"Thus, one ends up in the curious situation where the fossil record is taken seriously in providing a prior for the ages of nodes, but in retrospect is discovered to be wildly unreliable." — Graham Budd and Richard Mann, Systematic Biology, 20241
"Both scenarios can be distinguished using molecular-clock methods because old groups would have accumulated a substantial number of molecular substitutions, leading to long ancestral molecular branches." — Sandra Álvarez-Carretero and colleagues, Molecular Biology and Evolution, 20262
Summary
A molecular clock dates the split between living groups from their DNA differences. The fossil record dates the same groups by where their oldest members sit in the rocks. This article asks whether the two agree for the great radiations: animals, placental mammals, modern birds and flowering plants, with a look at other groups. It is a test of two mainstream methods against each other. All ages below are the mainstream's figures on the radiometric timescale, printed to show how far apart the two methods are on their own terms. The article does not adopt them. How clocks are built and calibrated is covered in the parent article, Molecular Clocks.
- The clock dates are older than the first fossils in every great radiation. The gap is about 75 to 260 million years for crown animals,31 13 to 18 for placental mammals in the newest study,4 about 27 to 66 for the root of modern birds, and about 14 to 126 for flowering plants, depending on the study. Budd and Mann list beetles, sand dollars and bryozoans with gaps of the same size.1
- The gaps have narrowed over time, but method choices move them as much. The placental gap fell from about 43 million years in 2003 to about 16 in 2022.54 Yet inside a single 2026 paper, changing only the calibration strategy moved the flowering-plant gap from about 102 to 17,6 and changing only the rate model moved the animal date by about 71.2 The trend across decades is mostly a record of those choices.
- Where the two agree, the agreement is often built in. Fossils set the minimum and maximum bounds a clock may use.7 When the maximum is set close to the fossils, the clock lands on the fossils.1 For flowering plants the crown age is "almost entirely conditioned by its own prior distribution".8
- The order agrees even where the timing does not. Animal fossils appear in the rocks in the order the molecular tree branches,1 and flowering-plant pollen types appear in the order the molecular tree predicts.9 Order matched between rock sections is evidence either side may use, and on order the two methods agree.
- Each of the four mainstream explanations has some independent support, and none closes the gap alone. The fossil record is incomplete, but fossil-only sampling models expect gaps of only a few to about a dozen million years for placentals and flowering plants. How well a group fossilizes does not predict how large its gap is: flowering plants leave abundant, durable pollen and still show one of the largest gaps. Bursts of faster evolution during radiations are measured in arthropods10 and proposed for birds,11 but not shown to account for the gaps. Calibration and prior choices are shown to move dates by tens of millions of years. Rate models are shown by simulation to inflate deep dates.12
- The experts dispute the result in print. Budd and Mann conclude that for animals and placentals the clock gives no "evidence for an origin of the clade deeper than what is suggested by the fossil record."1 The clock's authors reply that the young-origin models "have a posterior probability of ∼0."2 Their test assumes rate models that do not allow a short burst, which is the alternative Budd and Mann propose.
- The young-earth critique describes the disagreement correctly and draws more from it than it shows. Tomkins and Bergman write that "divergence dates commonly disagree with paleontology despite being calibrated by it."13 That is true. But both columns sit on the same timescale, so the gap between them measures how imprecise deep clock dates are, not how old the earth is. Young-earth geneticists hold that differences between created kinds were made, not accumulated, so on their own view the deep clock dates are not dates at all.1415 The agreement on order is the part of this evidence their model still has to explain.
Verdict. At the deepest nodes, clock dates and first fossils do not agree, and the disagreement has lasted thirty years. They agree better at shallower nodes and where the fossil record is written into the clock's priors. The gap is real, its size depends heavily on the analyst's choices, and no one explanation has been shown to account for it. It says that deep clock dates are imprecise and lean old. It says nothing about the age of the earth, and the fossils' agreement with the molecular branching order is the stronger signal in this evidence.
Part 1: What is being compared
Two dates for one event. The event is the split that produced the last common ancestor of all living members of a group, called the crown group. A clock estimates when that ancestor lived from the DNA differences among its living descendants. The fossil record gives the age of the oldest fossil that can be placed inside the crown group. Because a group must exist before its first fossil can form, the fossil gives a minimum. Some gap between the two is expected on any account. The question is how large it is, and whether its size can be explained.
The clock is not independent of the fossils. A clock needs a rate, and for deep nodes the rate comes from fossils. Michael Benton and Philip Donoghue, whose rules for fossil calibration are widely used, put it this way: "Fossils can provide good ‘‘minimum’’ age estimates for branches in the tree, but ‘‘maximum’’ constraints on those ages are poorer."7 A calibration, by the consensus rules of James Parham and colleagues, "uses the oldest assigned fossil of a taxon as the basis for its minimum age".16 So a clock analysis takes in the first fossils of many groups as bounds, and then estimates the ages of the nodes between them. When the node in question carries its own fossil minimum, a clock date younger than that fossil is ruled out in advance. The test is only how much older the date comes out.
The prior is not what the analyst typed. Programs combine the bounds on all nodes into a joint "effective prior," which can differ greatly from each calibration taken alone. Budd and Mann define it as "the prior probability distribution that is constructed from the calibration densities by enforcement of known topological constraints."1 Part 7 shows how much this matters. The parent article covers calibration and circularity in general; this article asks only whether the dates and the first fossils agree.
A gap is also predicted by sampling alone. Statistical models of how often fossils are found, using no DNA, put each group's origin somewhat before its oldest fossil. Such models exist for placental mammals and flowering plants, and they give the size of gap the fossil record's own sampling leads one to expect. How complete the record is, and how that is measured, belongs to the sibling article How Complete Is the Fossil Record?; this article uses only the figures those models print.
Part 2: Animals and the Cambrian explosion
The clock says animals are far older than their fossils, and has said so for three decades. The figures below are for the last common ancestor of living animals and of bilaterians, the animals with a left and right side. Each is set against the oldest fossils that the critics of the clock accept.
| Node | Study | Clock estimate, Ma | First accepted fossils, Ma | Gap, million years |
|---|---|---|---|---|
| Bilaterians | Ayala and colleagues 1998 | about 670, protostomes from deuterostomes17 | "≈555"18 | about 115 |
| Bilaterians | Peterson and colleagues 2004 | 573 to 65618 | "≈555" | 18 to 101 |
| Living animals | Peterson and colleagues 2008 | 766 (731 to 803)19 | "younger than about 575"1 | 156 to 228 |
| Bilaterians | Peterson and colleagues 2008 | 643 (617 to 671)19 | "≈555" | 62 to 116 |
| Living animals | Erwin and colleagues 2011 | "nearly 800"20 | "younger than about 575" | about 225 |
| Living animals | dos Reis and colleagues 2015, all strategies | 650 to 8333 | "younger than about 575" | 75 to 258 |
| Bilaterians | dos Reis and colleagues 2015, all strategies | 595.7 to 688.33 | "≈555" | 41 to 133 |
| Living animals | Álvarez-Carretero and colleagues 2026, two rate models | 688.1 (658.1 to 744.9) and 759.0 (694.7 to 825.8)2 | "ca. 564.8"2 | 83 to 261 |
Ayala and colleagues called their 1998 result "consistent with paleontological estimates,"17 and Michael Lee's 1999 recalibration of an older dataset gave animal splits "between 791 and 528 million years ago, a result broadly consistent with the palaeontological documentation of the 'Cambrian explosion.'"21 Since then the estimates have not moved steadily toward the fossils. The newest figures, from the group that wrote MCMCTree, one of the two main dating programs, put crown animals in the Cryogenian, some 80 to 260 million years before the oldest accepted animal fossils. Which rate model is chosen moves the date by about 70 million years on the same data.
The traces set a harder limit than the bodies. Soft-bodied animals rarely fossilize, but animals that crawl and burrow leave traces, and the trace record "is far more continuous than the body-fossil record, particularly with respect to that of soft-bodied organisms," as Gabriela Mángano and Luis Buatois put it.22 Their worldwide compilation found few, simple Ediacaran traces and a sharp rise at the base of the Cambrian. Their verdict: "Trace-fossil information strongly supports the Cambrian explosion, but allows for a short time of phylogenetic fuse during the terminal Ediacaran–Fortunian."22 The numbers are in the Cambrian explosion article. A long fuse of 80 to 260 million years therefore needs animals that left neither bodies nor traces. Kevin Peterson and colleagues proposed exactly that in 2004: "This cryptic precambrian history suggests that these stem-group bilaterians must have been 'micrometazoans,'"18 animals too small to leave burrows. The proposal explains the absence. It was made to explain the absence, and this article found no fossil that confirms it.
The critics' reanalysis, and the reply. Budd and Mann reran the 2015 dos Reis analysis "changing a single number, the soft maximum age of the Metazoa," to 580 million years. "The analysis accommodates this shift simply by bunching up the earliest nodes, including the bilaterians. Once again, there is a close prior–posterior correspondence."1 They concluded that "A fossil-based date of c. 555–540 Ma for the origin of the crown-group bilaterians would thus not be unreasonable."1 Álvarez-Carretero and colleagues answered with a formal comparison of the two hypotheses on the same data. Under the better-fitting rate model the late Ediacaran hypothesis has a posterior probability of 1.70 × 10⁻²⁰ against the unconstrained one,2 and they conclude that "the data provide strong evidence against the hypothesis that molecular data are uninformative about the age of crown Metazoa."2
That reply shows that the sequences prefer longer branches than the young hypothesis allows, under the rate models tested. It does not show that the longer branches mean more time rather than faster change. The authors themselves write that the comparison is "conditioned on the rate model used."2 Michael Lee and colleagues measured arthropod evolution in the Cambrian as "~4 times faster" in form and "~5.5 times faster" in molecules than in all later periods, and found that these fast rates "do not change substantially even if the radiation of arthropods is compressed entirely into the Cambrian (~542 mega-annum [Ma]) or telescoped into the Cryogenian (~650 Ma)."10 So a burst of rate during the radiation is measured, and its size does not decide between a short fuse and a long one. The animal case is open inside the mainstream. For bilaterians the two sides differ by about 40 to 150 million years on the mainstream timescale. Both sides read the same radiometric timescale, so neither brings the rocks any closer to a young-earth reading.
Part 3: Placental mammals and the end-Cretaceous extinction
Three models, named before the genomes arrived. Mark Springer and colleagues credit Kevin Archibald and Douglas Deutschman with "three competing models for the diversification of placental mammals, which are the Explosive, Long Fuse, and Short Fuse Models."23 On the explosive model, placental mammals and their orders arose after the extinction that ended the Cretaceous, about 66 million years ago on the mainstream scale. On the long-fuse model the orders split in the Cretaceous and diversified within themselves afterward. On the short-fuse model even the diversification within orders began in the Cretaceous. No crown placental fossil is accepted before the boundary. Maureen O'Leary and colleagues, scoring 4,541 characters on fossils and living species, named Protungulatum donnae, at 64.85 million years, "the oldest undisputed species within crown Placentalia."24 Budd and Mann add that "there are thousands of stem-group fossils known from before the boundary and no uncontroversial crown-group forms."1 The clocks have favoured the long fuse. Robert Meredith and colleagues wrote in 2011 that "Relaxed clock analyses support the long-fuse model of diversification".25 Springer and colleagues, themselves clock workers, summed up in 2019 that "The Long Fuse Model has received more support from relaxed clock studies than any of the other models, but this model is not supported by morphological cladistic studies that position Cretaceous eutherians outside of crown Placentalia."23
The placental gap has shrunk more than any other.
| Study | Method | Crown placental age, Ma | Gap from Protungulatum, million years |
|---|---|---|---|
| Springer and colleagues 2003 | relaxed clock | 107 (98 to 117)5 | about 42 |
| dos Reis and colleagues 2012 | Bayesian clock, genome-scale | 88 to 9026 | 23 to 25 |
| Álvarez-Carretero and colleagues 2022 | Bayesian clock, 4,705 species | 83.3 to 77.64 | 13 to 18 |
| Álvarez-Carretero and colleagues 2026, same data, two rate models | Bayesian clock | 80.0 (77.4 to 83.2) and 89.2 (85.1 to 94.0)2 | 13 to 18, and 20 to 29 |
| Carlisle and colleagues 2023, with and without Deccanolestes | fossils only, no DNA | 77 to 70, and 70.2 to 6627 | 5 to 12, and 1 to 5 |
| O'Leary and colleagues 2013 | fossils and anatomy | "just younger than the K-Pg boundary"24 | none |
The fall from about 42 million years to about 15 came as data grew from a few genes to genomes and calibrations were revised. O'Leary's team put the crown "just younger than the K-Pg boundary, or ~36 My younger than molecular clock–based mean estimates," while warning that "Ghost lineage estimates are minimum divergence dates and may underestimate the timing of actual splits."24 The fossil-only model of Emily Carlisle and colleagues splits the difference: it puts the root in the latest Cretaceous, but finds that "the majority of orders likely originated around or after the K-Pg boundary."27 Álvarez-Carretero and colleagues agree on the orders: "the latest Bayesian analyses provide diversification timings for ordinal-level crown groups that are close to their fossil records, but not for the common ancestor of the group."2 So for placentals the remaining disagreement is about one node, and it is now roughly 10 to 15 million years on the mainstream scale.
What the remaining gap rests on. Budd and Mann reran the 2022 analysis with the placental calibration set from the oldest fossil to the boundary and got "mean 66.47 Ma, 95% hdi 65.60–68.85 Ma" for uncorrelated rates.1 They argue that a post-extinction origin is the better reading, because survival odds rise after a mass extinction and the clock's own tree shows lineages crossing the boundary "with no apparent perturbation," although mammals suffered "up to 95% of species extinction."1 The reply found the post-extinction model's posterior probability "∼0," with the posterior "very narrow and concentrated at the maximum bound at 66 Ma (Fig. 5), which indicates conflict between the molecular data and the K-Pg constraint used."2 The authors' strongest physical argument is that "forcing divergence timings to equate the age of oldest fossils can result in estimated molecular rates that appear physiologically implausible."2 Matthew Phillips found the reverse risk in the clock: rate models for placentals show "substantial misspecification associated with life history rate correlates," and "divergences deep in the tree will tend to be inflated by the interaction of rate errors and asymmetric confidence in minimum and maximum bounds." His own tighter analysis gave dates "younger than others published recently," consistent with a radiation "near the 66 Ma Cretaceous-Paleogene extinction event."12 The rate question is the hinge of the placental case, and it has not been settled.
Part 4: Modern birds
Two nodes tell different stories. The root of modern birds, the split between the ostrich line and all other birds, still dates well before the oldest crown fossil. The great radiation of Neoaves, the branch that holds most living birds, now dates close to the end-Cretaceous boundary in some studies and well before it in others.
The oldest accepted crown bird is Asteriornis maastrichtensis, a skull from Belgium dated "66.8-66.7 million years ago," which "occupies a position close to the last common ancestor of Galloanserae," the ducks and fowl.28 Below the boundary there are many birds outside the crown. Jacob Berv and Daniel Field note that "The Mesozoic fossil record of crownward stem birds argues against the up to 100 million-year missing fossil record advocated by unconstrained divergence time analyses without a maximum age prior on the root."11
| Node | Study | Clock estimate, Ma | Gap from Asteriornis, million years |
|---|---|---|---|
| Root of modern birds | Brown and colleagues 2008, eight method and setting combinations | 102.8 to 133.229 | 36 to 66 |
| Root of modern birds | Jarvis and colleagues 2014 | "about 100"30 | about 33 |
| Root of modern birds | Kuhl and colleagues 2021 | 9431 | about 27 |
| Ducks against fowl | Kuhl and colleagues 2021 | 62.531 | 4 younger than the fossil |
| First split within Neoaves | Stiller and colleagues 2024 | 67.4 (66.2 to 68.9)32 | about 0 to 3 before the boundary |
| Orders of modern birds | Wu and colleagues 2024 | 24 of 39 orders before the boundary33 | most orders Cretaceous |
The duck-and-fowl row is the one place in this article where a clock date comes out younger than a fossil placed near the node. Kuhl's team say the 62.5 "fits well" with Asteriornis,31 which holds only if Asteriornis sits just outside the crown of ducks and fowl. Field's team placed it "close to" that ancestor. The point matters for the claim that clocks always run older: when a node carries no fossil minimum of its own, the clock can land on either side of a fossil.
Two genome studies of the same year disagree on whether most bird orders are Cretaceous. Josefin Stiller and colleagues, with 363 genomes, found that "Only two divergences occurred before the boundary," and wrote "In contrast to a recent study proposing a diversification of Neoaves during the Upper Cretaceous, we found that the early divergences in Neoaves were tightly associated with the K–Pg boundary."32 The study they answer, by Shaoyuan Wu and colleagues, found that "interordinal diversification of modern birds largely occurred in the Late Cretaceous," with about 15 orders after the boundary and "the remaining 24 orders" before it.33 Both used fossil calibrations and genome-scale data. Wu's team bounded the root between Asteriornis at 66.7 and Ichthyornis dispar at 94.3,33 so their root could not leave that window.
Part of the agreement was put in by hand. Of the bird study by Richard Prum and colleagues in 2015, which put the radiation after the boundary, Berv and Field write that it "employed an informative but soft-bound prior on the root of Neornithes, reflecting the conspicuous absence of avian crown group fossils in the Late Cretaceous Niobrara Formation," and that "Applying this prior compresses the age of the neornithine MRCA by 30 Ma."11 When the absence of fossils is written into the prior, an agreement with that absence is not an independent result.
The rate-burst explanation for birds. Berv and Field propose that birds that survived the extinction were small, that small birds have faster molecular rates, and that this "hidden" rate acceleration "may result in both strict and relaxed molecular clocks over-estimating the age of the avian crown group."11 Their evidence is partly independent of the clock. Body size reconstructed for the ancestor of modern birds came out at 553 grams, "50% smaller than our median body size estimate for the pre-K-Pg assemblage," a fossil bird fauna from the last stage of the Cretaceous.11 But the direct test did not run: "Despite lengthy attempts to reanalyze the avian evolutionary timescale in Coevol 1.4b to directly test for lineage-specific rate accelerations, these analyses failed to converge."11 The mechanism is plausible and partly supported. It has not been shown to close the bird gap.
Part 5: Flowering plants
The fossil side is unusually good. Flowering plants shed pollen in large numbers, and fossil pollen is the record this case turns on. The oldest accepted crown angiosperm fossils are pollen grains: "The earliest unequivocal fossil evidence of crown angiosperms dates to the Early Cretaceous (Valanginian stage; ∼135 Ma)."34 Mario Coiro, James Doyle and Jason Hilton describe what follows: "Pollen floras document a Northern Gondwanan appearance of monosulcate angiosperms in the Valanginian and subsequent poleward spread of monosulcates and tricolpate eudicots, accelerating in the Albian." They add that "supposed pre-Cretaceous angiosperms either represent other plant groups or lack features that might confidently assign them to the angiosperms."9
The clock side ranges over 130 million years.
| Study | Method | Crown angiosperm age, Ma | Gap from Valanginian pollen, million years |
|---|---|---|---|
| Smith and colleagues 2010 | uncorrelated relaxed clock | 217 (182 to 257), "in the Late Triassic"35 | 47 to 122 |
| Magallón and colleagues 2015 | relaxed clocks, crown bound from the fossil record | "An Early Cretaceous crown angiosperm age"36 | about 0 |
| Barba-Montoya and colleagues 2018 | Bayesian, five calibration strategies combined | 149 to 25637 | 14 to 121 |
| Wu and colleagues 2026, hard bounds | Bayesian | 212.30 to 261.266 | 77 to 126 |
| Wu and colleagues 2026, soft bounds | Bayesian | 158.45 to 172.536 | 23 to 38 |
| Wu and colleagues 2026, bounds from a fossil-sampling model | Bayesian | 151.46 to 152.99, "Late Jurassic"6 | 16 to 18 |
| Silvestro and colleagues 2021 | fossils only | probability of a Cretaceous crown "as low as P = 0.002"34 | pre-Cretaceous |
| Wu and colleagues 2026 | fossils only, all angiosperm fossils pooled | 150 to 133, mean 1416 | 0 to 15 |
Hervé Sauquet, Santiago Ramírez-Barahona and Susana Magallón reviewed the range in 2022: "Recent studies have reported a perplexing range of estimates for the crown-group age of angiosperms, from ~140 million years (Ma; Early Cretaceous) to 270 Ma (Permian)." Their diagnosis is that the clock estimates "have little to do with either the amount of molecular data or the number of internal fossil calibrations included. Instead, we argue that this age is almost entirely conditioned by its own prior distribution," so that "contrasting age estimates in the literature mostly reflect conflicting prior distributions."8 The 2026 study by Ruolin Wu and colleagues shows the point inside one paper: the same 644 species and 83 genes give a crown age between 212 and 261 million years under hard bounds and about 152 under bounds drawn from a model of the fossil record.6
The gap shrinks when the fossils are written into the clock. Magallón's Early Cretaceous result came after "The angiosperm crown age was bound within a confidence interval calculated with a method that considers the fossil record of the group."36 Wu's Late Jurassic result came from calibration densities built from 25,685 fossil occurrences, from which "We excluded all pollen records," because pollen is "difficult to attribute consistently to family-level clades."6 Each is a fair method. Neither is an independent check of the fossils, because the fossils set the bounds.
Why pollen makes this case hard to explain away. If flowering plants lived through the Jurassic or Triassic, they shed pollen there. Coiro and colleagues find that the order of pollen types in the rocks "agrees with molecular phylogenetic inferences on the course of pollen evolution, but it conflicts strongly with Triassic and early Jurassic molecular ages, and the discrepancy is difficult to explain by geographic or taphonomic biases."9 They leave one door open: "the record may allow the Late Jurassic existence of ecologically restricted angiosperms, like those seen in the basal ANITA grade."9 That reconciles the record with the youngest clock dates, those near 150 to 160 million years. It does not reconcile it with the Triassic dates that conventional calibrations still produce. The fossil-only estimates disagree among themselves. Silvestro's model found a pre-Cretaceous origin; Budd, Mann, Doyle, Coiro and Hilton disputed it in 2021, arguing that the result follows from the oldest fossil and the number of fossils;38 Wu's pooled run, without pollen, gives the Late Jurassic to earliest Cretaceous.6
Part 6: Other groups, large gaps and small
The pattern is general, but not uniform. Budd and Mann list recent gaps "between the first recognized fossil of a clade and the 95% credible interval of molecular clock estimates": beetles "55–134 myr older than the oldest fossil," sand dollars and sea biscuits "40–65 myr," cheilostome bryozoans "132–238 myr," angiosperms "23–121 myr" and bilaterians "50–143 myr."1 Three more groups show the range.
| Group | Clock estimate, Ma | Oldest fossil, Ma | Gap, million years |
|---|---|---|---|
| Land plants, crown | 515.1 to 470.0, "the majority are dated to around 500" (Morris and colleagues 2018)39 | spores of land-plant type, "Middle Ordovician [Dapingian; 469 Ma"39 | about 1 to 46 |
| Primates, crown | "∼71–63" (Springer and colleagues 2012);40 79.2 to 70.0 (dos Reis and colleagues 2018)41 | Altiatlasius, late Paleocene, "around 58"41 | about 5 to 21 |
| Eukaryotes, crown | 1866 to 1679 (Parfrey and colleagues 2011);42 1842 to 1210 (Betts and colleagues 2018)43 | the red alga Bangiomorpha, "1.047 +0.013/–0.017 Ga"44 | about 160 to 820 |
Land plants are the closest case, and the agreement is partly built in. Jennifer Morris and colleagues put crown land plants near 500 million years, against spores of land-plant type in the Middle Ordovician. But Cambrian spores "of unclear affinity," though "not considered unequivocally embryophyte, informed our soft maximum constraint (515.5 Ma)."39 The top of the clock's range is a fossil bound. The first body fossil, Cooksonia, comes much later, in the Silurian,39 so the land-plant clock agrees with the spores and not with the bodies.
Primate estimates have spanned some 30 million years. Mario dos Reis and colleagues write that before their study "the estimated age of the living primate clade has spanned a 30 My differential, ranging from as young as 55 Ma (Koop et al. 1989) to as old as 87 Ma (Perelman et al. 2011)."41 Springer's 2012 estimate is "markedly younger than most timetree dates (Table 4), some of which are as old at 85–90 Ma."40 The oldest well-preserved early primate skeleton, Archicebus from China, is "about 55 million years ago".45 The primate gap is small on the newest dates and large on the older ones, the same pattern as placentals.
Eukaryotes show how the reference fossil decides the gap. Laura Parfrey and colleagues found their eukaryote ancestor "consistent with the earliest microfossils interpreted with confidence as eukaryotic."42 Holly Betts and colleagues name the red alga Bangiomorpha as "The oldest fossil remains that can be ascribed to crown-Eukaryota".43 Against that fossil the gap is hundreds of millions of years, and Timothy Gibson and colleagues moved that fossil "150 m.y. younger than commonly held interpretations" in 2018.44 Whether a gap is large depends as much on which fossil is accepted as the first as on the clock.
Part 7: How the gap moved as methods changed
From strict clocks to relaxed clocks. Dos Reis, Donoghue and Yang describe the first genome-era conflict: "the clock estimates were much older than the dates suggested by the fossil record, sometimes twice as old."46 Relaxed clocks let rates differ between branches. They narrowed the placental and bird gaps over the next two decades, as the tables in Parts 3 and 4 show, but not the animal gap.
From node dating to tip dating. In tip dating, fossils enter the analysis as dated species with their own anatomy, rather than as bounds on nodes. It was expected to break up long branches and bring dates toward the fossils. For placental mammals it did the opposite. Robin Beck and Michael Lee found that morphological clocks "retrieve implausibly ancient, Late Jurassic–Early Cretaceous estimates for the initial diversification of Placentalia," and that "These dates are much older than all recent molecular and palaeontological estimates."47 Mark Puttick, Gavin Thomas and Michael Benton found the same with a different matrix: "the estimated date for the origin of crown mammals is much older, ∼130–145 million years ago (Ma), than fossil and molecular clock data (∼80–90 Ma)."48 Their title says it plainly: morphological clocks "fail to close the molecular fossil gap."48
Better priors. The newest move is to build calibration bounds from statistical models of the fossil record, as Wu and colleagues did for flowering plants. That closed most of the gap, from over 100 million years to about 17, inside one paper.6 The cost, as Part 5 notes, is that the clock then agrees with the fossils partly because the fossils set its bounds.
The gaps, priced. The script research/gap-pricing.ts takes every clock interval cited in this article for the five best-studied nodes and subtracts the first-fossil age each literature prints.49 Three results follow.
| Measure | Crown animals | Bilaterians | Placentals | Crown birds | Flowering plants |
|---|---|---|---|---|---|
| Studies | 5 | 4 | 5 | 3 | 5 |
| Midpoint gap, first study listed, million years | 192 (2008) | 115 (1998) | 43 (2003) | 51 (2008) | 85 (2010) |
| Midpoint gap, latest study, million years | 127 to 185 (2026) | 87 (2015) | 16 to 25 (2026) | 27 (2021) | 17 to 102 (2026) |
| Trend, million years per decade | −28 | −11 | −9 | −18 | −22 |
| Spread inside one paper, million years | 71, rate model only | none listed | 9, rate model only | none listed | 85, calibration only |
| Gap as a share of the fossil's age, latest | 22 to 32 % | 16 % | 24 to 38 % | 41 % | 13 to 75 % |
- Every group's gap has narrowed, at 9 to 28 million years per decade.
- But one analyst's choice inside one paper moves the gap by 9 to 85 million years, which is as much as one to four decades of that trend. The trend cannot be read as convergence on a true date until those choices are settled.
- Most gaps exceed the clock's own precision floor. Dos Reis and Yang showed that a deep date's 95 % interval cannot shrink below about ±16 % of the date, however much DNA is added.50 Of the 22 clock estimates in the script, 18 have a gap larger than that floor. The gap is not just noise.
Part 8: What explains the gaps, and which explanations are tested
The field offers four explanations. Each makes a prediction that can be checked against something other than the gap it was offered to explain.
| Explanation | What it predicts | Independent check | Result |
|---|---|---|---|
| The record is incomplete | Gaps larger where fossilization is poor; fossil-only models expect similar gaps | Ghost lineages, sampling models, preservation potential | Real, but explains only part of the size |
| Rates sped up during radiations | Faster rates at the base of each radiation, tied to body size or generation time | Rates measured on fossils and anatomy; body size across the extinction | Measured in one case, partly supported in another, not shown to close a gap |
| Calibration and priors drive the dates | Dates move with priors more than with data | Rerunning the same data with different bounds | Demonstrated, by tens of millions of years |
| Rate models are wrong | Dates move with the rate model; simulations recover inflated ages | Simulation and model comparison | Demonstrated |
1. Incompleteness is real, and smaller than the gaps. Ghost lineages, the stretches that a lineage must have existed because its sister group is older, give a fossil-only measure of what is missing. Álvarez-Carretero and colleagues count from O'Leary's fossil tree "61 ghost lineages among the basal branches that they sampled in placental mammal phylogeny, equating to a total of over ∼780 Myr of evolution with no record."2 That is about 13 million years per lineage on average, close to the remaining placental gap of 13 to 18.51 For placentals, then, incompleteness of the size needed is measured independently. The fossil-only sampling models say the same: Carlisle's model expects a gap of about 1 to 12 million years, and Wu's pooled model for flowering plants expects 0 to 15. The clock gaps in the newest studies are 13 to 29 for placentals and 17 to 126 for flowering plants.49 Sampling accounts for part of each gap, and in flowering plants only the gap under fossil-derived bounds.
The other prediction fails. If the gaps came from poor preservation, groups that fossilize badly should have the largest gaps. Ranking the five nodes by how well their first fossils preserve, with early animals lowest and pollen highest, gives a rank correlation with the relative gap of +0.39 across all 22 clock estimates and −0.15 across the latest estimates alone.49 Neither is the strong negative value that preservation alone predicts. Five groups and ranks of this article's choosing make that a weak test. The pollen case makes it a strong one. Coiro and colleagues judge the flowering-plant discrepancy "difficult to explain by geographic or taphonomic biases."9 For birds, Berv and Field note a rich record of stem birds just below the crown.11 For placentals, Budd and Mann count "thousands of stem-group fossils" before the boundary.1 In each case the rocks below the first crown fossil are full of the group's close relatives, and the crown members are the ones missing. An explanation by incompleteness has to say why the record missed only them.
2. Rate bursts are measured but not shown to close any gap. Lee and colleagues measured about 5.5 times faster molecular evolution in Cambrian arthropods, and found it robust whether the fuse was short or long.10 That makes a burst real and, for arthropods, uninformative about the fuse. For birds, Berv and Field's body sizes are partly independent support,11 and their direct test did not converge.11 For placentals, Budd and Mann argue that rates "could be much higher than the long-term mean when short intervals are considered,"1 and the clock's authors reply that the required rates "appear physiologically implausible."2 Neither paper prints the rate it means. The models used to reject the young-origin hypotheses, autocorrelated and independent lognormal rates, do not allow a short burst at one point in time.2 So the formal test has not yet been run against the alternative it is cited to defeat.
3. Priors are shown to move the dates by tens of millions of years. Rachel Warnock, Ziheng Yang and Philip Donoghue found that the bounds a user types "are not the effective priors implemented in calculating the posterior time estimates," and that changing a lognormal calibration's settings can move the estimates and their intervals "by hundreds of millions of years."52 Budd and Mann moved the animal date to the fossils by changing one bound.1 Sauquet's team found that the flowering-plant crown age is set mainly by its prior,8 and Wu's team moved it by about 85 million years with calibration choices alone.6 This explanation is tested, not fitted. It has a direction, too. Phillips argues that "asymmetric confidence in minimum and maximum bounds" inflates deep dates,12 and Francisco Rodríguez-Trelles, Rosa Tarrío and Francisco Ayala that the statistics of rate estimation "introduces a bias toward an overestimation of time since divergence."53
4. Rate models are shown to move the dates. On the same data and the same calibrations, the two rate models in the 2026 reply put crown animals 71 million years apart and crown placentals 9 apart.2 Phillips used fossil calibrations to test placental rate models and found "substantial misspecification associated with life history rate correlates."12 Beck and Lee warn that with relaxed clocks "relatively minor deviations in rate patterns can generate large effects on estimated divergence dates."47
Fitted after the fact. Two explanations were proposed to cover a particular absence and have no independent confirmation yet. One is Peterson's microscopic early bilaterians, which would leave no traces.18 The other is ecologically restricted early flowering plants, which Coiro and colleagues allow for the Late Jurassic only.9 Both are reasonable. Neither is evidence until its fossils are found.
Part 9: The young-earth reading
The positions, in their own words
The method critique. Jeffrey Tomkins and Jerry Bergman list among the clock's standing problems that "divergence dates commonly disagree with paleontology despite being calibrated by it,"13 and judge that "the prospects of achieving any type of evolutionary concordance has been dismal."13 Brian Thomas of the Institute for Creation Research, writing on the Cambrian in 2008, drew the young-earth conclusion from the spread of animal dates: "Either the molecular clocks are wrong, the rock dates are inaccurate, or both." He added that "creation scientists would suggest that assumptions of evolution may influence both of them too strongly to allow any confidence in either."54 Michael Oard notes that "The molecular clock often suggests that fossils should be found earlier," and that fossil ranges keep being pushed back.55
The burial-order model. Leonard Brand and Arthur Chadwick hold that first appearances record when habitats were destroyed in the Flood, not when groups arose: "These changes in the fossils do not represent the appearance of newly created organisms. They resulted from the successive final destruction of some habitats and the death and burial of organisms from each of those habitats as the catastrophe unfolds."56 Of birds, mammals and flowering plants: "Perhaps these groups were largely inhabitants of upland regions that were not affected until late in the geological catastrophe."56 Kurt Wise proposes separate pre-Flood biomes: "It is also likely that the gymnosperm-dinosaur biome was located at a lower altitude or closer to the shore of the antediluvian world. This would explain why members of the gymnosperm-dinosaur biome are consistently buried beneath members of the other biome."57
Created differences. For differences between created kinds, young-earth geneticists use no clock. Robert Carter writes that dating the great-ape lineages amounts to "uncovering the differences God built into their initial genomes."15 Nathaniel Jeanson writes of human-chimpanzee differences that "A creationist would respond by saying that these differences were created."14
An intelligent-design reading, kept separate. Stephen Meyer, who is not a young-earth creationist, argues that "Proponents of deep divergence don't deny that the fossil evidence has come up short. Instead, they adopt one of the versions of the artifact hypothesis to account for that missing evidence."58
The positions, tested
The description is right. The dates do disagree with the first fossils, by the amounts in Parts 2 to 6, and mainstream authors say so in print. Thomas took his range of "586 to over 1200 million years" for animal origins from a 2008 review by the biologist Jeffrey Levinton.54 Oard's remark that clocks predict earlier fossils is also correct, and it points the right way: each fossil found below an old first appearance shrinks a gap.
"Despite being calibrated by it" needs care. A clock is calibrated by first fossils at many nodes, as minima. The test is whether the nodes between them come out near their own first fossils. Where the node's own fossil sets a firm bound, agreement is built in. Where it does not, disagreement is a finding, not a contradiction in the method. Part 7's numbers support the critics on a narrower point: the dates move with the analyst's choices as much as with the data.
Thomas's "or both" is the consistent young-earth position, and it ends the test. Both columns of every table in this article rest on the same radiometric timescale. A reader who rejects that timescale rejects both columns, and the gap between them then measures nothing about the age of the earth. Inside the mainstream, the gap shows that deep clock dates are imprecise and lean old. It does not show that the earth is young, and the young-earth authors quoted here do not claim it does.
The burial-order model gets the sign right and nothing more yet. If every group lived from creation and was buried by habitat, every group is older than its first fossil, so a clock date older than the first fossil is expected. But the model as published gives no size for any gap, and no reason why the placental gap should shrink with better data while the animal gap does not. Its authors name its hardest case themselves: "why is there no flowering plant pollen below the Middle Cretaceous? Pollen is easily carried long distances by wind or water." Brand and Chadwick add, "Our current understanding of this model leaves many unanswered questions."56 The same pollen record is the strongest evidence against the Triassic clock dates for flowering plants,9 so it presses both readings at once.
The order is the harder test, and it is open to both sides. Order matched between rock sections does not depend on radiometric dates, and young-earth geologists accept it: Wise and Donna Richardson conclude that "the global biostratigraphic column seems to be real."59 On that order, the clock's branching and the rocks agree. Budd and Mann find "a compelling congruence between the order of appearance of fossils in the record and phylogeny,"1 and Coiro's team find that the sequence of pollen types "agrees with molecular phylogenetic inferences on the course of pollen evolution."9 A burial-order model sorts by habitat and elevation. Nothing in it, as published, says why habitat order should match the branching order of DNA trees. Whether ecological zonation reproduces the order of the fossil record is tested in The Order of the Fossil Record, which finds that it does not. Whether trees and rock order agree beyond chance is in Phylogenetic Inference and Congruence.
Created differences leave the clock dates without meaning, and leave the order to explain. On Carter's and Jeanson's view, sequence distances between kinds are not times, so the deep clock dates in this article are neither right nor wrong. That is a coherent position, and nothing here refutes it. It does leave the order unexplained: the branching order read from those differences is the order in which the groups first appear in the rocks. Designed differences need not nest in an order that burial follows. A created-kinds model that predicted the match would answer this article's strongest finding. None is published.
Objections and responses
1. "The clock is calibrated on fossils, so comparing it with fossils is circular." Partly true. Where a node carries a firm bound from its own first fossil, agreement is built in, as Parts 1 and 7 show. The comparison is informative only at nodes left loose, and those are the nodes where the gaps appear. The objection explains why some agreements mean little. It does not remove the disagreements.
2. "The fossil record is incomplete, so of course groups are older than their first fossils." A gap is expected, and fossil-only models give its expected size: a few million years for placentals and flowering plants. The clock gaps are larger, and they are not largest where fossilization is worst. Flowering plants leave abundant pollen and still have one of the biggest gaps, and the beds below the first crown birds and placentals are full of their stem relatives.
3. "The gaps are closing. Give the method time." They have narrowed by 9 to 28 million years per decade. But a single paper's choice of rate model or calibration moves them by 9 to 85 million years, and some of the closing came from building the fossil record into the priors. Closing by construction is not convergence on a true date.
4. "The 2026 model comparison settles it: the DNA rejects young origins." It shows that the sequences prefer longer branches than a young origin allows under the two rate models tested. Neither model allows a short burst at the radiation, which is the alternative Budd and Mann propose and which Lee and colleagues measured in arthropods. Until a burst model is compared, the question of time against rate is open.
5. "Clock dates miss the fossils by hundreds of millions of years, so the method is worthless and evolution is false." The misses show that deep clock dates are imprecise and lean old, which mainstream authors concede. They do not bear on common descent, because the same trees put the groups in the order the rocks show. They do not bear on the age of the earth, because both dates sit on one timescale.
6. "Rate acceleration is an ad hoc rescue." Not wholly. The Cambrian arthropod burst was measured from anatomy and DNA without reference to the gap, and the bird body-size shift is partly independent. The ad hoc explanations are the ones with no independent trace yet: microscopic early bilaterians and hidden Jurassic flowering plants.
7. "If every kind existed from creation, young-earth creation predicts that every group is older than its first fossil." It predicts the sign. It does not predict the sizes, why they differ between groups, or why the placental gap shrank with data while the animal gap did not. Brand and Chadwick concede that it does not yet explain the missing pollen. And it does not yet say why burial order matches the molecular branching order.
8. "The first fossils are disputed, so the gaps mean nothing." For placentals and birds the alternatives move the first-fossil age by a million years or two, and the gaps stand. For eukaryotes the choice of first fossil matters greatly, as Part 6 shows, and that case is weighted accordingly here.
9. "Fossil-only models also assume deep time." They do. They are used here only to show what the mainstream's own sampling models expect, so that the clock and the fossils can be compared on the mainstream's terms. They do not bear on the age of the earth either.
Issues and unanswered questions
- No fossil-only origin model was found for animals or birds. Such models exist for placentals, primates and flowering plants. An animal model would need the trace-fossil record as well as bodies.
- No published test compares node-age hypotheses under a rate-burst model on the animal or placental datasets. That test would separate time from rate, and both sides in the 2024 to 2026 exchange point toward it.
- The preservation test is weak. It uses five nodes and ranks set by this article. A proper test would use a measured preservation rate per group, such as the sampling rates estimated by fossil-only models.
- Two figures in the sources disagree with themselves. Álvarez-Carretero and colleagues give "at least 720 Myr" of placental ghost lineages in the abstract and "over ∼780 Myr" in the text.2 Carlisle and colleagues report 12 of 23 clades overlapping and 10 not, which sums to 22.27
- Some primary sources were not read in full: Wray and colleagues 1996 on billion-year animal dates; Prum and colleagues 2015, whose node ages are known here only through Berv and Field; Ksepka and Phillips 2015 on bird dating; Lee 1999 beyond its abstract; Heads 2009 on primate biogeography.
- A Jurassic flower is claimed. Brand and Chadwick report that "A perfect angiosperm flower has been found in the Jurassic."56 Coiro and colleagues judge every supposed pre-Cretaceous angiosperm to be another plant or unassignable.9 That specimen is not assessed here.
- No young-earth model predicts the size of any gap or the match between branching order and rock order. The burial-order models of Brand, Chadwick and Wise are qualitative. A model that ranked groups by pre-Flood elevation and predicted their first appearances, tested against the molecular branching order, would be the natural next step for that side.
Verdict
At the deepest nodes, clock dates and first fossils do not agree. For crown animals the clock runs 80 to 260 million years older than the oldest accepted animal fossils, on the mainstream timescale. For flowering plants it runs 14 to 126 older than the first pollen. For the root of modern birds it runs 27 to 66 older than Asteriornis. For placental mammals the gap has fallen from about 42 to about 15, and the placental orders and the main bird radiation now date close to their fossils. Land plants and primates show small to moderate gaps.
The disagreement is real, but its size is not a stable measurement. Within one paper, the analyst's choice of rate model or calibration strategy moves it by 9 to 85 million years. Where the clock and the fossils agree, the fossils often set the bounds. Of the mainstream explanations, calibration and prior effects and rate-model error are demonstrated by sensitivity tests. Incompleteness is measured and accounts for part of each gap, but not where fossils are most abundant. Rate bursts are measured in one group and proposed in another, and have not been shown to close any gap. The experts most involved disagree in print, and the decisive test, time against rate, has not been run.
For the young-earth question the gap is neutral, and the order is not. Both dates rest on the same timescale, so their disagreement says nothing about the age of the earth, as Thomas's "or both" already concedes. What this evidence does show is that the branching order read from DNA matches the order of first appearances in the rocks. That match is open to both sides, does not depend on radiometric dating, and is the part of the evidence a young-earth model has yet to explain.
Sources
- 1.
Budd, Graham E., and Richard P. Mann. “Two Notorious Nodes: A Critical Examination of Relaxed Molecular Clock Age Estimates of the Bilaterian Animals and Placental Mammals.” Systematic Biology 73(1):223–234. 2024. A reanalysis of the published clock datasets for the origin of bilaterian animals and of placental mammals. It argues that the deep dates come from how the calibrations and priors were set rather than from the sequences. Mirror: PMC
- 1a.
Section Animal Origins: “In contrast, the oldest generally accepted animal fossils are younger than about 575 Ma”. ↩ ↩ ↩
- 1b.
Section When the Priors and Posteriors Disagree: the Case of the Placental Mammals: “mean 66.47 Ma, 95% hdi 65.60–68.85 Ma for uncorrelated runs”. With the placental calibration set as a uniform range from the oldest fossil down to the end-Cretaceous boundary. ↩
- 1c.
Section Animal Origins: “We then re-ran the focal analysis of dos Reis et al. (2015) but changing a single number, the soft maximum age of the Metazoa, to reflect the diversification seen in the fossil record, that is, to 580 Ma”. “the analysis accommodates this shift simply by bunching up the earliest nodes, including the bilaterians. Once again, there is a close prior–posterior correspondence.” ↩ ↩ ↩
- 1d.
Section Abstract: “We conclude that in neither case does the molecular clock analysis provide any evidence for an origin of the clade deeper than what is suggested by the fossil record.” ↩
- 1e.
Section Animal Origins: “A fossil-based date of c. 555–540 Ma for the origin of the crown-group bilaterians would thus not be unreasonable. In addition, there is a compelling congruence between the order of appearance of fossils in the record and phylogeny”. ↩ ↩ ↩
- 1f.
Section Introduction: “Thus, one ends up in the curious situation where the fossil record is taken seriously in providing a prior for the ages of nodes, but in retrospect is discovered to be wildly unreliable.” ↩
- 1g.
Section Introduction: “effective prior: the prior probability distribution that is constructed from the calibration densities by enforcement of known topological constraints.” ↩
- 1h.
Section When the Priors and Posteriors Disagree: the Case of the Placental Mammals: “the subsequent lineages rapidly increase across the K-Pg boundary with no apparent perturbation”. “numbers up to 95% of species extinction being suggested”. ↩
- 1i.
Section Introduction: “To give some recent examples of the estimated gaps between the first recognized fossil of a clade and the 95% credible interval of molecular clock estimates: beetles (55–134 myr older than the oldest fossil (Cai et al. 2022)); sand dollar and sea biscuit echinoids (40–65 myr (Mongiardino Koch et al. 2022)); cheilostome bryozoans (132–238 myr (Orr et al. 2022)); angiosperms (23–121 myr (Barba-Montoya et al. 2018)); and bilaterians (50–143 myr (dos Reis et al. 2015)).” ↩ ↩
- 1j.
Section Discussion: “it is entirely plausible that all these rates could be much higher than the long-term mean when short intervals are considered.” ↩
- 1k.
Section When the Priors and Posteriors Disagree: the Case of the Placental Mammals: “there are thousands of stem-group fossils known from before the boundary and no uncontroversial crown-group forms”. ↩ ↩
- 1a.
- 2.
Álvarez-Carretero, Sandra et al. “On the origin of animals and placental mammals: a critique of literalist readings of the fossil record.” Molecular Biology and Evolution 43(9):msag211. 2026. The reply of the MCMCTree group to Budd and Mann 2024, testing their constrained models by Bayesian model selection.
- 2a.
Section Introduction: “Both scenarios can be distinguished using molecular-clock methods because old groups would have accumulated a substantial number of molecular substitutions, leading to long ancestral molecular branches.” “conditioned on the rate model used”. ↩ ↩
- 2b.
Section Introduction: “O’Leary et al. (2013) identified 61 ghost lineages among the basal branches that they sampled in placental mammal phylogeny, equating to a total of over ∼780 Myr of evolution with no record”. The abstract of the same paper gives at least 720 Myr. The figure is read from O'Leary's fossil-based tree, not from a clock. ↩ ↩
- 2c.
Section A Bayesian model test of diversification timings in animals: “ca. 564.8 Ma, as of Μatthews et al. 2020”. The oldest Ediacaran animal fossils, as Budd and Mann date them. ↩
- 2d.
Section A Bayesian model test of diversification timings in animals: “Thus, the data provide strong evidence against the hypothesis that molecular data are uninformative about the age of crown Metazoa.” ↩
- 2e.
Section A critique of the literalist interpretation of the fossil record: “the latest Bayesian analyses provide diversification timings for ordinal-level crown groups that are close to their fossil records, but not for the common ancestor of the group.” ↩
- 2f.
Section A Bayesian model test of diversification timings in mammals: “Budd and Mann's post-K-Pg origination model for Placentalia has a posterior probability of ∼0. Furthermore, the posterior distribution of Placentalia's age is very narrow and concentrated at the maximum bound at 66 Ma (Fig. 5), which indicates conflict between the molecular data and the K-Pg constraint used.” ↩
- 2g.
Section On the nature of molecular rates: “As pointed out by Springer et al. (2013), forcing divergence timings to equate the age of oldest fossils can result in estimated molecular rates that appear physiologically implausible.” ↩ ↩
- 2h.
Section Abstract: “the explosive diversification models previously proposed for animals and placental mammals have a posterior probability of ∼0. We show the characteristic pathologies purportedly discovered do not exist”. ↩
- 2i.
Table 1: Bayesian selection of the animal origination model on the dos Reis 2015 data. Unconstrained model, 833 to 552.85 Ma: autocorrelated rates (GBM), posterior model probability 1.00, crown Metazoa 688.1 (658.1, 744.9) Ma; independent lognormal rates (ILN), probability 1.04e-10, 759.0 (694.7, 825.8) Ma. Late Ediacaran model, 580 to 552.85 Ma: GBM probability 1.70e-20, 579.4 Ma; ILN probability 1.46e-39, 579.3 Ma. ↩ ↩ ↩ ↩ ↩
- 2j.
Table 2: Bayesian selection of the placental origination model on the Álvarez-Carretero 2022 data. Unconstrained model, 162.5 to 61.66 Ma: GBM, posterior model probability 1.00, crown Placentalia 80.0 (77.4, 83.2) Ma; ILN, probability 1.96e-45, 89.2 (85.1, 94.0) Ma. ↩
- 2a.
- 3.
dos Reis, Mario et al. “Uncertainty in the Timing of Origin of Animals and the Limits of Precision in Molecular Timescales.” Current Biology 25(22):2939–2950. 2015. A Bayesian dating study of animal origins from 203 nuclear proteins in 71 species, run under four fossil-calibration strategies, two relaxed-clock models, five ways of partitioning the data and 161 alternative trees. It measures how much each choice moves the dates and finds the combined uncertainty too large to decide between rival causes of the Cambrian radiation. Mirror: PMC
- 4.
Álvarez-Carretero, Sandra et al. “A species-level timeline of mammal evolution integrating phylogenomic data.” Nature 602:263–267. 2022. A Bayesian timetree of 4,705 mammal species, combining 72 genomes with a large sequence alignment and fossil calibrations, by the group behind MCMCTree. Cited here by section because the copy read is the authors' accepted manuscript.
- 5.
Springer, Mark S. et al. “Placental mammal diversification and the Cretaceous–Tertiary boundary.” Proceedings of the National Academy of Sciences 100(3):1056–1061. 2003. An early relaxed-clock study of placental mammals on a large nuclear and mitochondrial dataset, dating the base of the placental tree to the Cretaceous.
- 6.
Wu, Ruolin et al. “Integrated analysis of fossils and molecular divergence time estimates a latest Jurassic origin of angiosperms.” Nature Plants 12:1242–1251. 2026. A dating of 644 flowering-plant species from 83 genes in which calibration bounds for 110 nodes were built from a statistical model of 25,685 fossil occurrences, compared with conventional bounds. It is by the Bristol and Gothenburg groups on both sides of the fossil-only dispute.
- 6a.
Page 1243: “Disregarding variations in sampling rates among clades by aggregating all angiosperm fossils results in an estimate for the origin of crown angiosperms that ranges from 150 to 133 Ma (Kimmeridgian–Valanginian; mean of 141 Ma)”. ↩ ↩
- 6b.
Page 1246, figure 3: Crown angiosperms, 95% HPD by calibration strategy: hard bounds 212.30–261.26 Ma; soft bounds 158.45–172.53 Ma; fossil-model densities 151.46–152.99 Ma. ↩ ↩ ↩ ↩ ↩ ↩
- 6c.
Page 1245: “Analyses using calibration strategy 3 estimated crown angiosperms to have originated in the Late Jurassic (152.99–151.46 Ma)”. ↩
- 6d.
Pages 1243–1244: “We excluded all pollen records from our analysis because they are difficult to attribute consistently to family-level clades within angiosperms”. ↩
- 6a.
- 7.
Benton, Michael J., and Philip C. J. Donoghue. “Paleontological Evidence to Date the Tree of Life.” Molecular Biology and Evolution 24(1):26–53. 2007. A review by two Bristol palaeontologists that sets out how fossils should be used to calibrate molecular clocks and supplies justified minimum and maximum ages for 30 key nodes across the tree of life. It argues that fossils give firm minimum ages and only loose maximum ages.
- 8.
Sauquet, Hervé et al. “What is the age of flowering plants?” Journal of Experimental Botany 73(12):3840–3853. 2022. A review by three plant systematists, two of whom have published flowering-plant timescales themselves, of why estimates of the crown age range so widely. It argues that the estimates track their priors.
- 8a.
Section Abstract: “Recent studies have reported a perplexing range of estimates for the crown-group age of angiosperms, from ~140 million years (Ma; Early Cretaceous) to 270 Ma (Permian).” “fossil-calibrated molecular dating estimates of the crown-group age of angiosperms have little to do with either the amount of molecular data or the number of internal fossil calibrations included. Instead, we argue that this age is almost entirely conditioned by its own prior distribution”. “contrasting age estimates in the literature mostly reflect conflicting prior distributions.” ↩ ↩ ↩
- 8a.
- 9.
Coiro, Mario et al. “How deep is the conflict between molecular and fossil evidence on the age of angiosperms?” New Phytologist 223(1):83–99. 2019. A review by three palaeobotanists of the fossil evidence on when flowering plants arose, set against molecular clock dates. It reads the order and spread of early pollen types, weighs every claimed pre-Cretaceous flowering plant, and examines known biases in clock dating.
- 9a.
Section Abstract: “However, the record may allow the Late Jurassic existence of ecologically restricted angiosperms, like those seen in the basal ANITA grade.” ↩ ↩
- 9b.
Section Abstract: “Pollen floras document a Northern Gondwanan appearance of monosulcate angiosperms in the Valanginian and subsequent poleward spread of monosulcates and tricolpate eudicots, accelerating in the Albian. The sequence of pollen types agrees with molecular phylogenetic inferences on the course of pollen evolution, but it conflicts strongly with Triassic and early Jurassic molecular ages, and the discrepancy is difficult to explain by geographic or taphonomic biases. Critical scrutiny shows that supposed pre-Cretaceous angiosperms either represent other plant groups or lack features that might confidently assign them to the angiosperms.” ↩ ↩ ↩ ↩ ↩ ↩ ↩
- 9a.
- 10.
Lee, Michael S. Y. et al. “Rates of phenotypic and genomic evolution during the Cambrian explosion.” Current Biology 23(19):1889–1895. 2013. A clock study of arthropods that estimated rates of anatomical and of molecular change through time from a large anatomical and genomic dataset. It is the clearest measurement of a rate burst during a radiation.
- 10a.
Section Abstract: “Assuming an Ediacaran origin for arthropods, phenotypic evolution was ~4 times faster, and molecular evolution ~5.5 times faster, during the Cambrian explosion compared to all subsequent parts of the Phanerozoic.” “these fast early rates do not change substantially even if the radiation of arthropods is compressed entirely into the Cambrian (~542 mega-annum [Ma]) or telescoped into the Cryogenian (~650 Ma).” ↩ ↩ ↩
- 10a.
- 11.
Berv, Jacob S., and Daniel J. Field. “Genomic Signature of an Avian Lilliput Effect across the K-Pg Extinction.” Systematic Biology 67(1):1–13. 2018. A study of rate variation across a genome-scale bird dataset that ties substitution rates to body size and metabolic rate, and argues that birds shrank across the end-Cretaceous extinction, speeding up their molecular rates and making clocks date the bird radiation too old. Mirror: PMC
- 11a.
Section Materials and Methods: “Despite lengthy attempts to reanalyze the avian evolutionary timescale in Coevol 1.4b to directly test for lineage-specific rate accelerations, these analyses failed to converge”. ↩ ↩
- 11b.
Section Abstract: “This “hidden” rate acceleration may result in both strict and relaxed molecular clocks over-estimating the age of the avian crown group”. ↩ ↩
- 11c.
Section Discussion: “Prum et al. (2015) employed an informative but soft-bound prior on the root of Neornithes, reflecting the conspicuous absence of avian crown group fossils in the Late Cretaceous Niobrara Formation.” “Applying this prior compresses the age of the neornithine MRCA by 30 Ma”. ↩
- 11d.
Section Results: “The Neornithes MRCA (median) was reconstructed as 553 g in the upper bound fossil calibrated analysis ( 617 g in uncalibrated), 50% smaller than our median body size estimate for the pre-K-Pg assemblage.” The pre-boundary assemblage is a fossil bird fauna from the last stage of the Cretaceous, described by Longrich and colleagues in 2011. ↩ ↩
- 11e.
Section Introduction: “The Mesozoic fossil record of crownward stem birds argues against the up to 100 million-year missing fossil record advocated by unconstrained divergence time analyses without a maximum age prior on the root”. ↩ ↩
- 11a.
- 12.
Phillips, Matthew J. “Geomolecular dating and the origin of placental mammals.” Systematic Biology 65(3):546–557. 2016. A test of placental rate models against fossil calibrations, followed by a dating with tight, conservative bounds. It found the common models misspecified and its own dates younger than most.
- 12a.
Section Abstract: “reveal substantial misspecification associated with life history rate correlates”. “divergences deep in the tree will tend to be inflated by the interaction of rate errors and asymmetric confidence in minimum and maximum bounds”. “The resulting divergence estimates are younger than others published recently, and provide the long-anticipated molecular signature for the placental mammal radiation observed in the fossil record near the 66 Ma Cretaceous-Paleogene extinction event.” ↩ ↩ ↩ ↩
- 12a.
- 13.
Tomkins, Jeffrey P., and Jerry Bergman. “Evolutionary molecular genetic clocks: a perpetual exercise in futility and failure.” Journal of Creation 29(2):26–35. 2015. The fullest young-earth critique of molecular clocks as a method, by a geneticist at the Institute for Creation Research and a biologist. It surveys rate differences between genes and lineages, clock dates that disagree with fossils, and argues that calibration is circular.
- 13a.
Page 26, abstract: “Since its first use in the early 1960s, molecular genetic clock methodologies assume evolution and deep-time calibrations taken from paleontology.” “3) divergence dates commonly disagree with paleontology despite being calibrated by it.” ↩ ↩
- 13b.
Page 28: “With either method of determining the genetic clock rate, the prospects of achieving any type of evolutionary concordance has been dismal.” ↩
- 13a.
- 14.
Jeanson, Nathaniel T. Replacing Darwin: The New Origin of Species. Master Books. 2017. A book by a young-earth creationist biologist with a Harvard doctorate proposing a creation model of species origins built on genetic clocks. Its chapters on humans and chimpanzees conclude that mitochondrial differences arose by mutation over 6,000 years while most nuclear differences between species were created.
- 15.
Carter, Robert W. “Genealogical vs phylogenetic mutation rates: Answering a challenge.” Proceedings of the Ninth International Conference on Creationism, 168–180. Cedarville University International Conference on Creationism. Edited by J. H. Whitmore. 2023. A young-earth paper by a geneticist at Creation Ministries International arguing that mutation rates measured in families are the true rates and that selection and drift cannot explain why long-term rates are slower. It models the question with the program Mendel's Accountant.
- 16.
Parham, James F. et al. “Best Practices for Justifying Fossil Calibrations.” Systematic Biology 61(2):346–359. 2012. A consensus statement by 25 palaeontologists and molecular biologists that sets five requirements a fossil calibration should meet, from museum numbers to an explained numeric age. It reports that most published calibrations do not meet them.
- 16a.
Page 347: “An appropriately constructed fossil calibration uses the oldest assigned fossil of a taxon as the basis for its minimum age”. ↩
- 16a.
- 17.
Ayala, Francisco J. et al. “Origin of the metazoan phyla: molecular clocks confirm paleontological estimates.” Proceedings of the National Academy of Sciences 95(2):606–611. 1998. A clock analysis of 18 protein-coding genes that answered the billion-year animal dates of the mid-1990s, arguing that slower vertebrate rates had inflated them. It is the earliest study in this article's animal table, and its authors read it as agreeing with the fossils. Mirror: PMC
- 18.
Peterson, Kevin J. et al. “Estimating metazoan divergence times with a molecular clock.” Proceedings of the National Academy of Sciences 101(17):6536–6541. 2004. A clock study of animal origins from seven nuclear genes with invertebrate fossil calibrations, which argues that earlier billion-year dates came from extrapolating slow vertebrate rates. Mirror: PMC
- 18a.
Section Abstract: “The earliest unequivocally bilaterian fossils are ≈555 million years old.” ↩ ↩
- 18b.
Section Abstract: “we estimate that the last common ancestor of bilaterians arose somewhere between 573 and 656 Ma, depending on the value assigned to the parameter scaling molecular substitution rate heterogeneity”. ↩
- 18c.
Section Discussion: “This cryptic precambrian history suggests that these stem-group bilaterians must have been “micrometazoans,””. ↩ ↩
- 18a.
- 19.
Peterson, Kevin J. et al. “The Ediacaran emergence of bilaterians: congruence between the genetic and the geological fossil records.” Philosophical Transactions of the Royal Society B 363(1496):1435–1443. 2008. A clock study of animal origins with revised calibrations, by a molecular palaeobiologist and an Ediacaran palaeontologist among others. It argued that bilaterians arose in the Ediacaran, and showed that dropping one fossil maximum moved crown animals from 766 to 904 Ma. Mirror: PMC
- 20.
Erwin, Douglas H. et al. “The Cambrian Conundrum: Early Divergence and Later Ecological Success in the Early History of Animals.” Science 334(6059):1091–1097. 2011. A combined fossil, molecular-clock and developmental study that dates 118 animal taxa from seven housekeeping genes with 24 fossil calibrations and argues that most Precambrian animals were stem groups of the living phyla.
- 20a.
Page 1092: “We estimate that the last common ancestor of all living animals arose nearly 800 Ma and that the stem lineages leading to most extant phyla had evolved by the end of the Ediacaran (541 Ma).” ↩
- 20a.
- 21.
Lee, Michael S. Y. “Molecular clock calibrations and metazoan divergence dates.” Journal of Molecular Evolution 49(3):385–391. 1999. A reanalysis of an earlier clock study that had put animal origins far back in the Precambrian, showing that it rested on a single fossil calibration and recalibrating it with several. The recalibrated dates came much closer to the Cambrian.
- 21a.
Section Abstract: “Recalibration of the protein data using these fossil dates indicates that metazoans diverged between 791 and 528 million years ago, a result broadly consistent with the palaeontological documentation of the "Cambrian explosion."” ↩
- 21a.
- 22.
Mángano, M. Gabriela, and Luis A. Buatois. “Decoupling of body-plan diversification and ecological structuring during the Ediacaran–Cambrian transition: evolutionary and geobiological feedbacks.” Proceedings of the Royal Society B 281:20140038. 2014. A research article that built a worldwide database of trace fossils (the tracks, burrows and trails that animals leave in sediment) from 369 rock units. Its authors found that body-plan diversification was complete by the Fortunian stage, while the restructuring of seafloor ecology came in Cambrian Stage 2. The article uses it as the one line of evidence that needs no exceptional preservation. Mirror: PMC
- 22a.
Section Abstract: “Trace-fossil information strongly supports the Cambrian explosion, but allows for a short time of phylogenetic fuse during the terminal Ediacaran–Fortunian”. ↩
- 22b.
Page 2: The record of animal–sediment interactions. “is far more continuous than the body-fossil record, particularly with respect to that of soft-bodied organisms”. Their figure caption makes the same point about the contrast with fossil Lagerstätten. ↩
- 22a.
- 23.
Springer, Mark S. et al. “Evolutionary models for the diversification of placental mammals across the KPg boundary.” Frontiers in Genetics 10:1241. 2019. A review by clock workers of the five models of placental origins, with the molecular and fossil evidence for each and a reanalysis of the largest anatomical matrix. Mirror: PMC
- 23a.
Section Abstract: “Archibald and Deutschman described three competing models for the diversification of placental mammals, which are the Explosive, Long Fuse, and Short Fuse Models.” ↩
- 23b.
Section Abstract: “The Long Fuse Model has received more support from relaxed clock studies than any of the other models, but this model is not supported by morphological cladistic studies that position Cretaceous eutherians outside of crown Placentalia.” ↩
- 23a.
- 24.
O'Leary, Maureen A. et al. “The Placental Mammal Ancestor and the Post–K-Pg Radiation of Placentals.” Science 339(6120):662–667. 2013. A study scoring 4,541 anatomical and other observable characters for 86 fossil and living species together with molecular data, which places the origin of crown placentals just after the end-Cretaceous extinction from the fossils.
- 24a.
Page 662, abstract: “Placentalia is just younger than the K-Pg boundary, or ~36 My younger than molecular clock–based mean estimates”. Page 662: “Ghost lineage estimates are minimum divergence dates and may underestimate the timing of actual splits.” ↩ ↩
- 24b.
Page 665: “We recognize Protungulatum donnae as the oldest undisputed species within crown Placentalia”. Table 1: “64.85”. ↩ ↩
- 24a.
- 25.
Meredith, Robert W. et al. “Impacts of the Cretaceous Terrestrial Revolution and KPg extinction on mammal diversification.” Science 334(6055):521–524. 2011. A family-level mammal tree from a large molecular supermatrix, dated with relaxed clocks and many fossil calibrations. It is a standard source for the long-fuse placental timescale.
- 25a.
Section Abstract: “Relaxed clock analyses support the long-fuse model of diversification and highlight the importance of including multiple fossil calibrations that are spread across the tree.” ↩
- 25a.
- 26.
dos Reis, Mario et al. “Phylogenomic datasets provide both precision and accuracy in estimating the timescale of placental mammal phylogeny.” Proceedings of the Royal Society B 279(1742):3491–3500. 2012. A Bayesian clock study of 36 nuclear genomes and 274 mitochondrial genomes of mammals, 20.6 million base pairs in all, with fossil calibrations.
- 26a.
Section Abstract: “Placentalia diverged 88–90 Ma”. ↩
- 26a.
- 27.
Carlisle, Emily et al. “A timescale for placental mammal diversification based on Bayesian modeling of the fossil record.” Current Biology 33(15):3073–3082. 2023. A fossil-only estimate of when placental mammals and their main groups arose, from a Bayesian model of fossil occurrences with no molecular data.
- 27a.
Section Results: “although Placentalia likely originated in the Late Cretaceous, the majority of orders likely originated around or after the K-Pg boundary”. ↩
- 27b.
Section Results: “Out of 23 clades from recent molecular clock analyses (Figure 6), 12 have overlapping credible intervals compared with the total group BBB analysis”. “Of the 10 that do not overlap, many are estimated to be older under BBB model”. ↩
- 27c.
Section Results: “the root age is estimated to be 77–70 millions of years before present (Ma), whereas without Deccanolestes, this drops to 70.2–66 Ma.” With the Indian fossil Deccanolestes counted as a placental relative for the first figure. ↩
- 27a.
- 28.
Field, Daniel J. et al. “Late Cretaceous neornithine from Europe illuminates the origins of crown birds.” Nature 579:397–401. 2020. The description of Asteriornis maastrichtensis, a nearly complete skull from Belgium that is one of the only well-supported crown birds from the age of dinosaurs.
- 29.
Brown, Joseph W. et al. “Strong mitochondrial DNA support for a Cretaceous origin of modern avian lineages.” BMC Biology 6:6. 2008. A dating of modern birds from whole mitochondrial genomes under several dating programs and settings, built to test whether the bird orders arose before or after the end-Cretaceous extinction. All of its methods put the root of modern birds deep in the Cretaceous. Mirror: PMC
- 29a.
Table 3: “A Neognaths-Paleognaths 125.7 (12.4) 131.1 (10.7) 114.3 (6.9) 102.8 (6.1) 129.9 (11.0) 132.4 (10.7) 132.9 (11.6) 133.2 (8.1)”. Root ages in Ma under r8s, PATHd8, Multidivtime, Dating5 and BEAST settings, with standard deviations in brackets. ↩
- 29a.
- 30.
Jarvis, Erich D. et al. “Whole-genome analyses resolve early branches in the tree of life of modern birds.” Science 346(6215):1320–1331. 2014. The first whole-genome tree of the orders of modern birds, dated with MCMCTree and fossil calibrations. Mirror: PMC
- 30a.
Section Dating the radiation of Neoaves: “Our results suggest that after the Palaeogna-thae and Neognathae divergence about 100 Ma in the Late Cretaceous”. ↩
- 30a.
- 31.
Kuhl, Heiner et al. “An unbiased molecular approach using 3′-UTRs resolves the avian family-level tree of life.” Molecular Biology and Evolution 38(1):108–127. 2021. A family-level bird tree from noncoding gene ends in 429 species, dated with fossil calibrations. Its root and deep splits came out much older than the 2015 study that placed modern birds after the extinction. Mirror: PMC
- 31a.
Section Time Calibration of the Family-Level Phylogeny: “shows divergence dates for Palaeognathae and Neognathae (94 Ma) and Galloanserae and Neoaves (81 Ma) that are much earlier than those suggested previously ( Prum et al. 2015 ).” ↩
- 31b.
Section Time Calibration of the Family-Level Phylogeny: “the estimated divergence time of Galliformes and Anseriformes of 62.5 Ma (million years ago) fits well with a recently reported Mesozoic fossil (66.7 Ma) that is close to the last common ancestor of Galloanserae”. ↩ ↩
- 31a.
- 32.
Stiller, Josefin et al. “Complexity of avian evolution revealed by family-level genomes.” Nature 629:851–857. 2024. A tree of 363 bird species built from whole genomes and dated with fossil calibrations.
- 32a.
Page 853: “In contrast to a recent study proposing a diversification of Neoaves during the Upper Cretaceous11, we found that the early divergences in Neoaves were tightly associated with the K–Pg boundary”. “Only two divergences occurred before the boundary”. ↩
- 32b.
Section Results: “Only two divergences occurred before the boundary: Mirandornithes diverged from the remaining Neoaves 67.4 Ma (95% credible interval 66.2–68.9) and Columbaves diverged 66.5 Ma (95% credible interval 65.2–67.9). All subsequent divergences postdate the bound”. ↩
- 32a.
- 33.
Wu, Shaoyuan et al. “Genomes, fossils, and the concurrent rise of modern birds and flowering plants in the Late Cretaceous.” Proceedings of the National Academy of Sciences 121(8):e2319696121. 2024. A genome-scale bird tree dated with 20 fossil constraints, which put most bird orders in the Late Cretaceous. It is the study that Stiller and colleagues answer in the same year. Mirror: PMC
- 33a.
Section Results: “we assigned the age of the earliest known member of crown Aves, Asteriornis maastrichtensis (66.7 Ma) ( 19 ), as the minimum bound and the age of the earliest known member of Ichthyornithes, Ichthyornis dispar (94.3 Ma) ( 20 ), as the maximum bound”. ↩
- 33b.
Section Results: “Our results show that interordinal diversification of modern birds largely occurred in the Late Cretaceous and underwent a continuous radiation across the KPg boundary. Based on the lower level of 95% CIs, there are approximately 15 avian orders that likely originated in the Paleogene, whereas the origin of the remaining 24 orders falls within the Late Cretaceous”. ↩ ↩
- 33a.
- 34.
Silvestro, Daniele et al. “Fossil data support a pre-Cretaceous origin of flowering plants.” Nature Ecology & Evolution 5:449–457. 2021. A Bayesian model of the fossil record of 198 flowering plant families, using no molecular data, to estimate when each family and the crown group arose. Cited here from the authors' accepted manuscript.
- 35.
Smith, Stephen A. et al. “An uncorrelated relaxed-clock analysis suggests an earlier origin for flowering plants.” Proceedings of the National Academy of Sciences 107(13):5897–5902. 2010. A relaxed-clock dating of land plants with fossil calibrations that put the origin of crown flowering plants in the Late Triassic.
- 35a.
Section Results: “We estimated the origin of crown angiosperms to be 217 Myr (95% HPD: 182–257 Myr), in the Late Triassic.” ↩
- 35a.
- 36.
Magallón, Susana et al. “A metacalibrated time-tree documents the early rise of flowering plant phylogenetic diversity.” New Phytologist 207(2):437–453. 2015. A dating of flowering plants sampling 87% of families with 137 fossil calibrations, in which the crown age was bounded by an interval computed from the group's fossil record. It is the main clock study to land the crown in the Early Cretaceous.
- 36a.
Section Abstract: “While available molecular clock estimates of angiosperm age range from the Paleozoic to the Late Cretaceous, the fossil record is consistent with angiosperm diversification in the Early Cretaceous.” “The angiosperm crown age was bound within a confidence interval calculated with a method that considers the fossil record of the group. An Early Cretaceous crown angiosperm age was estimated with high confidence.” ↩ ↩
- 36a.
- 37.
Barba-Montoya, Jose et al. “Constraining uncertainty in the timescale of angiosperm evolution and the veracity of a Cretaceous Terrestrial Revolution.” New Phytologist 218(2):819–834. 2018. A Bayesian dating of flowering plants from 83 genes in 644 species under several calibration strategies, reporting how much the strategy moves the origin.
- 37a.
Table 1: “This study (composite) Loci: 77‐plastid, 4‐mt, 2‐nuc. Taxa: 644. Calib: 52./IR 149–256 Ma”. ↩
- 37a.
- 38.
Budd, Graham E. et al. “Fossil data do not support a long pre-Cretaceous history of flowering plants.” bioRxiv. 2021. A critique of the fossil-only model behind the pre-Cretaceous result for flowering plants, arguing that its answer follows from the age of the oldest fossil and the number of fossils rather than from the data. ↩
- 39.
Morris, Jennifer L. et al. “The timescale of early land plant evolution.” Proceedings of the National Academy of Sciences 115(10):E2274–E2283. 2018. A Bayesian dating of land plants with 37 fossil calibrations, run across seven possible trees and several calibration strategies. Its authors include palaeobotanists who study the earliest spores, and they compare the result with the spore and body-fossil records directly. Mirror: PMC
- 39a.
Section Comparisons with the Fossil Record: “The first unequivocal embryophyte body fossil taxon, Cooksonia cf. pertoni, appears in the Wenlock”. ↩
- 39b.
Section Discussion: “the origin of crown embryophytes is dated to 515.1 Ma to 470.0 Ma (middle Cambrian–Early Ordovician). However, all of the mean estimated ages are resolved within the Phanerozoic across all alternative topologies and dating strategies, and the majority are dated to around 500 Ma”. ↩
- 39c.
Section Comparisons with the Fossil Record: “The earliest known fossils of embryophyte affinity are permanently fused tetrahedral tetrad cryptospores”. “from the Middle Ordovician [Dapingian; 469 Ma (50)]. Cryptospores of unclear affinity from the Cambrian [sensu stricto Strother (51)], while not considered unequivocally embryophyte, informed our soft maximum constraint (515.5 Ma).” ↩ ↩
- 39a.
- 40.
Springer, Mark S. et al. “Macroevolutionary dynamics and historical biogeography of primate diversification inferred from a species supermatrix.” PLoS ONE 7(11):e49521. 2012. A dated tree of 367 primate species from a molecular supermatrix with hard and soft fossil bounds. Its crown primate date is younger than most earlier clock estimates.
- 41.
dos Reis, Mario et al. “Using phylogenomic data to explore the effects of relaxed clocks and calibration strategies on divergence time estimation: primates as a test case.” Systematic Biology 67(4):594–615. 2018. A genome-scale dating of 372 primate species under two clock models and two calibration strategies, written partly to show how much those choices move the dates. It also reviews five decades of primate clock estimates.
- 41a.
Section Results: “Under calibration strategy A and the AR model, we find that crown Primates originated 79.2–70.0 million years ago (Ma), before the K–Pg event at 66 Ma.” ↩
- 41b.
Section Appendix 1: “the basal primate Altiatlasius from Ouarzazate in Morocco which is considered to represent the late Paleocene (Thanetian) and dating to around 58”. The paper's calibration table gives 55.6 Ma for the same fossil. ↩
- 41c.
Section Five Decades of Primate Molecular Timetrees: “Prior to the study reported here, the estimated age of the living primate clade has spanned a 30 My differential, ranging from as young as 55 Ma (Koop et al. 1989) to as old as 87 Ma (Perelman et al. 2011).” ↩
- 41a.
- 42.
Parfrey, Laura Wegener et al. “Estimating the timing of early eukaryotic diversification with multigene molecular clocks.” Proceedings of the National Academy of Sciences 108(33):13624–13629. 2011. A relaxed-clock dating of the eukaryote tree from 15 proteins with 23 fossil calibrations, by a team that includes the palaeontologist Andrew Knoll. It compares its result with the oldest microfossils thought to be eukaryotic. Mirror: PMC
- 43.
Betts, Holly C. et al. “Integrated genomic and fossil evidence illuminates life's early evolution and eukaryote origin.” Nature Ecology & Evolution 2(10):1556–1562. 2018. A dated tree of all life from conserved genes with fossil calibrations, run under several priors and clock models. It is used here for its interval on the last common ancestor of living eukaryotes and for its choice of the oldest crown-eukaryote fossil. Mirror: PMC
- 44.
Gibson, Timothy M. et al. “Precise age of Bangiomorpha pubescens dates the origin of eukaryotic photosynthesis.” Geology 46(2):135–138. 2018. A rhenium-osmium dating of the shales above and below the red alga Bangiomorpha on Baffin Island, the oldest fossil assigned to a living eukaryote group. It made the fossil much younger than had been assumed, which changes every clock calibrated on it.
- 45.
Ni, Xijun et al. “The oldest known primate skeleton and early haplorhine evolution.” Nature 498(7452):60–64. 2013. The description of Archicebus achilles, a nearly complete skeleton from the early Eocene of China placed at the base of the tarsier line.
- 45a.
Page 60, abstract: “a nearly complete and partly articulated skeleton of a primitive haplorhine primate from the early Eocene of China, about 55 million years ago, the oldest fossil primate of this quality ever recovered.” ↩
- 45a.
- 46.
dos Reis, Mario et al. “Bayesian molecular clock dating of species divergences in the genomics era.” Nature Reviews Genetics 17(2):71–80. 2016. A review of molecular-clock dating by three of its leading practitioners, two of whom wrote MCMCTree and much of the theory of Bayesian dating. It traces the method from Zuckerkandl and Pauling to genome-scale analyses, explains why sequence data cannot separate rate from time, and lists the main programs. Cited here by section because the copy read is the authors' final manuscript.
- 46a.
Section Introduction: “These studies generated considerable controversy because the clock estimates were much older than the dates suggested by the fossil record, sometimes twice as old”. ↩
- 46a.
- 47.
Beck, Robin M. D., and Michael S. Y. Lee. “Ancient dates or accelerated rates? Morphological clocks and the antiquity of placental mammals.” Proceedings of the Royal Society B 281:20141278. 2014. A tip-dating study that ran relaxed clocks on a large anatomical matrix of mammals, alone and with an 8.5-kilobase nuclear dataset. Its dates for the first placental split came out older than every molecular and fossil estimate, which the authors read as a warning about the clock models rather than as a true age. Mirror: PMC
- 47a.
Section Abstract: “retrieve implausibly ancient, Late Jurassic–Early Cretaceous estimates for the initial diversification of Placentalia (crown-group Eutheria). These dates are much older than all recent molecular and palaeontological estimates.” “relatively minor deviations in rate patterns can generate large effects on estimated divergence dates.” ↩ ↩
- 47a.
- 48.
Puttick, Mark N. et al. “Dating placentalia: morphological clocks fail to close the molecular fossil gap.” Evolution 70(4):873–886. 2016. A tip-dating study of placental mammals on an osteological matrix of 3,660 characters, comparing tree priors including the fossilized birth-death process. It found tip dates older than both the fossils and the molecular clocks. Mirror: PMC ↩
- 48a.
Section Abstract: “We find that the estimated date for the origin of crown mammals is much older, ∼130–145 million years ago (Ma), than fossil and molecular clock data (∼80–90 Ma).” The paper is about crown placentals, as its title says. ↩
- 48a.
- 49.
The gaps, trends, spreads, floor ratios and rank correlations in this article are computed by
research/gap-pricing.tsfrom the intervals printed in the sources cited in Parts 2 to 6, each against the first-fossil age its own literature prints: 575 Ma for crown animals,1 555 for bilaterians,18 64.85 for placentals,24 66.7 for crown birds28 and 135 for flowering plants.34 The trend is an ordinary least-squares slope of midpoint gap on publication year; the precision floor is half of dos Reis and Yang's 0.322 t; the preservation ranks are this article's. The script has self-tests and a negative control. All figures are the mainstream's, on its timescale. ↩ ↩ ↩ - 50.
dos Reis, Mario, and Ziheng Yang. “The unbearable uncertainty of Bayesian divergence time estimation.” Journal of Systematics and Evolution 51(1):30–43. 2013. A methods paper showing, with theory and a genome-scale primate example, that Bayesian date estimates do not converge on the true dates as sequence data grow. Their error falls only to a floor proportional to the age of each node, set by the uncertainty of the fossil calibrations.
- 50a.
Page 31, Figure 1 legend: “for every one million years of divergence, 0.322 million years are added to the CI width of time estimates.” Page 31: “nearly all the uncertainty in the posterior time estimate is due to uncertainties in the fossil calibrations, and adding more sequence data is unlikely to produce more precise estimates.” ↩
- 50a.
- 51.
780 Myr over 61 ghost lineages is 12.8 Myr per lineage; the abstract's 720 Myr gives 11.8. Ghost lineages are minimums, so the true average is larger. ↩
- 52.
Warnock, Rachel C. M. et al. “Exploring uncertainty in the calibration of the molecular clock.” Biology Letters 8(1):156–159. 2012. A test of how the shape of the probability curve used for fossil calibrations changes Bayesian dates, run on a vertebrate dataset in the two main programs, BEAST and MCMCTree. It shows that the programs truncate the curves users specify, and that dates swing widely with the curve's parameters when only a minimum age is given.
- 52a.
Page 157: “It will surprise some to discover that user-specified (initial) prior probabilities on the timing of clade divergence are not the effective priors implemented in calculating the posterior time estimates”. “changing the mean or the standard deviation of the lognormal distribution can cause the mean divergence estimates, as well as lower and upper 95% posterior intervals, to differ by hundreds of millions of years”. ↩
- 52a.
- 53.
Rodríguez-Trelles, Francisco et al. “A methodological bias toward overestimation of molecular evolutionary time scales.” Proceedings of the National Academy of Sciences 99(12):8112–8115. 2002. A short methods paper arguing that the statistics of rate and date estimation push clock dates older than the truth.
- 53a.
Page 8112, abstract: “This introduces a bias toward an overestimation of time since divergence”. ↩
- 53a.
- 54.
Thomas, Brian. “Cambrian Clash: Fossils and Molecular Clocks Disagree.” Institute for Creation Research. Oct 20, 2008. A news article by a science writer at the Institute for Creation Research on Jeffrey Levinton's 2008 review of the Cambrian explosion. It gives the young-earth reading of the clock-fossil gap for animals.
- 54a.
Paragraph 4: “The estimates range from 586 to over 1200 million years!” “Either the molecular clocks are wrong, the rock dates are inaccurate, or both.” Paragraph 5: “creation scientists would suggest that assumptions of evolution may influence both of them too strongly to allow any confidence in either.” ↩ ↩
- 54a.
- 55.
Oard, Michael J. “More expansion of fossil time ranges.” Journal of Creation 33(3):3–4. 2019. A short news column by a young-earth geologist collecting recent finds that extend fossil ranges downward, read as evidence that the fossil record is broader than evolutionary accounts expected.
- 55a.
Page 3: “The molecular clock often suggests that fossils should be found earlier, but regardless pushing chrysophytes back 120 million evolutionary years still shows that the fossil record is broader than they earlier thought.” ↩
- 55a.
- 56.
Brand, Leonard, and Arthur Chadwick. Faith, Reason, & Earth History: A Paradigm of Earth and Biological Origins by Intelligent Design. Andrews University Press, Berrien Springs. 3rd edition. 2016. A college textbook by two young-earth scientists, a biologist and a geologist, who have published taphonomic field work in secular journals. Its chapter on a biblical model of earth history explains the order of first appearances as the destruction of habitats in turn during the Flood, and states plainly which observations the model does not yet explain.
- 56a.
Page 376: “A perfect angiosperm flower has been found in the Jurassic.” Their note cites Liu and Wang 2015 in Historical Biology. ↩
- 56b.
Page 377: “These changes in the fossils do not represent the appearance of newly created organisms. They resulted from the successive final destruction of some habitats and the death and burial of organisms from each of those habitats as the catastrophe unfolds.” ↩
- 56c.
Page 375: “why is there no flowering plant pollen below the Middle Cretaceous? Pollen is easily carried long distances by wind or water. Why are there not at least a few mice or sparrows in Paleozoic or Mesozoic deposits?” “Our current understanding of this model leaves many unanswered questions. But in broad strokes, it offers a beginning on which to build better answers.” ↩
- 56d.
Page 376: “Perhaps these groups were largely inhabitants of upland regions that were not affected until late in the geological catastrophe.” The groups are birds, mammals and flowering plants. ↩
- 56a.
- 57.
Wise, Kurt P. Faith, Form, and Time: What the Bible Teaches and Science Confirms about Creation and the Age of the Universe. Broadman & Holman, Nashville. 2002. A book for general readers by a young-earth palaeontologist with a Harvard doctorate. Its chapter on the pre-Flood world proposes separate biomes whose burial in turn would give the order of the fossil record.
- 57a.
Chapter The Antediluvian World: “It is also likely that the gymnosperm-dinosaur biome was located at a lower altitude or closer to the shore of the antediluvian world. This would explain why members of the gymnosperm-dinosaur biome are consistently buried beneath members of the other biome.” ↩
- 57a.
- 58.
Meyer, Stephen C. Darwin's Doubt: The Explosive Origin of Animal Life and the Case for Intelligent Design. HarperOne, New York. Paperback edition. 2014. Section Copyright page: “Copyright © 2013 by Stephen C. Meyer. Epilogue © 2014 by Stephen C. Meyer.” A book that argues for intelligent design from the Cambrian explosion, the sudden appearance of most animal body plans in the fossil record. The edition cited is the 2014 paperback, whose epilogue, answering critics of the 2013 first edition, runs to roughly 63,000 words and does not exist in the hardcover, so any claim that the book never mentions something must name its edition. In the book, Cloudina does not appear anywhere, including the bibliography; Tommotian and Atdabanian are absent from the first edition and each appears once, in one epilogue sentence; small shelly appears eight times in the whole book but not in the first-edition main text; and Manykaian appears only in endnote 39 to chapter 4. The article uses it as the book under review, since its main text and epilogue claim that the main pulse of Cambrian body-plan innovation occupied no more than six million years.
- 58a.
Chapter 5, section Deep Divergence: “Proponents of deep divergence don’t deny that the fossil evidence has come up short. Instead, they adopt one of the versions of the artifact hypothesis to account for that missing evidence.” ↩
- 58a.
- 59.
Wise, Kurt P., and Donna Richardson. “What Biostratigraphic Continuity Suggests about Earth History.” Proceedings of the Ninth International Conference on Creationism 9:611–625. 2023. A paper by Kurt Wise, a young-earth creationist with a Harvard paleontology doctorate, and Donna Richardson, doing original quantitative work on the same open database the order article uses. Their method: at each of 114 globally defined stratigraphic boundaries they computed the number of local stage-straddling species (NLSSS), meaning species occurring in the stage immediately below and immediately above a boundary within two degrees of latitude and longitude of each other, using the Paleobiology Database, a public fossil database, and they checked the counts at 1°, 3° and 4°. The order article calls it the strongest recent concession from inside the creationist movement, and recomputes its statistic in its Part 6 because the pre-Flood/Flood use does not survive.
- 59a.
Page 623, right column: “the global biostratigraphic column seems to be real.” Their conclusion on the column itself. ↩
- 59a.