The last common bilaterian ancestor
Open accessAbstract
Many regulatory genes appear to be utilized in at least superficially similar ways in the development of particular body parts in Drosophila and in chordates. These similarities have been widely interpreted as functional homologies, producing the conventional view of the last common protostome-deuterostome ancestor (PDA) as a complex organism that possessed some of the same body parts as modern bilaterians. Here we discuss an alternative view, in which the last common PDA had a less complex body plan than is frequently conceived. This reconstruction alters expectations for Neoproterozoic fossil remains that could illustrate the pathways of bilaterian evolution.
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Claims the paper makes
4 of 36 cited sources checked · 1 supported · 0 flagged · accuracy not yet measured
- Premisecitation
Molecular clock estimates mostly place the latest PDA much deeper in time than the latest Neoproterozoic, with dates ranging from 600 to 1200 Mya depending on proteins, calibration points, and substitution models.
“Depending on the proteins, calibration points and rate substitution models used, these estimates range from 600 to 1200 Mya (Wray et al., 1996;Ayala et al., 1998;Bromham et al., 1998;Gu, 1998;Cutler, 2000).”
Quote in the paperSubstantial claim1 of 3 sources checked- cites Wray et al., 1996 source
- cites Ayala et al., 1998 source
- cites Bromham et al., 1998 source
The Results report divergence estimates older than 680 Ma and say estimates differ between the two sequence datasets; the Methods also describe multiple fossil calibrations and a substitution model. But the paper does not give the attributed 600–1200 Ma range, and its estimates have wide confidence intervals rather than precise dates. The cited paper therefore supports the general point about older, method-dependent molecular dates, not the stated range. (LLM, conf=0.84)
Experimental — accuracy not yet measured
- Rests on an assumption: Molecular clock estimates, despite potential problems, are reliable enough to indicate that the latest PDA is much older than the latest Neoproterozoic.
- Premisecitation
There is clear evidence for the prevalence of sponges and cnidarians in the Doushantuo and Ediacaran assemblages.
“and indeed there is clear evidence for the prevalence of sponges and cnidarians from both the Doushantuo and Ediacaran assemblages (Gehling and Rigby, 1996;Brasier et al., 1997;Chen et al., 2000;Xiao et al., 2000;Chen et al., 2002).”
Quote in the paperNot a substantial claim1 of 3 sources checked- cites Gehling and Rigby, 1996 source
- cites Brasier et al., 1997
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- cites Chen et al., 2000 source
The Methods and Discussion report apparent cnidarian forms and microscopic sponges in Doushantuo samples, though the authors caution that the fossil interpretations are not certain. The paper does not establish their prevalence across the Doushantuo assemblage; its background discussion describes the affinities of Ediacaran remains as enigmatic rather than establishing a prevalence of sponges and cnidarians there. Page numbers were not available in our copy. (LLM, conf=0.86)
Experimental — accuracy not yet measured
- Premisecitation
Across Bilateria, cell type-specific gene expression in striated muscle cells is controlled by members of the MyoD family of bHLH transcriptional regulators and by regulators of the Mef2 class.
“Across the Bilateria, for example, cell type-specific gene expression in striated muscle cells is controlled by members of the MyoD family of bHLH transcriptional regulators and by regulators of the Mef2 class (Molkentin et al., 1995) (reviewed by Zhang et al., 1999).”
Quote in the paperSubstantial claim1 of 2 sources checked- cites Molkentin et al., 1995 source
- cites Zhang et al., 1999 source
The Abstract and Results discuss Hairy, Groucho, and dCtBP in early Drosophila embryos, not regulation of striated-muscle gene expression across Bilateria. Searches for MyoD, Mef2, and muscle-specific regulators found no passage addressing the attributed claim. (LLM, conf=0.77)
Experimental — accuracy not yet measured
- Findingreasoning
There is absence of evidence for macroscopic benthic adult bilaterian forms prior to 555 Mya.
“More compelling is the absence of any evidence for macroscopic benthic adult bilaterian forms prior to 555 Mya.”
Quote in the paperSubstantial claim - Premisecitation
The oldest convincing bilaterian remains are Kimberella and various bilaterian trace fossils, dating only to about 555 Mya.
“The oldest convincing remains of bilaterians are fossils of an already well-developed animal, Kimberella, which we discuss below, and of various bilaterian trace fossils.These date back only to about 555 Mya (Martin et al., 2000)”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Martin et al., 2000 source
- Premisenone given
There is an enigmatic lack of fossil evidence that can be tied to a likely PDA in rocks older than the Cambrian.
“but there is an enigmatic absence of fossil evidence that can be related to a likely PDA in older rocks.”
Quote in the paperSubstantial claim - Interpretationreasoning
This alternative reconstruction changes what should be expected from Neoproterozoic fossils that might document bilaterian evolution.
“This reconstruction alters expectations for Neoproterozoic fossil remains that could illustrate the pathways of bilaterian evolution.”
Quote in the paperSubstantial claim - Interpretationreasoning
The authors argue for an alternative reconstruction in which the last common PDA had a less complex body plan than commonly assumed.
“Here we discuss an alternative view, in which the last common PDA had a less complex body plan than is frequently conceived.”
- Also stated in: § INTRODUCTION · plain
“We propose an alternative view of the PDA, an animal much more simply constructed than is customary in current treatments.”
Quote in the paperSubstantial claim - Also stated in: § INTRODUCTION · plain
- Interpretationnone given
A morphologically complex PDA would very likely have left detectable traces in the relatively well-studied Neoproterozoic fossil record.
“Second, to put the matter in a nutshell, the paleontological problem is that a morphologically complex PDA would very likely have left traces of its presence in the relatively wellstudied fossil record of the Neoproterozoic.”
Quote in the paperNot a substantial claim- Rests on an assumption: The Neoproterozoic fossil record has been sampled well enough that a complex PDA would likely have been detected if it existed.
- Interpretationnone given
For many developmental regulatory systems, the evidence does not convincingly show that specific morphogenetic pathways themselves are conserved, as opposed to cell-type specification and differentiation.
“First, for many developmental regulatory systems there is no convincing evidence that it is specific morphogenetic pathways per se that are conserved, rather than cell-type specification and differentiation processes.”
Quote in the paperNot a substantial claim - Interpretationcitation
An alternative interpretation of the observations used to support morphogenetic conservation may apply to many of the most prominent examples.
“But there is an alternative interpretation for the observations used to support morphogenetic conservation, which may apply to many of the most prominent examples.”
Quote in the paperSubstantial claim - Interpretationcitation
If Kimberella is indeed an advanced protostomial animal, then the last common PDA must significantly antedate 555 Mya.
“first, if Kimberella is indeed an advanced protostomial animal (Fedonkin and Waggoner, 1997), then the last common PDA must significantly antedate 555 Mya”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Fedonkin and Waggoner, 1997 source
- Interpretationreasoning
The regulatory toolkit of Bilateria can construct all sorts of body plans, as shown by diverse usage.
“The regulatory 'toolkit' of the Bilateria contains elements that can be used to construct all sorts of body plans, as amply shown by their diverse current usage.”
Quote in the paperSubstantial claim - Interpretationreasoning
Under the revised topology, the last common PDA is also the last common bilaterian ancestor.
“For this discussion the most significant change is that the last common PDA is now also the last common bilaterian ancestor.”
Quote in the paperSubstantial claim - Interpretationreasoning
Genomic regulatory programs for cell differentiation predate clade-specific bilaterian morphogenetic pattern formation programs.
“In general, it may be concluded from the pleisiomorphy of conserved genes and cell differentiation states that genomic regulatory programs for cell differentiation predate any clade-specific bilaterian programs for morphogenetic pattern formation.”
Quote in the paperSubstantial claim - Interpretationreasoning
The PDA utilized the same homeodomain regulatory genes in initial CNS formulation as insects and vertebrates.
“We can also infer that the PDA utilized the same set of homeodomain regulatory genes in the initial embryonic formulation of its central nervous system as do both insects and vertebrates.”
Quote in the paperSubstantial claim - Interpretationreasoning
At least some differentiation gene batteries date to the last common ancestor of cnidarians and bilaterians.
“and thus at least some of these differentiation gene batteries date to the last common ancestor of cnidarians and bilaterians.”
Quote in the paperSubstantial claim - Interpretationcitation
Although heads, hearts, eyes, and other structures carry out analogous functions, their developmental morphogenesis and functional anatomies are not actually very similar when considered in detail.
“Although the heads, hearts, eyes, etc., of insects, vertebrates and other creatures carry out analogous functions, neither their developmental morphogenesis, nor their functional anatomies are actually very similar if considered in any detail.”
Quote in the paperSubstantial claim - Interpretationreasoning
Later in evolution, transcriptional regulators of differentiation gene batteries were coopted for increasingly complex, clade-specific programs of gene regulation controlling pattern formation.
“Later in evolution the transcriptional regulators of these differentiation gene batteries would have been coopted for use in increasingly complex, clade-specific programs of gene regulation that control pattern formation processes.”
Quote in the paperSubstantial claim - Interpretationreasoning
Other network subelements are common to all bilaterians and part of shared heritage from the last common bilaterian ancestor.
“Other classes of network subelement are also common to all bilaterians, and hence are part of the shared genomic regulatory heritage from the last common bilaterian ancestor.”
Quote in the paperSubstantial claim - Interpretationreasoning
These shared properties must have been attributes of the last common PDA.
“These shared properties must have been attributes of the last common PDA.”
Quote in the paperSubstantial claim - Interpretationreasoning
Regulatory processes underlying development of specialized differentiated cells are very old, conserved, plesiomorphic features.
“The regulatory processes that underlie development of specialized differentiated cells are indeed very old, conserved, plesiomorphic features.”
Quote in the paperSubstantial claim - Premisecitation
Vertebrates and insects use the same transcriptional regulators for differentiation of neuroblasts, motoneurons, pioneer neurons, and glial cells.
“they use the same transcriptional regulators for the differentiation of neuroblasts, motoneurons, pioneer neurons, and glial cells (reviewed by Chan and Jan, 1999;Arendt and Nübler-Jung, 1999)”
Quote in the paperSubstantial claim0 of 2 sources checked - Premisecitation
Body fossils and molecular biomarkers establish that numerous crown-group eukaryote lineages existed between 1200 and 740 Mya, including green and red algae, dinoflagellates, ciliates, and testate amoebae.
“Body fossils and evidence from molecular biomarkers has established the presence of numerous crown-group eukaryote lineages between 1200 Mya and 740 Mya, including green and red algae, dinoflagellates, ciliates and testate amoebae (which form fossilizable chambers) (Butterfield, 2000;Porter and Knoll, 2000).”
Quote in the paperNot a substantial claim0 of 2 sources checked- cites Butterfield, 2000
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- cites Porter and Knoll, 2000
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- Interpretationnone given
The progressive division of morphological space into transcriptional states is the fundamental mechanism by which bilaterians organize body parts and body plans.
“The programmed and progressive division of morphological space into diverse transcriptional states is the fundamental mechanism by which bilaterians organize the morphogenesis of all of their major body parts and thus their body plans during development.”
Quote in the paperNot a substantial claim - Interpretationreasoning
Evolution of adult body plans in the bilaterian stem group would have had to await more favorable late Neoproterozoic environments.
“Evolution of adult body plans in the bilaterian stem group would have had to await the more favorable late Neoproterozoic environments.”
Quote in the paperSubstantial claim - Interpretationreasoning
It cannot safely be assumed that the PDA had a full range of complex body parts homologous to extant Bilateria.
“it cannot safely be assumed that the PDA possessed a full range of complex body parts homologous to those of extant Bilateria.”
Quote in the paperSubstantial claim - Interpretationnone given
The fossil record shows a clear pattern of progressive appearance of more complex bilaterian lineages through the Early and Middle Cambrian.
“The fossil record reveals a clear pattern of progressive appearance of more complex bilaterian lineages through the Early and Middle Cambrian.”
Quote in the paperNot a substantial claim - Interpretationreasoning
The fossil record provides no constraints on when such an animal appeared because evidence is hard to find.
“The fossil record provides no constraints on when such an animal could have appeared because it will be hard to find palentological evidence of it.”
Quote in the paperSubstantial claim - Interpretationreasoning
Taking Drosophila and vertebrates as surrogates for Bilateria, the Bilateria clearly share detailed regulatory control systems for many specific differentiated cell types.
“Taking Drosophila and vertebrates as surrogates for Bilateria (this leaves aside Lophotrochozoa, for which there is as yet only relatively sparse molecular biological evidence), the Bilateria clearly share detailed regulatory control systems for many specific differentiated cell types.”
Quote in the paperSubstantial claim- The authors state this as an assumption: Drosophila and vertebrates are adequate surrogates for Bilateria in comparative analysis of conserved regulatory systems.
- Interpretationreasoning
Comparative developmental data do not require a morphologically complex PDA.
“But the comparative developmental data do not require a morphologically complex PDA.”
Quote in the paperSubstantial claim - Interpretationreasoning
Some likely PDA properties are implied by general bilaterian synapomorphies: two-ended gut, mesodermal layers, central nervous system.
“The most important of these are a twoended gut with mouth and anus; the major use of mesodermal layers and structures in building the body plan; and a central nervous system.”
Quote in the paperSubstantial claim - Interpretationreasoning
The lack of horizontal burrowing in rocks older than about 575 Mya and vertical burrowing older than 543 Mya is strong evidence that no animals about 1 cm or longer capable of disturbing sediments existed before then.
“The lack of any evidence of horizontal burrowing in rocks older than about 575 Mya and of vertical burrowing in rocks older than 543 Mya is a strong argument that there existed no animals about 1 cm or longer that were capable of disturbing sedimentary layers before this time.”
Quote in the paperSubstantial claim- Rests on an assumption: The absence of trace fossils of burrowing by animals about 1 cm or longer before 575/543 Mya reflects absence of such animals rather than preservation bias.
- Interpretationcitation
Many toolkit items are used multiple times in constructing different parts of the same body plan.
“many items in the toolkit are used multiple times in the developmental construction of different parts of the same body plan.”
Quote in the paperSubstantial claim - Interpretationreasoning
Morphogenetic pattern formation programs by which body parts develop their form are clade-specific within phyla or classes.
“In contrast, the morphogenetic pattern formation programs by which the body parts develop their form are clade-specific within phyla or classes.”
Quote in the paperSubstantial claim - Interpretationreasoning
In other prominent examples interpreted as conservation of morphogenetic pathways, the transcriptional regulator is also still serving to control specific downstream differentiation programs at the terminus of development, as with pax6.
“However, in each case the transcriptional regulator on which the argument is based is also (still) serving to control specific downstream differentiation programs at the terminus of the developmental process, just as is the case of pax6.”
Quote in the paperSubstantial claim - Interpretationreasoning
In all scenarios of Fig. 3, the protostome-deuterostome divergence could have taken place hundreds of millions of years earlier.
“In all the scenarios of Fig. 3 that divergence could have taken place hundreds of millions of years earlier.”
Quote in the paperSubstantial claim - Interpretationnone given
The bilaterian morphogenesis toolkit consists of devices that establish spatial domains of transcriptional expression.
“the bilaterian toolkit for morphogenesis consists essentially of devices whose function is to establish spatial domains of transcriptional expression.”
Quote in the paperSubstantial claim - Interpretationnone given
Convincing arguments for conservation of adult body plan genetic programs between deuterostomes and protostomes pertain only to general features.
“convincing, and non-controversial, arguments for conservation of genetic regulatory programs for features of the adult body plan that are shared between deuterostomes and protostomes pertain only to general features”
Quote in the paperSubstantial claim - Premisenone given
Claims for pre-600 Mya metazoan fossils have not achieved general acceptance among paleontologists.
“There are also a variety of claims for pre-600 Mya metazoan fossils, but these have not achieved general acceptance among paleontologists.”
Quote in the paperNot a substantial claim - Interpretationreasoning
These morphogenetic programs have often continued to be utilized at the same location in the embryo.
“These morphogenetic programs have often continued to be utilized at the same location in the embryo.”
Quote in the paperNot a substantial claim - Interpretationcitation
Because crown-group cnidarians are already present in Doushantuo deposits, the cnidarian-bilaterian divergence must have occurred significantly before about 570-580 Mya.
“and second, since crown group cnidarians are already present in Duoshantuo deposits (Chen et al., 2002), the prior cnidarian-bilaterian divergence must have occurred significantly before about 570-580 Mya.”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Chen et al., 2002
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- Interpretationreasoning
Evolution of given body parts probably began with installation of cell differentiation programs to deploy specific cell types in a certain position, initially in a very simple morphological context.
“The evolution of given body parts probably began with the installation of cell differentiation programs to deploy specific cell types in a certain position in an organism, initially in a very simple morphological context.”
Quote in the paperSubstantial claim - Premisenone given
Diagnostic, unequivocally bilaterian trace fossils are associated only with latest Neoproterozoic and younger faunas.
“Diagnostic, unequivocally bilaterian trace fossils are associated only with the latest Neoproterozoic and younger faunas.”
Quote in the paperSubstantial claim - Premisecitation
Vertebrates and insects share contractile heart cell types that do not use MyoD, but in which certain homeodomain transcriptional regulators of the NK2 family plus Mef2 factors are required for differentiation.
“Vertebrates and insects share contractile heart cell types that do not use MyoD, but in which certain homeodomain transcriptional regulators of the NK2 family plus the Mef2 factors are required for differentiation (Olson and Srivnastava, 1996) (reviewed by Bodmer and Venkatesh, 1998)”
Quote in the paperSubstantial claim0 of 2 sources checked- cites Olson and Srivnastava, 1996 source
- cites Bodmer and Venkatesh, 1998
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- Interpretationreasoning
Building even the simple PDA required the same toolkit sufficient for all bilaterian diversification.
“To build the adult body plan of even the simple PDA that we discuss here would have required the same toolkit that has proved sufficient for the whole diversification of the bilaterians.”
Quote in the paperNot a substantial claim - Interpretationreasoning
A complex benthic vermiform PDA with appendages and segments would almost certainly have been detected in Neoproterozoic deposits.
“a complex, benthic, vermiform PDA equipped with appendages and segments would almost certainly have been detected, either as a trace or body fossil in Neoproterozoic deposits”
Quote in the paperSubstantial claim - Interpretationreasoning
Morphogenetic gene networks in Bilateria are evolutionarily very flexible.
“One thing that is clear from post-Cambrian evolution is that the morphogenetic gene networks used in the Bilateria are evolutionarily very flexible.”
Quote in the paperSubstantial claim
- May combine two claims. 33
- Classified as probably not a substantial claim. 7
- The quote only partly states this claim. 1
May combine two claims.
InterpretationcitationWhat is actually homologous in the role of pax6 in diversely constructed eyes is only its function in controlling genes encoding visual pigments; the gene was later coopted for different morphogenetic programs.
“The alternative is that what is actually homologous in the role of the pax6 gene in the diversely constructed eyes of various bilaterians is only its function in the control of genes encoding visual pigments.”
Quote in the paperSubstantial claim1 of 3 sources checkedProbably not a substantial claim: may not pass S1 (asserted), S4 (based).
PremisereasoningThese comparable deployment patterns have often been treated as evidence of functional homology, leading to the standard reconstruction of the PDA as a complex animal with some body parts like those of living bilaterians.
“These similarities have been widely interpreted as functional homologies, producing the conventional view of the last common protostome-deuterostome ancestor (PDA) as a complex organism that possessed some of the same body parts as modern bilaterians.”
Quote in the paperNot a substantial claimProbably not a substantial claim: may not pass S4 (based).
Premisenone givenA number of regulatory genes show broadly comparable deployment patterns in the formation of specific structures in flies and chordates.
“Many regulatory genes appear to be utilized in at least superficially similar ways in the development of particular body parts in Drosophila and in chordates.”
Quote in the paperNot a substantial claimMay combine two claims.
PremisecitationFurther analyses of rDNA phylogeny, Hox gene sequences, and morphology place flatworms within lophotrochozoa and acoels within flatworms.
“However, much further analysis of rDNA phylogeny as well as of Hox gene sequences and of morphological characters clearly places the flatworms within the lophotrochozoa, and the acoels within the flatworms (Adoutte, 2000;Berney et al., 2000;Littlewood et al., 2001;Peterson and Eernisse, 2001).”
Quote in the paperSubstantial claim0 of 3 sources checkedMay combine two claims.
PremisecitationSignal transduction pathways from ligands and receptors to transcription factors are orthologous and conserved across Bilateria.
“the signal transduction pathways leading from intercellular ligands and cell surface receptors to transcription factors are orthologous, despite the variety of names, and are conserved across Bilateria.”
Quote in the paperSubstantial claimMay combine two claims.
PremisecitationMultiple major continental glaciations occurred between about 760 and 580 Mya, and at least one possibly involved sea-level glaciation at near-equatorial latitudes.
“The latest Neoproterozoic is immediately preceded by multiple major continental glaciations that occurred between about 760 and 580 Mya (Kaufman et al., 1997).At least one and possibly more of these events involved glaciation at sea level at near-equatorial latitudes, a highly unusual occurrence.”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Kaufman et al., 1997 source
May combine two claims.
PremisecitationDifferentiation, or cell-type specification, is due to expression of specific batteries of structural genes and is a property of individual cells.
“Differentiation, or cell-type specification, is due to expression of specific batteries of structural genes that encode the proteins needed to endow that cell type with its functional properties; differentiation is a property of individual cells.”
Quote in the paperSubstantial claimMay combine two claims.
InterpretationreasoningSpatial control elements are like 'chips' that can build various patterning apparatuses; what they build depends on network architecture, not their identity.
“We may think of these spatial control elements (and many others that could be mentioned) as 'chips' in the toolkit, that can be used to build a great variety of developmental patterning apparatuses.”
Quote in the paperSubstantial claimMay combine two claims.
PremisecitationKimberella is interpreted as a benthic animal, most likely of protostome affinity and possibly a mollusk, with numerous specimens well dated at 555.3 Mya in northern Russia.
“This is interpreted as a benthic animal, most likely of protostome affinity, and possibly a mollusk (Fedonkin and Waggoner, 1997).Numerous specimens of Kimberella have been well dated at 555.3 Mya in northern Russia (Martin et al., 2000).”
Quote in the paperNot a substantial claim0 of 2 sources checkedMay combine two claims.
InterpretationreasoningBecause diverse crown-group bilaterians already existed in the Early Cambrian, bilaterian stem-group ancestors including the PDA must be significantly older than the Cambrian boundary and should predate Kimberella.
“The most general argument, independent of molecular clock extrapolations, is that given the diversity of crown group bilaterians already present in the Early Cambrian, bilaterian stem group ancestors, including the PDA, must be significantly more ancient than the Cambrian boundary; and they should also predate Kimberella.”
Quote in the paperSubstantial claimMay combine two claims.
InterpretationreasoningIf the protostome-deuterostome divergence occurred before these worldwide glaciations, the glaciations likely imposed a severe ecological constraint on the forms that could have survived.
“Although the extent of glaciation remains uncertain, if the protostome-deuterostome divergence occurred before these world-wide glaciations, they are likely to have imposed a severe ecological constraint on the forms that could have survived.”
Quote in the paperSubstantial claimMay combine two claims.
InterpretationreasoningPrimary differentiated cell types occur in all bilaterians, so the genome of the PDA must have included the respective differentiation gene batteries and their controllers.
“Primary differentiated cell types such as muscle cells, neurons of various sorts, photosensitive cells, etc., occur in all bilaterians and it follows that the genome of the PDA must have included the respective differentiation gene batteries and their controllers.”
Quote in the paperSubstantial claimMay combine two claims.
Premisenone givenThe first appearance and diversification of crown-group bilaterian clades occurred during the Early Cambrian.
“The first appearance and diversification of crown group bilaterian clades occurs during the Early Cambrian.”
Quote in the paperNot a substantial claimMay combine two claims. The quote only partly states this claim.
InterpretationreasoningIf the last common PDA predates 555 Mya, it is very unlikely to have been constructed in a complex manner, or its remains would probably have been found.
“One constraint is that if the last common PDA predates 555 Mya it is very unlikely to have been constructed in a complex manner”
Quote in the paperSubstantial claim- Rests on an assumption: The Neoproterozoic fossil record has been sampled well enough that a complex PDA would likely have been detected if it existed.
May combine two claims.
InterpretationreasoningThe revised topology has fundamentally changed inferred morphological homologies among Bilateria and interpretations of conserved developmental programs.
“The revised topology of Fig. 1B has fundamentally altered inferred morphological homologies among the Bilateria, and in consequence has affected interpretations of conserved developmental programs.”
Quote in the paperSubstantial claimMay combine two claims.
InterpretationreasoningA variety of bilaterian differentiation gene batteries are shared with cnidarians, while others are bilaterian-only.
“In summary, we conclude that a variety of bilaterian differentiation gene batteries are also shared with cnidarians, while others occur only among bilaterians.”
Quote in the paperSubstantial claimMay combine two claims.
InterpretationreasoningIn development, morphogenetic regulatory programs for pattern formation precede cell differentiation programs, but the reverse is likely to have been true in the evolution of body parts.
“In development, morphogenetic regulatory programs for pattern formation precede the institution of cell differentiation programs, but it is likely to have been the reverse in the evolution of body parts.”
Quote in the paperSubstantial claimMay combine two claims.
InterpretationreasoningThe Neoproterozoic fossil record does not yet explicitly illuminate the PDA and its antecedents, but it leaves valuable limits.
“So, in summary, although the Neoproterozoic fossil record does not yet explicitly illuminate the PDA and its antecedents, it leaves us with some valuable limits.”
Quote in the paperNot a substantial claimMay combine two claims.
InterpretationcitationThe bottom-up view holds that controlling differentiation gene batteries was primitive for each body part, while morphogenetic programs are not conserved but evolved independently and were added later.
“The 'bottom-up' view of body part evolution considered here (Davidson, 2001) is one in which the function of controlling differentiation gene batteries was primitive for each body part, while the morphogenetic programs are not conserved, but instead evolved independently and were added in later.”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Davidson, 2001
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
May combine two claims.
InterpretationreasoningThe PDA had an AP axis, two-ended gut, mesodermal layers, and central/peripheral nervous system with sensory cells.
“It had an AP axis, a twoended gut, mesodermal layers and a central and peripheral nervous system with sensory cell types.”
Quote in the paperSubstantial claim- The authors state this as an assumption: The PDA had all the general features discussed: a two-ended gut, mesodermal layers and structures, and a central nervous system.
May combine two claims.
InterpretationreasoningThe most reasonable conclusion is that the Neoproterozoic PDA was small and probably constructed simply and delicately.
“The most reasonable conclusion is that the Neoproterozoic PDA must have been small and was probably constructed simply and delicately.”
Quote in the paperSubstantial claimProbably not a substantial claim: may not pass S1 (asserted), S3 (load-bearing).
PremisecitationRunnegar argued that conditions even within glacial refugia would have allowed survival only of small, simply constructed, pelagic bilaterian stem-group forms.
“Runnegar (Runnegar, 2000) argued that conditions even within the refugia would have allowed survival only of small, simply constructed, pelagic bilaterian stem group forms”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Runnegar, 2000
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
May combine two claims.
PremisecitationMolecular phylogenetics reveals metazoan relationships very different from traditional topologies, including placement of acoelomates and pseudocoelomates within protostomes, recognition of ecdysozoa and lophotrochozoa, and alliance of annelids with molluscs rather than arthropods.
“Among the most important changes are the transfer of the acoelomates and pseudocoelomates to within the protostomes; the recognition of two great protostome subclades, the ecdysozoa and lophotrochozoa, and the alliance of annelids with molluscs and other lophotrochozoans rather than with arthropods (ecdysozoa).”
Quote in the paperSubstantial claimMay combine two claims.
Premisenone givenThe Ediacaran fauna appears in the latest Neoproterozoic (c. 575-543 Mya), with a few forms extending into the earliest Cambrian, and includes at least one demonstrable adult bilaterian, Kimberella.
“A new assemblage of organisms, known as Ediacaran fauna, appears in the latest Neoproterozoic (c 575-543 Mya), with a few extending into the earliest Cambrian.”
- Also stated in: § THE 'MISSING' FOSSIL RECORD · boosted
“at least one demonstrable adult bilaterian form, Kimberella (Fig. 2).”
Quote in the paperSubstantial claim- Also stated in: § THE 'MISSING' FOSSIL RECORD · boosted
Probably not a substantial claim: may not pass S2 (checkable).
InterpretationreasoningIt is dangerous to assume such chips have a hard pan-bilaterian morphogenetic assignment conserved from an ancestral form.
“Therefore it is particularly dangerous to assume that such chips have a hard, pan-bilaterian morphogenetic assignment, that has been conserved from an ancestral form.”
Quote in the paperNot a substantial claimMay combine two claims.
InterpretationreasoningWhen bilaterian traces appear, they indicate animals with anteroposterior differentiation, but there is no evidence of limbs.
“When they do appear, these bilaterian traces indicate the presence of animals that had AP differentiation, but there is no evidence of limbs.”
Quote in the paperNot a substantial claimMay combine two claims.
Premisenone givenAfter 550 Mya metazoan diversity and complexity accelerated, beginning with simple skeletonized tubes and more complex trace fossils, followed at about 530 Mya by the first brachiopods, molluscs, and arthropods.
“After 550 Mya there is an accelerating increase in metazoan diversity and complexity, beginning with simple skeletonized tubes and more complex trace fossils, followed at about 530 Mya by the first brachiopods, molluscs and arthropods.”
Quote in the paperNot a substantial claimMay combine two claims.
InterpretationreasoningThe absence of bilaterian fossils before the very end of the Neoproterozoic does not prove that bilaterians were absent then, but it does place meaningful constraints on what they could have been like.
“The absence of bilaterian fossils prior to the very end of the Neoproterozoic, at least so far, does not mean that bilaterians did not exist then, but it does place meaningful constraints on their possible nature.”
Quote in the paperSubstantial claim- Rests on an assumption: The Neoproterozoic fossil record has been sampled well enough that a complex PDA would likely have been detected if it existed.
May combine two claims.
InterpretationcitationPhosphatic preservation like that in the Doushantuo appears restricted to a relatively narrow Neoproterozoic interval, and similar preservation opportunities may not occur in older rocks.
“Such unusual phosphatic preservation has been described from elsewhere in the Phanerozoic, but this complex preservational environment (Xiao and Knoll, 1999) appears restricted to a relatively narrow interval of the Neoproterozoic (Cook and Shergold, 1984).”
Quote in the paperSubstantial claim0 of 2 sources checkedProbably not a substantial claim: may not pass S2 (checkable), S3 (load-bearing), S4 (based). May combine two claims.
InterpretationreasoningFossilization of this simple PDA might require rare special circumstances, and its fossils might not be recognized as such unless prior expectations are adjusted.
“Fossilization of this animal might require rare special circumstances, and unless prior expectations are suitably adjusted, its fossils might not be recognized as such even if found.”
Quote in the paperNot a substantial claimProbably not a substantial claim: may not pass S1 (asserted).
InterpretationreasoningIf the canonical body parts of all bilaterians are truly homologous, then the PDA should have had all these same parts.
“If the canonical body parts of all bilaterians are truly homologous, as suggested by commonalties in their developmental regulatory programs, then the PDA should have had all these same parts.”
Quote in the paperNot a substantial claimMay combine two claims.
Interpretationnone givenOther Ediacaran fossils assigned to annelids, echinoderms, or arthropods lack characters convincingly supporting those assignments; most lack appendages, mouth, digestive system, or other bilaterian characters and can be interpreted as cnidarian-grade.
“Other elements of the Ediacaran fauna have been assigned to the annelids, echinoderms and arthropods, but none display characters convincingly supporting such phylogenetic assignments.Most Ediacaran fossils lack clearly identifiable appendages, mouth, indications of a digestive system or other characters supporting assignment to the Bilateria.But most can be comfortably interpreted as cnidarian-”
Quote in the paperSubstantial claimMay combine two claims.
InterpretationreasoningIn each body part, the same differentiated cell types are employed across Bilateria, and this fact underlies their analogous functions.
“However, in each of the body parts, respectively, the same differentiated cell types are employed across the Bilateria, and it is this fact that underlies their analogous functions”
Quote in the paperSubstantial claimMay combine two claims.
InterpretationreasoningThe PDA possessed the essential bilaterian toolkit and deployed many differentiation gene batteries.
“In conclusion, the PDA possessed the essential bilaterian toolkit for morphogenetic pattern formation, and it deployed many of the differentiation gene batteries that its modern descendants continue to rely on.”
Quote in the paperSubstantial claimMay combine two claims.
InterpretationcitationThere is already some convincing evidence that the architecture of gene networks controlling formation of many analogous body parts in unrelated bilaterian clades is clade-specific, while the regulatory apparatus for differentiated cell types is conserved across Bilateria.
“Though yet fragmentary, there is already some convincing evidence on both scores (Davidson, 2001).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Davidson, 2001
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.
PremisecitationMany new acritarch groups appeared during the earliest Cambrian, perhaps linked to the origin of zooplankton.
“Many new acritarch groups appeared during the earliest Cambrian (Vidal and Moczydl ⁄ owska-Vidal, 1997), perhaps linked to the origination of zooplankton (Butterfield, 1997).”
Quote in the paperNot a substantial claim0 of 2 sources checkedMay combine two claims.
PremisecitationAcritarchs are the most diverse and heterogeneous fossils from the 1200-740 Mya interval and increased greatly in diversity during the Neoproterozoic.
“The most diverse and heterogeneous fossils from this interval are resistant, organic-walled microfossils known as acritarchs, which increased greatly in diversity during the Neoproterozoic (Knoll, 1996; Vidal and Moczydl ⁄ owska-Vidal, 1997; Xiao and Knoll, 1999; Zhou et al., 2001).”
Quote in the paperNot a substantial claim0 of 3 sources checkedMay combine two claims.
PremisecitationMany conserved developmental control genes and some differentiated cell types are present in Cnidaria.
“Many of the highly conserved developmental control genes are also present in Cnidaria, as are some of these same differentiated cell types”
Quote in the paperSubstantial claim
The authors state this as an assumption.
Drosophila and vertebrates are adequate surrogates for Bilateria in comparative analysis of conserved regulatory systems.
“Taking Drosophila and vertebrates as surrogates for Bilateria (this leaves aside Lophotrochozoa, for which there is as yet only relatively sparse molecular biological evidence)”
The paper relies on this without support.
The Neoproterozoic fossil record has been sampled well enough that a complex PDA would likely have been detected if it existed.
“Paleontologists have extensively explored vast areas of the Neoproterozoic (1000-543 Mya) fossil record, including a variety of marine environments and styles of preservation.”
The paper relies on this without support.
Molecular clock estimates, despite potential problems, are reliable enough to indicate that the latest PDA is much older than the latest Neoproterozoic.
“Despite various potential problems with molecular clock estimates (Smith and Peterson, 2002), most of these suggest that the age of the latest PDA must lie much deeper in time than the latest Neoproterozoic.”
The paper relies on this without support.
The absence of trace fossils of burrowing by animals about 1 cm or longer before 575/543 Mya reflects absence of such animals rather than preservation bias.
“The lack of any evidence of horizontal burrowing in rocks older than about 575 Mya and of vertical burrowing in rocks older than 543 Mya is a strong argument that there existed no animals about 1 cm or longer that were capable of disturbing sedimentary layers before this time.”
The authors state this as an assumption.
The PDA had all the general features discussed: a two-ended gut, mesodermal layers and structures, and a central nervous system.
“We may assume that the PDA had all these general features, and that it utilized some of the genetic control apparatus which modern Bilateria display in the development of these basal aspects of their body plans.”
The authors state this as an assumption.
The PDA utilized some of the genetic control apparatus that modern Bilateria display in developing these basal body plan features.
“We may assume that the PDA had all these general features, and that it utilized some of the genetic control apparatus which modern Bilateria display in the development of these basal aspects of their body plans.”
The authors state this as an assumption.
The PDA's morphology was unprepossessing.
“the safest assumption is that its morphology was unprepossessing.”
Claims other papers make about this one
Each is checked against this paper’s own text: does it say what the citing paper says it does?
At least 600 million years of evolution separate cnidarians from bilaterians.
“Though at least 600 million years of evolution separate cnidarians from bilaterians [9–16], many aspects of the nervous system are conserved, including neuropeptides and neurotransmitters [15–20].”
From The Jellyfish Cassiopea Exhibits a Sleep-like State (2017) · cites this paper as [12]
Bilaterians employ similar fundamental approaches in constructing their varied body forms.
“Over the last decade, an enormous amount of evidence from developmental molecular biology, much of it presented in these same pages, has shown that all Bilateria share a common genetic toolkit for development, and that all Bilateria use the same basic strategies to build their diverse body plans (reviewed in Davidson, 2001;Erwin and Davidson, 2002;Carroll et al., 2001).”
From Precambrian Animal Life: Probable Developmental and Adult Cnidarian Forms from Southwest China (2002) · cites this paper as Erwin and Davidson, 2002
Bilaterians possess a shared set of developmental genes.
“Over the last decade, an enormous amount of evidence from developmental molecular biology, much of it presented in these same pages, has shown that all Bilateria share a common genetic toolkit for development, and that all Bilateria use the same basic strategies to build their diverse body plans (reviewed in Davidson, 2001;Erwin and Davidson, 2002;Carroll et al., 2001).”
From Precambrian Animal Life: Probable Developmental and Adult Cnidarian Forms from Southwest China (2002) · cites this paper as Erwin and Davidson, 2002
Fossil evidence is nearly absent regarding the shape, age, or stem lineage of the last common bilaterian ancestor.
“But of the form of this ancestor, or the time at which it lived, or the nature of the stem group lineage from which it derived, there is yet almost no information in the fossil record (see Erwin and Davidson, 2002 for discussion).”
From Precambrian Animal Life: Probable Developmental and Adult Cnidarian Forms from Southwest China (2002) · cites this paper as Erwin and Davidson, 2002
These results suggest that the biochemical functions of CnASH have been conserved despite the divergence of cnidarians and arthropods before the Cambrian explosion.
“These results suggest that although cnidarians and arthropods diverged prior to the Cambrian explosion (Erwin and Davidson, 2002), the biochemical functions of the CnASH protein have been conserved”
From Evolutionary aspects of developmentally regulated helix-loop-helix transcription factors in striated muscle of jellyfish (2003) · cites this paper as Erwin and Davidson, 2002
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- Authors
- Erwin, D. H., & Davidson, E. H.
- Affiliations
- National Museum of Natural History (US)
- California Institute of Technology (US)
- ORCID
- Erwin, Douglas H.
- Venue
- Development
- Publisher
- The Company of Biologists
- Published
- 2002-07-01
- Volume
- 129
- Issue
- 13
- Pages
- 3021-3032
- ISSN
- 0950-1991 (ISSN-L)
- 1477-9129
- OpenAlex
- W2165511961
- Cited by
- 320
- References
- 153
- Citations by year
- 2026: 3 · 2025: 3 · 2024: 13 · 2023: 7 · 2022: 8 · 2021: 6 · 2020: 4 · 2019: 13 · 2018: 9 · 2017: 7 · 2016: 4 · 2015: 15 · 2014: 10 · 2013: 9 · 2012: 10
- FWCI
- 1.42
- Type
- journal article
- Language
- en
- Open access
- green
- resolver.caltech.edu
- Locations
- Development (publishedVersion)
- PubMed (publishedVersion)
- CaltechAUTHORS (California Institute of Technology) (submittedVersion)
- HighWire Press Open Archive (submittedVersion)
- Funders
- Eugene V. and Claire E. Thaw Charitable Trust
- Santa Fe Institute
- Topics
- Life Sciences › Biochemistry, Genetics and Molecular Biology › Molecular Biology › Developmental Biology and Gene Regulation
- Physical Sciences › Earth and Planetary Sciences › Paleontology › Marine Invertebrate Physiology and Ecology
- Social Sciences › Arts and Humanities › History and Philosophy of Science › Philosophy and History of Science
- Keywords
- bilaterian evolution, regulatory genes, functional homology, Drosophila, Chordata, body plan, evolutionary developmental biology
- MeSH terms
- Animals, Chordata, Nonvertebrate, Drosophila, Biological Evolution, Fossils, Morphogenesis, Phylogeny, Gene Expression Regulation, Developmental
About these checks
Registry, retraction, Cites and statistics results are computed, not judged, with one exception: a printed reference that no index lists is read once by a model before it counts as a citation error. Claims are extracted by a model and kept only when their quote appears verbatim in the text; each cited source is then checked, with a model’s help, for whether it says what the claim relies on. Those verdicts can be wrong — read the claim’s quote and the cited source before relying on one.
Claims read by deepseek-v4.1-flash under definition version 3 (extract-v3) on 2026-09-30.
Classified by jev-1.13-20260917 (gold-guideline-v0.2) on 2026-09-30.
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