral Recombination Graphs.”  PLoS Genetics 10(5):e1004342.  2014.  The methods paper for ARGweaver, a program that infers the family tree at every point along the genome and dates each branch point.  It applied the program to 54 human genomes from Complete Genomics, dating the trees against a prior that assumes a single constant population size.  The 495,000-year TMR4A figure was computed with this program.

    1. 26a.

      Page 9:  “For this analysis, we assumed K=19, sK=1,000,000 generations, 4Nμ=5.8×10−4, and μ=1.26×10−8, implying N=11,534.”  K is the number of time points on which branch points are dated, sK the oldest of them, and N the constant population size of the prior. ↩ ↩

  27. 27.

    For TMR4A against a creation about 6,000 years ago, 495,000 ÷ 6,000 = 83 and 358,000 ÷ 6,000 = 60.  For TMR10A against a Flood about 4,500 years ago, 180,000 ÷ 4,500 = 40.  Each factor is the average speed-up of the mutation clock over the per-generation rate the trees were dated with, holding generation time fixed. ↩ ↩

  28. 28.

    Hössjer, Ola et al.  “Genetic Modeling of Human History Part 1: Comparison of Common Descent and Unique Origin Approaches.”  BIO-Complexity 2016(3):1–15.  2016.  The first of two review articles comparing a common-descent model of human history with a unique-origin model, in which humanity came from one couple with created diversity; it is a review, not a simulation paper, and the fitting is in Hössjer & Gauger 2019.  Its two unique-origin scenarios are dated at about a million years and at 100,000–200,000 years, neither of them a young-earth date.  The article uses it for the sixth mechanism, created founder diversity, which is the creationist answer to the four-allele ceiling.

    1. 28a.

      Page 5:  “VI. Created founder diversity is biologically plausible for DNA of non-sex chromosomes.  Since there are four copies of each non-sex chromosome in two individuals, we may think of one of the founding male's two copies as a reference or template for the other three.  All differences between his reference chromosome and the other three founder chromosomes can be thought of as a very large number of mutations, all of which occurred in one generation.”. ↩

    2. 28b.

      Page 11:  “the blocks are several thousand nucleotides long, and in spite of this there is very little variation within them.  Each block comes in a few variants, four for many parts of the genome.”.  The sentence carries no citation at all.  The nearest reference, [64] Rosenfeld, Mason & Smith 2012, is attached to the preceding sentence about block length and is about the limitations of the human reference genome.  The measurement Hössjer needed is in Gabriel et al. 2002, which reports averages of 5.0 common haplotypes in Yoruban and African-American samples, 4.2 in European samples and 3.5 in Asian samples.  Four is roughly supported, but the African value exceeds it. ↩

    3. 28c.

      Page 12:  “the Middle East ancestry model faces some challenges, in particular to explain why African DNA looks older than non-African DNA.” ↩

  29. 29.

    Carter, Robert W.  “Effective population sizes and loss of diversity during the Flood bottleneck.”  Journal of Creation 32(2):124–127.  2018.  A short research note by Robert Carter in the Journal of Creation, a creationist journal.  It uses HapMap genotype data, a catalogue of common human genetic variants from 1,301 people in 11 populations, to estimate how many people would carry a given share of today's variation, and finds that 1.97 to 4.75 individuals would capture 55% to 77% of the allelic diversity.  The article grants that this result is real and useful, then separates its sampling statistic from the question of how many deep lineages a single locus holds.

    1. 29a.

      Page 126:  “Across all 11 populations, a single individual carried an average of 31.1% (± 0.005 SD) of the allelic diversity.  It required a minimum of 21 individuals to carry 95% of worldwide SNP diversity and 57 individuals to carry 99%.  This is far greater than the effective population size during the Flood.”  Figure 2 caption, on the arrows marking the minimum and maximum effective population sizes:  “1.97 to 4.75 individuals would capture 55% to 77% of the allelic diversity”.  The running text on the same page gives the same result:  “of only 4.75 people the Ark passengers could easily have contained up to 77% of the then-circulating alleles.  In the worst-case scenario, with an effective population size slightly less than 2.0, approximately 55% of all created alleles would be expected to have been captured.”  Read the statistic carefully.  "Allelic diversity" here is a sampling statistic over SNP sites, not a per-locus allele count: it measures what fraction of catalogued SNP alleles a set of people carries, and says nothing about the number of deep lineages at any locus.  The dataset is HapMap phase 3 release 2, not 1000 Genomes.  Carter also concedes in his note 1 that his use of "effective population size" is non-standard; the correct term for what he computed is founder genome equivalents. ↩

    2. 29b.

      Page 126:  “there are other, more complex models where, for instance, God engineered multiple different genomes into the reproductive cells of both Adam and Eve. … there is really no limit to the amount of diversity that could have been front-loaded into Adam and Eve.”. ↩

  30. 30.

    Gabriel, Stacey B. et al.  “The structure of haplotype blocks in the human genome.”  Science 296(5576):2225–2229.  2002.  A large study of haplotype blocks (stretches of DNA inherited together with little recombination inside them), covering 51 regions with about 13 million base pairs of the genome, in samples from Africa, Europe and Asia.  It finds that the genome can be split into blocks, each carrying only a few common haplotypes, and that the block boundaries and haplotypes are highly correlated across populations.  The article uses its counts of common haplotypes per block, which it sets against the four variants per block that Hössjer assumes.

    1. 30a.

      Section Science Express page 3 of 10:  “Haplotype diversity was greatest in the Yoruban and African-American samples, with an average of 5.0 common haplotypes observed.  Lower diversity was observed in the European samples (4.2 common haplotypes), and the smallest number of common haplotypes (3.5) was observed in the Asian samples.” ↩ ↩

    2. 30b.

      Section Science Express page 3 of 10:  “Each single population sample contained 3.1 – 4.9 haplotypes with a frequency >5%.  The union of these sets, however, contained only 5.3 haplotypes (Fig. 4E).”  In the discussion:  “Within each block, a very small number of common haplotypes (three to five) typically capture ≥90% of all chromosomes in each population.” ↩

  31. 31.

    Wall, Jeffrey D., and Jonathan K. Pritchard.  “Assessing the Performance of the Haplotype Block Model of Linkage Disequilibrium.”  American Journal of Human Genetics 73(3):502–515.  2003.  A study that proposes three quantitative criteria for how blocklike linkage disequilibrium (the tendency of nearby DNA sites to be inherited together) is, and applies them to real data sets and to computer simulations.  It finds that real data show only a partial fit to the block model, and that simulations with recombination concentrated in narrow hotspots fit much better than uniform recombination.  The article uses it because hotspots of recombination can reproduce the block pattern, which weakens the block pattern as evidence of founders.

    1. 31a.

      From the abstract.  “Analyses of several large data sets indicate that real data show a partial fit to the haplotype block model;  some regions conform quite well, whereas others do not. … although the real data are only moderately blocklike, our simulations indicate that, under a model of uniform recombination, the structure of LD would actually fit the block model much less well.  Simulations of a model in which much of the recombination occurs in narrow hotspots provide a much better fit to the observed patterns of LD.” ↩

  32. 32.

    Jeanson, Nathaniel T., and Jason Lisle.  “On the Origin of Eukaryotic Species' Genotypic and Phenotypic Diversity.”  Answers Research Journal 9:81–122.  2016.  A creationist research article in the Answers Research Journal, by Nathaniel Jeanson of Answers in Genesis and Jason Lisle of the Institute for Creation Research.  It argues that created heterozygosity (diversity built into the created kinds) together with natural processes explains the diversity of species, and that comparing mitochondrial and nuclear DNA clocks requires that created diversity.  The article gives it credit for a numerical prediction about haplotype blocks, and argues that the prediction concerns the number of blocks, which is not the quantity in dispute.

    1. 32a.

      Page 98:  “these common variants are inexplicable by constant rates of nuclear SNV mutation over 6000 years (Fig. 3B), but the assumption of created heterozygosity followed by constant rates of recombination and gene conversion plausibly explains the number of haplotype blocks present in the human population today (Fig. 8).”  The prediction is about how many haplotype blocks there are.  That is not the quantity in dispute, because recombination and gene conversion reshuffle existing variation and do not create divergence between versions. ↩

    2. 32b.

      Page 99:  “Since only four DNA base-pairs exist, virtually every possible genomic position allele (i.e., far more than 4–28 gene unit alleles) could have been present at the time of the Flood, if the individuals were heterozygous.” ↩

  33. 33.

    Swamidass, S. Joshua.  The Genealogical Adam and Eve: The Surprising Science of Universal Ancestry.  IVP Academic, Downers Grove.  2019.  A book by S. Joshua Swamidass on the genetics of universal ancestry.  It says that Adam and Eve, if created recently in the Middle East, would be consistent with the genetic evidence, provided their offspring interbred with people outside the Garden.  The article uses it for this recent-Adam model, which gives up sole genetic ancestry for Adam and Eve.

    1. 33a.

      Page 97:  “ENTIRELY CONSISTENT WITH THE GENETIC EVIDENCE, Adam and Eve, ancestors of us all, could have been de novo created in the Middle East.  As long as their offspring eventually interbred with people outside the Garden, it is possible as recently as six thousand years ago.” ↩

    2. 33b.

      Page 133:  “They would also be genetic ghosts, who leave us no identifiable DNA. … The only way science presses on our understanding of Adam and Eve is by suggesting that there were people outside the Garden.”  Page 79:  “The genetic evidence that we descend from Adam and Eve, however, would be lost long ago.  For this reason, there is no genetic evidence for or against the de novo creation of Adam and Eve.” ↩

  34. 34.

    Nei, Masatoshi et al.  “The Bottleneck Effect and Genetic Variability in Populations.”  Evolution 29(1):1–10.  Mar 1975.  A theory paper on what happens to genetic variation when a population shrinks suddenly and then grows fast.  It gives a formula for average heterozygosity (the share of gene copies at a locus that differ from each other) as population size changes, and a simulation for the number of alleles (versions of a gene), and it shows that a fast-growing founder population keeps most of its heterozygosity and loses most of its allelic richness.  The article uses this to explain why a brief crash keeps normal heterozygosity with fewer alleles, and the work is worked through in detail in Population Bottlenecks. ↩