{
  "study_id": "DISC-114-BASAL-METAZOA-CTENO-PORIFERA-CNIDARIA-TIMETREE",
  "slug": "cteno-porifera-cnidaria-timetree",
  "preprint_slug": "basal-metazoa-timetree-preprint",
  "preprint_id": "PREPRINT-05",
  "validation_set_id": "VAL-SET-044-BASAL-METAZOA-CTENO-PORIFERA-CNIDARIA-TIMETREE-64TAXA-1.64MB",
  "generated_at": "2026-11-04T23:12:00.000Z",
  "engine": "RuNIE v1.27.0 (Rust Native Inference Engine)",
  "mode": "Dual-Pass Blinded Dry-Lab Mode (SHA-256 + D_ctx) -> Full Step-Up (Tier 1 -> Tier 2 -> Tier 3) -> Final-Report-Only Publication Mode",
  "bootstrap_replicates": 500,
  "prior_beakers": 5,
  "posterior_beakers": 5,
  "tier_trajectory": {
    "tier_1_exploratory": {
      "taxa": 18,
      "cohorts": 8,
      "aligned_bp": 312000,
      "rate_cv": 0.3842,
      "ctenophore_aa_rate_subs_per_site_myr": 0.00188,
      "porifera_aa_rate_subs_per_site_myr": 0.00055,
      "anthozoa_aa_rate_subs_per_site_myr": 0.00058,
      "cteno_to_sponge_rate_ratio": 3.42,
      "strict_clock_20state_root_ma": 1148.0,
      "relaxed_clock_dayhoff6_root_ma": 792.4,
      "strict_clock_inflation_ma": 355.6,
      "posterior_z": 3.62,
      "posterior_beakers": 3
    },
    "tier_2_blinded_validation": {
      "taxa": 42,
      "cohorts": 24,
      "dev_cohorts": 12,
      "reserve_cohorts": 12,
      "aligned_bp": 884000,
      "rcfv_20state": 0.142,
      "rcfv_dayhoff6": 0.023,
      "rcfv_reduction_pct": 83.8,
      "concordant_cohorts": 20,
      "mean_abs_delta_z_concordant": 0.0,
      "dev_mean_z": 4.45,
      "reserve_mean_z": 4.39,
      "overall_posterior_z": 4.42,
      "posterior_beakers": 4,
      "intercepted_anomalies": [
        {
          "token": "BLIND-METAZOA-04",
          "split": "DEV",
          "accession": "NC_016117.1",
          "issue": "Hyper-compact 10,326 bp Mnemiopsis leidyi mitochondrial genome (84.2% AT, missing atp6/atp8, >12x rate acceleration) co-aligned under a single strict clock with slow 18-20 kb anthozoan/sponge mtDNA, inflating Crown Metazoa to 1,284.0 Ma",
          "d_ctx_sd": 5.46
        },
        {
          "token": "BLIND-METAZOA-09",
          "split": "DEV",
          "accession": "GCF_000002865.3",
          "issue": "High-RCFV choanoflagellate Monosiga brevicollis outgroup + unrecoded 20-state amino-acid ribosomal supermatrix inducing Porifera-sister LBA inversion and +294.0 Ma basal stem stretch",
          "d_ctx_sd": 5.12
        },
        {
          "token": "BLIND-METAZOA-16",
          "split": "RESERVE",
          "accession": "CRYOGENIAN-24IPC-STERANE",
          "issue": "Cryogenian 24-isopropylcholestane sterane biomarker applied as a hard 660.0 Ma Crown Demospongiae minimum bound despite rhizarian/algal sterol diagenesis, stretching the sponge stem by +138.0 Ma",
          "d_ctx_sd": 4.88
        },
        {
          "token": "BLIND-METAZOA-21",
          "split": "RESERVE",
          "accession": "GCF_013753865.1-SYMBIO",
          "issue": "Endosymbiotic dinoflagellate (Symbiodinium) transcriptomic contig chimera embedded in an Acropora reef-coral ortholog supermatrix",
          "d_ctx_sd": 4.76
        }
      ]
    },
    "tier_3_scientific_synthesis": {
      "taxa": 64,
      "extant_taxa": 54,
      "fossil_tips": 10,
      "cohorts": 32,
      "aligned_bp": 1642800,
      "nuclear_orthologs": 212,
      "ancestral_linkage_groups": 29,
      "irreversible_bcns_fusions_benthozoa": 7,
      "bcns_fisher_exact_p": 1.4e-21,
      "bootstrap_replicates": 500,
      "map_topology_support_pct": 99.6,
      "map_topology_support_sd_pct": 0.2,
      "posterior_z_pos": 5.48,
      "posterior_z_all": 4.82,
      "posterior_beakers": 5,
      "dated_nodes": [
        {
          "node": "Choanozoa (Choanoflagellata vs. Metazoa)",
          "age_ma": 892.0,
          "ci_95_ma": [858.0, 926.0],
          "geological_stage": "Early Tonian",
          "genomic_synapomorphy": "29 Ancestral Linkage Groups (ALGs); cadherin extracellular repeats; proto-tyrosine kinase signaling"
        },
        {
          "node": "Crown Metazoa (Ctenophora vs. Benthozoa/Myriazoa)",
          "age_ma": 792.4,
          "ci_95_ma": [762.0, 822.8],
          "geological_stage": "Late Tonian (Pre-Sturtian)",
          "genomic_synapomorphy": "Collagen IV NC1 sulfilimine basement membrane (Met93-Hyl211, PDB:1LI1); ancestral DEEA-pore Cav/Nav1 channels (PDB:6A90); Wnt/beta-catenin polarity"
        },
        {
          "node": "Benthozoa / Myriazoa (Porifera vs. ParaHoxozoa)",
          "age_ma": 738.6,
          "ci_95_ma": [710.4, 766.8],
          "geological_stage": "Terminal Tonian / Sturtian Onset",
          "genomic_synapomorphy": "7 irreversible BCnS chromosome fusion-with-mixing events (A1a+A1b, C1+C2, E1+E2, G+H, J1+J2, L+M, N+O; p = 1.4e-21); primary pelago-benthic life cycle"
        },
        {
          "node": "ParaHoxozoa (Placozoa vs. Planulozoa)",
          "age_ma": 684.2,
          "ci_95_ma": [658.0, 710.4],
          "geological_stage": "Cryogenian Interglacial",
          "genomic_synapomorphy": "ParaHox/Hox (Gsx, Xlox, Cdx) ANTP homeobox cluster; Classical Cadherin EC1-EC2 3xCa2+ hinge (PDB:3Q2V); PSD-95/DLG scaffold (PDB:1BE9)"
        },
        {
          "node": "Planulozoa (Cnidaria vs. Bilateria)",
          "age_ma": 642.5,
          "ci_95_ma": [618.0, 667.0],
          "geological_stage": "Terminal Cryogenian / Early Ediacaran",
          "genomic_synapomorphy": "Striated muscle myosin heavy chain (ST-MHC) + Troponin/Tropomyosin; canonical Neuroligin-Neurexin chemical synapses; DEKA-pore Nav selectivity filter"
        },
        {
          "node": "Crown Porifera (Silicea vs. Calcarea + Homoscleromorpha)",
          "age_ma": 614.8,
          "ci_95_ma": [590.2, 639.4],
          "geological_stage": "Early Ediacaran",
          "genomic_synapomorphy": "Choanocyte aquiferous collar complex; silicatein/carbonic anhydrase biomineralization toolkit"
        },
        {
          "node": "Crown Cnidaria (Anthozoa vs. Medusozoa)",
          "age_ma": 586.4,
          "ci_95_ma": [564.8, 608.0],
          "geological_stage": "Mid-Ediacaran (Pre-Gaskiers)",
          "genomic_synapomorphy": "Cnidocyte minicollagen/nematogalectin capsule; Chordin/BMP-2/4 directive axis; anchored by 560 Ma †Auroralumina attenboroughii"
        },
        {
          "node": "Crown Bilateria (Protostomia vs. Deuterostomia)",
          "age_ma": 568.2,
          "ci_95_ma": [549.0, 587.4],
          "geological_stage": "Late Ediacaran",
          "genomic_synapomorphy": "Collinear 7-13 gene Hox cluster; nephrozoan excretory/triploblastic mesoderm; anchored by 555 Ma †Kimberella & 518 Ma †Haikouichthys"
        },
        {
          "node": "Crown Ctenophora (Cydippida/Lobata/Beroida Radiation)",
          "age_ma": 384.6,
          "ci_95_ma": [352.0, 417.2],
          "geological_stage": "Middle Devonian (after 407.8 Myr stem survival)",
          "genomic_synapomorphy": "Syncytial Innexin nerve net (12-14 paralogs); colloblasts; 8-comb-row macrocilia; stem anchored by 551 Ma †Eoandromeda & 518 Ma †Daihua"
        }
      ]
    }
  }
}
