<p>Accurate estimation of speciation (<InlineEquation ID="IEq1"> <EquationSource Format="TEX">\(\lambda \)</EquationSource> <EquationSource Format="MATHML"><math> <mi>λ</mi> </math></EquationSource> </InlineEquation>) and extinction (<InlineEquation ID="IEq2"> <EquationSource Format="TEX">\(\mu \)</EquationSource> <EquationSource Format="MATHML"><math> <mi>μ</mi> </math></EquationSource> </InlineEquation>) rates from phylogenetic trees is central to studies of diversification, yet it remains unclear whether commonly used estimators are unbiased. Here we examine two sources of error: (1) statistical bias in the estimators themselves, and the (2) structural bias introduced by how small trees are handled in likelihood calculations. For the Yule process, we re-derive the expected bias of the standard estimator, showing that <InlineEquation ID="IEq3"> <EquationSource Format="TEX">\(\hat{\lambda }\)</EquationSource> <EquationSource Format="MATHML"><math> <mover accent="true"> <mi>λ</mi> <mo stretchy="false">^</mo> </mover> </math></EquationSource> </InlineEquation> underestimates <InlineEquation ID="IEq4"> <EquationSource Format="TEX">\(\lambda \)</EquationSource> <EquationSource Format="MATHML"><math> <mi>λ</mi> </math></EquationSource> </InlineEquation> by a factor of <InlineEquation ID="IEq5"> <EquationSource Format="TEX">\((n-2)/(n-1)\)</EquationSource> <EquationSource Format="MATHML"><math> <mrow> <mo stretchy="false">(</mo> <mi>n</mi> <mo>-</mo> <mn>2</mn> <mo stretchy="false">)</mo> <mo stretchy="false">/</mo> <mo stretchy="false">(</mo> <mi>n</mi> <mo>-</mo> <mn>1</mn> <mo stretchy="false">)</mo> </mrow> </math></EquationSource> </InlineEquation>. Extending to the general birth–death model, we use symbolic regression to find functional forms that minimize the bias in both <InlineEquation ID="IEq6"> <EquationSource Format="TEX">\(\lambda \)</EquationSource> <EquationSource Format="MATHML"><math> <mi>λ</mi> </math></EquationSource> </InlineEquation> and <InlineEquation ID="IEq7"> <EquationSource Format="TEX">\(\mu \)</EquationSource> <EquationSource Format="MATHML"><math> <mi>μ</mi> </math></EquationSource> </InlineEquation>. The best-performing correction for <InlineEquation ID="IEq8"> <EquationSource Format="TEX">\(\lambda \)</EquationSource> <EquationSource Format="MATHML"><math> <mi>λ</mi> </math></EquationSource> </InlineEquation> is identical to the Yule result, while the bias in <InlineEquation ID="IEq9"> <EquationSource Format="TEX">\(\mu \)</EquationSource> <EquationSource Format="MATHML"><math> <mi>μ</mi> </math></EquationSource> </InlineEquation> depends on both sample size and the estimated extinction fraction (<InlineEquation ID="IEq10"> <EquationSource Format="TEX">\(\mu /\lambda \)</EquationSource> <EquationSource Format="MATHML"><math> <mrow> <mi>μ</mi> <mo stretchy="false">/</mo> <mi>λ</mi> </mrow> </math></EquationSource> </InlineEquation>). Applying these corrections substantially improves the fit between the estimated and generating values. When these corrected estimators are used to derive other diversification-related parameters, turnover is nearly unbiased, but net diversification (<InlineEquation ID="IEq11"> <EquationSource Format="TEX">\(\lambda - \mu \)</EquationSource> <EquationSource Format="MATHML"><math> <mrow> <mi>λ</mi> <mo>-</mo> <mi>μ</mi> </mrow> </math></EquationSource> </InlineEquation>) remains systematically underestimated due to the slight overestimation of <InlineEquation ID="IEq12"> <EquationSource Format="TEX">\(\mu \)</EquationSource> <EquationSource Format="MATHML"><math> <mi>μ</mi> </math></EquationSource> </InlineEquation>. On the whole, these results begin to clarify the statistical and structural sources of bias in diversification rate estimation and provide a general framework for improving inference under birth-death models.</p>

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Statistical and Structural Bias in Birth-Death Models

  • Jeremy M. Beaulieu,
  • Brian C. O’Meara

摘要

Accurate estimation of speciation ( \(\lambda \) λ ) and extinction ( \(\mu \) μ ) rates from phylogenetic trees is central to studies of diversification, yet it remains unclear whether commonly used estimators are unbiased. Here we examine two sources of error: (1) statistical bias in the estimators themselves, and the (2) structural bias introduced by how small trees are handled in likelihood calculations. For the Yule process, we re-derive the expected bias of the standard estimator, showing that \(\hat{\lambda }\) λ ^ underestimates \(\lambda \) λ by a factor of \((n-2)/(n-1)\) ( n - 2 ) / ( n - 1 ) . Extending to the general birth–death model, we use symbolic regression to find functional forms that minimize the bias in both \(\lambda \) λ and \(\mu \) μ . The best-performing correction for \(\lambda \) λ is identical to the Yule result, while the bias in \(\mu \) μ depends on both sample size and the estimated extinction fraction ( \(\mu /\lambda \) μ / λ ). Applying these corrections substantially improves the fit between the estimated and generating values. When these corrected estimators are used to derive other diversification-related parameters, turnover is nearly unbiased, but net diversification ( \(\lambda - \mu \) λ - μ ) remains systematically underestimated due to the slight overestimation of \(\mu \) μ . On the whole, these results begin to clarify the statistical and structural sources of bias in diversification rate estimation and provide a general framework for improving inference under birth-death models.