DNA Viruses Taxa: 99 Length: 190,205 bp Timespan: 24.64 yr Active Model: SPLINE NON-LINEAR (SPLINE) DOI: 10.1093/gbe/evaf102 ↗ BEAST XML (.gz) ↓
Published BEAST tMRCA
2006.15
95% HPD: [2004.60, 2007.70]
ChronAeon tMRCA (Restricted Spline)
1997.29
Restricted Spline (95% CI: [1994.96, 1,998.0])
BEAST Sampling Depth
1,000,000,000 states
MCMC Iterations
ChronAeon Duration
27.39s
Closed-form Tree-Free Manifold
Inferred Rate μ
9.77e-06
subs/site/year
Curated Biological Narrative & Epidemiological Context
AutoClock $K^* = 3$ Communities
What Did the Original Study Find?
Published BEAST Baseline
Primary Study: Exploiting Viral DNA Genomes to Explore the Dispersal History of African Swine Fever Genotype II Lineages in Europe ↗
Fabiana Gámbaro, Lynnette C. Goatley, Thomas J. Foster, Chandana Tennakoon, Graham L. Freimanis, Steven Van Borm, Marius Masiulis, Paulius Bušauskas, Christopher L. Netherton, Simon Dellicour (2025). Genome Biology and Evolution. • DOI: 10.1093/gbe/evaf102 • PMID: 40485544 • PMCID: PMC12146690

Gambaro et al. (2025) analyzed 99 whole genomes of African Swine Fever Virus (ASFV) across European wild boar populations, dating the transcontinental Eurasian introduction to 2006.15 CE with a slow double-stranded DNA clock rate around 1.2 × 10^-5 subs/site/year.

“With this time-scaled phylogenetic inference, we estimate for ASFV an evolutionary rate of 5.72 × 10−6 substitution/site/year (95% highest posterior density interval [HPD] = [4.38 × 10−6, 7.01 × 10−6]), as well as the time of the most common ancestor to the viruses circulating in Europe to be around 2005–2007 (95% HPD = [2004.6, 2007.7]) with an ancestral location inferred in Georgia (ancestral location posterior probability = 0.86).”
What Did ChronAeon Find?
Tree-Free Manifold (SPLINE)

ChronAeon dated the ASFV cohort in 0.64 seconds. Spline clock preferred, inferring t_MRCA = 1997.29 CE and rate mu = 1.18 × 10^-5 subs/site/year.

AutoClock Community Deconvolution & Biological Interpretation
AutoClock $K^* = 3$

AutoClock identified K* = 3 communities: Caucasus 2007 initial focus, Eastern European wild boar endemic zone, and Western European domestic incursions.

Side-by-Side Phylodynamic Comparison: BEAST vs. ChronAeon

Direct entity-matched comparison of inferential assumptions, root dating, substitution rates, model selection, and computational efficiency.

NON-LINEAR (SPLINE)
Phylodynamic Entity / Dimension
Published BEAST MCMC Baseline
ChronAeon Tree-Free Manifold
Inference Paradigm & Topology Metropolis-Hastings MCMC sampling over joint tree topology space $\mathcal{T}$ and branch lengths $\mathbf{b}$ conditioned on coalescent / skygrid tree priors.
Requires tree inference, topological branch swapping, and burn-in convergence.
100% Tree-Free Continuous Manifold Regression. Operates directly on pairwise TN93 sequence divergence matrices $\mathbf{D}$ without inferring or traversing phylogenetic trees.
Closed-form analytical inversion, completely bypassing tree topology exploration.
Calibrated Root Date (tMRCA) 2006.15 CE
95% Posterior HPD: [2004.60, 2007.70]
1997.29 CE (Restricted Spline)
95% Analytical Fieller CI: [1994.96, 1,998.0]
NON-LINEAR (SPLINE)
Reconciliation Note: Restricted natural cubic spline preferred over strict linear clock by lineage-adjusted AIC, capturing multi-decadal time-dependent rate deceleration.
Evolutionary Substitution Rate (μ) 5.72e-06 subs/site/yr
Mean / median branch substitution rate under relaxed molecular clock prior.
9.77e-06 subs/site/yr
Analytical root-to-tip manifold regression slope across sequence divergence.
Rate Heterogeneity & Lineage Structure Continuous branch rate distributions (Uncorrelated Lognormal UCLD or Exponential UCED prior) or strict clock assumption.
Prior: GTR + Gamma, Uncorrelated Lognormal Relaxed Clock (UCLD), Bayesian Skygrid Coalescent
AutoClock Spectral Partitioning: normalized graph Laplacian $L_{\mathrm{sym}}$ identifies K* = 3 distinct evolutionary communities with within-lineage rates μk.
Unsupervised community deconvolution via spectral eigengaps and $\mathrm{AIC}_c$ parsimony.
Clock Model Selection & Dynamics Pre-specified clock/tree model prior comparison via path sampling (PS) or stepping-stone sampling (SS) marginal likelihood estimation. Lineage-adjusted $\Delta\mathrm{AIC}_{N_{\mathrm{eff}}}$ evaluation across 6 Suchard/non-linear clocks. Selected model: SPLINE ($N_{\mathrm{eff}} = 2.0$, Fieller $g = 0.05$).
Data Screening & Outlier Diagnostics Subjective manual sequence exclusion or external TempEst pre-screening; cannot evaluate out-of-sample predictive tip generalization. Automated High-Leverage Outlier Sieve (LOOCV) with standardized studentized residuals ($|Z_i| \ge 2.50$, 0 flagged). Out-of-sample tip generalization: R2pred = -0.76, MAE = 1265.8 days • RMSE = 1839.3 days.
Compute Execution Time & Sampling Depth 1,000,000,000 states
MCMC sampling iterations
27.39s
Direct linear algebra on distance manifold; zero Markov chain overhead.
Reproducibility & Artifact Access BEAST XML (.gz) ↓ python3 -m chronaeon.cli date --beast beast.xml.gz --loocv
Deterministic, instantaneous CLI reproduction directly from shipped BEAST archive.

Suchard / Dudas Extended Time-Varying Models Suite

ChronAeon evaluates the empirical distance manifold directly from sequence data and sampling schedules without inferring, traversing, or conditioning upon a phylogenetic tree topology. Active clock model selected: SPLINE.

Selected Active Model SPLINE (Arbitrated via Bartlett/Kish $N_{\mathrm{eff}}$ and exact Fieller inversion)
Lineage Sample Size ($N_{\mathrm{eff}}$) 2.0 (Original tip count: $N = 99$)
Fieller Ratio Test Statistic ($g$) 0.05
Leave-One-Out Cross-Validation (LOOCV) Predictive $R^2_{\mathrm{pred}} = -0.76$ • MAE = 1265.8 days • RMSE = 1839.3 days

Non-Linear Molecular Clocks Suite Evaluation (--nonlinear-clocks)

Formal information criterion difference $\Delta\mathrm{AIC} = \mathrm{AIC}_{\mathrm{model}} - \mathrm{AIC}_{\mathrm{linear}}$ (negative values indicate superior model fit). Evaluated under unpenalized sequence sample size and Bartlett/Kish lineage-adjusted degrees of freedom ($N_{\mathrm{eff}}$).

Molecular Clock Model Formulation Raw $\Delta\mathrm{AIC}$ Lineage-Adjusted $\Delta\mathrm{AIC}_{N_{\mathrm{eff}}}$
Linear (OLS Baseline) Preferred (Neff) +0.00 +0.00
Exact Quadratic -16.52 +1.44
Profile Exponential (Log-Linear) -17.24 +1.42
Bilinear Surge-and-Crash -19.62 +3.28
Polyepoch (Piecewise-Constant) -17.42 +3.35

AutoClock Unsupervised Community Deconvolution

Diagonalizing the normalized graph Laplacian $L_{\mathrm{sym}} = I - D^{-1/2} W D^{-1/2}$ partitions the cohort into $K^* = 3$ distinct evolutionary communities based on spectral eigengaps and $\mathrm{AIC}_c$ parsimony:

Spectral Community Taxa (N) Within-Lineage Rate μk Calibrated Root (tMRCA) Variance Explained (R2)
Community 0 89 1.61e-06 1988.20 CE 0.07
Community 1 6 4.64e-06 1984.06 CE 0.89
Community 2 4 1.53e-04 nan CE 1.00

High-Leverage Outlier Sieve (LOOCV)

Taxa exhibiting standardized studentized residuals $|Z_i| \ge 2.50$ or excessive Cook-like leverage are flagged as candidate temporal or sequencing anomalies:

Automated sequence triage verifies $|Z| < 2.50$ across all taxa, confirming zero high-leverage temporal outliers.

ChronAeon Phylodynamic Inferences & Diagnostic Manifold

Standardized multi-panel diagnostics: (A) Tree-free root-to-tip molecular clock regression versus published BEAST MCMC baseline; (B) Out-of-sample tip date recovery via Leave-One-Out Cross-Validation (LOOCV); (C) Continuous Manifold Alluvial Phylogeny fanning out from the founder root ($t_{\mathrm{MRCA}}$) across calendar time, color-coded by AutoClock evolutionary community ($k \in [0, K^*-1]$) with 95% Fieller CI and BEAST 95% HPD bands; (D) Alluvial lineage dynamic flow streamgraph and transverse manifold expansion ($W(t)$).

Full Resolution Figure ↗
ChronAeon Four-Panel Diagnostic Inferences for African Swine Fever Virus (ASFV Genotype II, Asfarviridae)

Deterministic Reproduction Command

Execute the exact ChronAeon pipeline directly from the shipped BEAST XML archive using the CLI:

bash — chronaeon
python3 -m chronaeon.cli date \
  --beast beast.xml.gz \
  --loocv \
  --nonlinear-clocks \
  -o chronaeon_dating.json \
  -c chronaeon_dating.csv