Mammalian sexual selection under the lens of adaptation-inertia methods:

A reCAP of software
1 2 2 2 1 3
Jesualdo A. Fuentes-G. *, Bayu Brahmantio , Hao Chi Kiang , Krzysztof Bartoszek , Janna Fierst , Thomas F. Hansen , Jason Pienaar 1
1Florida International University, 2Linköpings Universitet, 3Universitetet i Oslo
INTRODUCTION REGIME SOFTWARE

Given their role in sperm production, testes are expected to be relatively
larger in systems with higher levels of female promiscuity where male-male
competition is higher[1,2,3]:
An evolutionary test of this hypothesis requires studying the historical effects
of mating system on relative testes mass. A way to accomplish this involves
reconstructing (here, under maximum likelihood[4]) the levels of sperm
competition[3] on a time-calibrated phylogenetic tree[5]:
Sperm competition allows for tackling this association as a sexual selection
problem. The adaptation-inertia method[6,7] is an ideal framework to analyze
it because, unlike methods based solely on Brownian motion (BM), they
allow for evolution towards specific states without accumulating variance
unboundedly with time. It accomplishes this by using a mean reverting
stochastic Ornstein-Uhlenbeck (OU) process:

BM

x
x

x
Sperm

x
T

x
High competition level Low dXT = σ2 * dWT

x
OU

x
x
x x
Polyandry

x
Polygyny μ

x
Polygynandry Mating systems

x
Monogamy dXT = σ2 * dWT +  (θ - XT)

x
Promiscuty T

x
x
Hypothesized relative

x
x
x x
Large testes mass Small
θ
We offer a compilation of software developed under this adaptation-inertia
We test this hypothesis in mammals by using continuous measures of testes We use this mapping as a historical regime to understand the evolutionary framework and present some of their key findings for the mammalian sexual
mass and body mass (reported in g[3] ) under different variable systems association between mating system and relative testes size in mammals selection hypothesis (Bold: no longer available, Underline: not developed in
R [8])

Discretely mapped levels of sperm Discretely mapped levels of sperm Discretely mapped levels of sperm Discrete shift configurations inferred
SELECTIVE competition used to estimate different RATE competition used to estimate different MIXED competition used in conjunction with ESTIMATED from the data in absence of prior
primary optima evolutionary rates continuously evolving variables evolutionary hypotheses

COMPARE, pmc, maticce Brownie (RBrownie), OUwie slouch surface, phylolm, l1ou, bayou
High

High

CONCLUDING REMARKS
High

σ2 High = 0.12
• The growth of phylogenetic comparative methods in recent years is
well reflected in adaptation-inertia methods: From a univariate
Testes mass (ln[g])

approach with a binary regime, the framework has grown towards

multivariate formulations that allow mixtures of continuous and
Low

Low
Low

discrete multistate regimes, estimates of the amount and location of

UNIVARIATE

σ2 Low = 0.26 shifts, and mosaics of parameters and evolutionary models across
the tree.
σ2 = 0.19
• All reviewed implementations have made valuable contributions to
the development of the framework and offer complementary
insights on phenotypic evolution.
θ θ Testes mass (ln[g])
Body mass (ln[g])
Body mass (ln[g])
• σ2High < σ2Low : Increased disparity with shift • The ground covered so far, as well as the ground yet to be covered,
• Variable: Testes mass (ln[g]) / Body mass (ln[g])
• θHigh > θLow: consistent with sexual selection hypothesis • Negative allometry (slope between 0-1) with body mass • Estimated shifts (7) lead to remarkably small (e.g. makes the adaptation-inertia a reliable and exciting framework to
• θHigh > θLow: Consistent with sexual selection hypothesis modeled as BM Eurasian shrew) or large (muskrat) mammals tackle macroevolutionary questions about adaptation.
• Limited suport: 95% CI overlap extensivelly • Limited suport: 95% CI overlap extensivelly
• θHigh > θLow: consistent with sexual selection hypothesis • Estimated regime may reflect remnants of scaling
• Strong suport: 95% CI little overlap effects
PhylogeneticEM, PCMFit (PCMBase), ACKNOWLEDGMENTS
ouch mvMORPH mvSLOUCH (PCMBase) GLinvCI, PCMkappa
• Pienaar lab for discussions.
• Art: Laura Fuentes (main: title), Bing Image Creator (niches & background).
• This material is based on work supported by the National Science
Foundation under grant no. 2225683 to JP; ELLIIT Call C and the Swedish
Research Council (Vetenskapsrådet) grant no. 2017-04951 to KB and BB.
Testes mass (ln[g])
Testes mass (ln[g])

BM

OU REFERENCES
MULTIVARIATE

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primates. Nature 293(5827):55-57.
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Mammalogy 67(1):1-22.
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mammals. Journal of Evolutionary Biology 22(11):2215-2221.
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characters on phylogenies. Systematic Biology 48(3):612-622.
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diversification. Molecular Biology and Evolution 32(4):835-845.
Body mass (ln[g])
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• Unequal rate matrices and root states (BM) tissue hypothesis: trade-off between brain and body mass 1351.
• Bivariate macroevolutionary landscape with two • [7] Hansen TF, Pienaar J, Orzack SH. 2008. A comparative method for studying adaptation to a randomly
adaptive peaks: ▲ & ▲ • ● Slightly tighter correlation (0.81 > 0.75) but similar • Positive conditional expectations except for brain and • Evolutionary process shifts (OU → BM) in the Old evolving environment. Evolution 62(8):1965-1977.
rates (variance) body mass (consistent with hypothesis) • [8] R Development Core Team. 2022. R: a language and environment for statistical computing. Version 3.1.1. R
• ▲ Larger size Aligned with sexual World monkeys (Cercopithecidae) Foundation for StatisticalComputing, Vienna. http://www.R-project.org.
• ● Similar size but larger testes (caution with adaptive • Negative conditional expectations close to zero (support • Prevalence of uni-male groups and pre-copulatory male
• ▲ Larger testes selection hypothesis
interpretations, as these estimates do not reflect optimal for hypothesis is weak) competition may have shifted the selective pressures on
states) sperm size for this group *jefuente@fiu.edu