The aim of this project is to implement a versatile high-performance version of the BPP software. It should have the following properties:

• open-source code with an appropriate open-source license.
• 64-bit multi-threaded design that handles very large datasets.
• easy to use and well-documented.
• SIMD implementations of time-consuming parts.
• Linux, Mac and Microsoft Windows compatibility.

BPP currently implements four methods:

• Estimation of the parameters of species divergence times and population sizes under the multi-species coalescent (MSC) model when the species phylogeny is given (Rannala and Yang, 2003)

• Inference of the species tree when the assignments are given by the user (Rannala and Yang, 2017)

• Species delimitation using a user-specified guide tree (Yang and Rannala, 2010; Rannala and Yang, 2013)

• Joint species delimitation and species tree estimation (Yang and Rannala 2014)

BPP can also accommodate variable mutation rates among loci (Burgess and Yang, 2008) and heredity multipliers (Hey and Nielsen, 2004). Finally, BPP supports diploid data. Phasing is done analytically as described by Gronau et al, 2011.

Update 2019-01-02

BPP now also implements the Multispecies-coalescent-with-introgression (MSci) model (see Flouri et al, 2020), an extension of the multispecies coalescent model to incorporate introgression/hybridization. For more information on usage please see the BPP manual and/or the wiki documentation.

Update 2020-04-20

Since v4.3.8 BPP implements the following:

• Parallelized computation using POSIX threads.
• Nucleotide substitution models JC69, K80, F81, HKY, T92, TN93, F84 and GTR.
• 19 amino acid substition models (Dayhoff, LG, DCMUT, JTT, etc)
• Site rate variation (+Gamma)
• Partitioned analyses
• Relaxed clock (independent rates model and correlated clock) as described in (Zhu et al, 2015).
• Easy to use tool for creating MSci networks that accepts a species tree and a list of definitions specifying introgressions/hybridizations. For information on using those options check the BPP manual

Binary distribution Starting with version 4.1.3, binary distribution files containing pre-compiled binaries will be available as part of each release. The included executables are statically compiled whenever possible such that no library dependencies are necessary.

Binary distributions are provided for x86-64 systems running GNU/Linux, macOS (version 10.13 or higher) and Windows (64-bit, version 7 or higher).

Download the appropriate executable for your system using the following commands if you are using a Linux x86_64 system:

wget https://github.com/bpp/bpp/releases/download/v4.6.2/bpp-4.6.2-linux-x86_64.tar.gz
tar zxvf bpp-4.6.2-linux-x86_64.tar.gz

Or these commands if you using a Mac:

wget https://github.com/bpp/bpp/releases/download/v4.6.2/bpp-4.6.2-macos-x86_64.tar.gz
tar zxvf bpp-4.6.2-macos-x86_64.tar.gz

Or if you are using Windows, download and extract (unzip) the contents of this file:

https://github.com/bpp/bpp/releases/download/v4.6.2/bpp-4.6.2-win-x86_64.zip

Linux and Mac: You will now have the binary distribution in a folder called bpp-4.6.2-linux-x86_64 or bpp-4.6.2-macos-x86_64. The binary file is located in the bin subfolder, i.e. bin/bpp. We recommend making a copy or a symbolic link to the binary in a folder included in your $PATH.

Windows: You will now have the binary distribution in a folder called bpp-4.6.2-win-x86_64. The bpp executable is called bpp.exe.

Compiling from source You can either download the source distribution for a particular version or clone the repository.

Source distribution To download the source distribution from a release and build the executable and documentation, use the following commands:

wget https://github.com/bpp/bpp/archive/v4.6.2.tar.gz
tar zxvf v4.6.2.tar.gz
cd bpp-4.6.2/src
make

Cloning the repo Instead of downloading the source distribution as a compressed archive, you could clone the repo and build it as shown below.

git clone https://github.com/bpp/bpp.git
cd bpp/src
make

Compiling BPP requires that your system has GCC version 4.7 or newer, as AVX and AVX-2 optimized functions are compiled even if your processor does not support them. This is fine, as BPP will automatically select the right instruction set that your processor supports at run-time. This means, you can compile on one system, and run BPP on any other compatible system.

However, if your compiler is older than 4.7, you will get errors such as:

cc1: error: unrecognized command line option "-mavx2"

or

cc1: error: unrecognized command line option "-mavx"

If your compiler is GCC 4.6.x then you can compile BPP using:

make clean
make -e DISABLE_AVX2=1

In case your compiler is older than GCC 4.6 then compile using:

make -e DISABLE_AVX2=1 DISABLE_AVX=1

You can check your compiler version with:

gcc --version

After creating the control file, one can run BPP as follows:

bpp --cfile [CONTROL-FILE]

If you would like to resume a checkpoint file, please run:

bpp --resume [CHECKPOINT-FILE]

If you would like to run the simulator (previously MCcoal), please run:

bpp --simulate [CONTROL-FILE]

If you would like to run the MSci network generator, please run:

bpp --msci-create [DEFS-FILE]

If you would like to only summarize the results of an analysis, please run:

bpp --summary [CONTROL-FILE]

For an example of a DEFS-FILE see the MSci generator notes

More documentation regarding control files, will be available soon on the wiki.

The most up-to-date documentation of BPP is bppDOC.pdf distribution together with BPP.

A tutorial on BPP was recently published as a book chapter: A Tutorial on the Use of BPP for Species Tree Estimation and Species Delimitation

For information on the MSci model please read:

• Flouri T., Jiao X., Rannala B., Yang Z. (2020) A Bayesian Implementation of the Multispecies Coalescent Model with Introgression for Phylogenomic Analysis. Molecular Biology and Evolution, 37(4):1211-1223.

• Jiao X., Flouri T., Rannala B., Yang Z. (2020) The Impact of Cross-Species Gene Flow on Species Tree Estimation. Systematic Biology (in press). doi:10.1093/sysbio/syaa001

Please cite the following publication if you use BPP:

Flouri T., Jiao X., Rannala B., Yang Z. (2018) Species Tree Inference with BPP using Genomic Sequences and the Multispecies Coalescent. Molecular Biology and Evolution, 35(10):2585-2593. doi:10.1093/molbev/msy147

Please note, citing the corresponding of the four underlying methods, may also be appropriate.

If you use the MSci model please also cite:

Flouri T., Jiao X., Rannala B., Yang Z. (2020) A Bayesian Implementation of the Multispecies Coalescent Model with Introgression for Phylogenomic Analysis. Molecular Biology and Evolution, 37(4):1211-1223. doi:10.1093/molbev/msz296

The code is currently licensed under the GNU Affero General Public License version 3.

• Tomáš Flouri
• Bruce Rannala
• Anna Nagel
• Ziheng Yang

Special thanks to:

• Paul M. Hime
• Jiayi Ji
• Paschalia Kapli
• Mario dos Reis Barros
• Yuttapong Thawornwattana,

for testing and bug reports.

• Burgess R., Yang Z. (2008) Estimation of hominoid ancestral population sizes under Bayesian coalescent models incorporating mutation rate variation and sequencing errors. Molecular Biology and Evolution, 25(9):1979-1994. doi:10.1093/molbev/msn148

• Flouri T., Carrasco FI, Darriba D., Aberer AJ, Nguyen LT, Minh BQ, Haeseler A., Stamatakis A. (2015) The Phylogenetic Likelihood Library. Systematic Biology, 64(2):356-362. doi:10.1093/sysbio/syu084

• Flouri T., Jiao X., Rannala B., Yang Z. (2018) Species Tree Inference with BPP using Genomic Sequences and the Multispecies Coalescent. Molecular Biology and Evolution, 35(10):2585-2593. doi:10.1093/molbev/msy147

• Flouri T., Jiao X., Rannala B., Yang Z. (2020) A Bayesian Implementation of the Multispecies Coalescent Model with Introgression for Phylogenomic Analysis. Molecular Biology and Evolution, 37(4):1211-1223. doi:10.1093/molbev/msz296

• Gronau I., Hubisz MJ, Gulko B., Danko CG, Siepel A. (2011) Bayesian inference of ancient human demography from individual genome sequences. Nature Genetics, 43(10):1031-1035. doi:10.1038/ng.937

• Hey J., Nielsen R. (2004) Multilocus methods for estimating population sizes, migration rates and divergence time, with applications to the divergence of Drosophila pseudoobscura and D. persimilis. Genetics, 167(2):747-760. doi:10.1534/genetics.103.024182

• Jiao X., Flouri T., Rannala B., Yang Z. (2020) The Impact of Cross-Species Gene Flow on Species Tree Estimation. Systematic Biology (in press). doi:10.1093/sysbio/syaa001

• Rannala B., Yang Z. (2013) Improved reversible jump algorithms for Bayesian species delimitation. Genetics, 194:245-253. doi:10.1534/genetics.112.149039

• Rannala B., Yang Z. (2017) Efficient Bayesian Species Tree Inference under the Multispecies Coalescent. Systematic Biology, 66(5):823-842. doi:0.1093/sysbio/syw119

• Yang Z., Rannala B. (2003) Bayes Estimation of Species Divergence Times and Ancestral Population Sizes using DNA Sequences From Multiple Loci. Genetics, 164:1645-1656. Available at: http://www.genetics.org/content/164/4/1645.long

• Yang Z., Rannala B. (2010) Bayesian species delimitation using multilocus sequence data. Proceedings of the National Academy of Sciences, 107(20):9264-9269. doi:10.1073/pnas.0913022107

• Yang Z., Rannala B. (2014) Unguided species delimitation using DNA sequence data from multiple loci. Molecular Biology and Evolution, 31(12):3125-3135. doi:10.1093/molbev/msu279