Genetic Analysis of Populations with Mixed Reproduction

Population genetic analyses for hierarchical analysis of partially clonal populations built upon the architecture of the 'adegenet' package. Originally described in Kamvar, Tabima, and Grünwald (2014) with version 2.0 described in Kamvar, Brooks, and Grünwald (2015) .

In Development: Build Status Coverage Status

CRAN Status: CRAN version CRAN check status

Usage: Downloads from Rstudio mirror

What is poppr?

Poppr is an R package designed for analysis of populations with mixed modes of sexual and clonal reproduction. It is built around the framework of adegenet's genind and genlight objects and offers the following implementations:

  • clone censoring of populations at any of multiple levels of a hierarchy
  • convenient counting of multilocus genotypes and sub-setting of populations with multiple levels of hierarchy
  • define multilocus genotypes
  • calculation of indices of genotypic diversity, evenness, richness, and rarefaction
  • drawing of dendrograms with bootstrap support for genetic distances
  • drawing of minimum spanning networks for genetic distances
  • calculation of the index of association (Index of association) or (Standardized index of association)
  • batch processing on any server that has R ( ≥ 2.15.1) installed
  • calculation of Bruvo's distance for microsatellite (SSR) markers (implemented in C for speed)
  • import of data from and export to GenAlEx

New in version 2.0:

  • handling of genomic SNP data
  • custom multilocus genotype definitions
  • collapse multilocus lineages by genetic distance
  • calculate reticulate minimum spanning networks
  • calculate index of association in a sliding window across snps
  • bootstrapping of MLG diversity statistics
  • interactive exploration of minimum spanning networks
  • and more!

For full details, see the NEWS file or type in your R console:

news(Version >= "2.0.0", package = "poppr")


If you use poppr at all, please specify the version and cite:

analysis of populations with clonal, partially clonal, and/or sexual reproduction. PeerJ 2:e281

If you use poppr in a presentation please mention it as the poppr R package, specify the version, and use our logo: (png) | (svg).

Additionally, if you use any following functionalities:

  • minimum spanning networks with reticulation
  • collapsing multilocus genotypes into multilocus lineages with mlg.filter()
  • custom multilocus genotype definitions with mlg.custom()
  • index of association for genomic data with win.ia() or samp.ia()
  • bootstrapping any genetic distance with genind, genlight, or genpop objects with aboot()

Please also cite:

Kamvar ZN, Brooks JC and Grünwald NJ (2015) Novel R tools for analysis of genome-wide population genetic data with emphasis on clonality. Front. Genet. 6:208. doi: 10.3389/fgene.2015.00208

You can obtain citation information in R by typing:

citation(package = "poppr")



Binary versions for mac and windows are available for R ≥ 2.15.1 here.

To install, make sure R is at least version 2.15.1 (the authors recommend ≥ 3.0), and in your console, type:


If you want the absolute latest version of poppr, see about installing from github below.

Stable and Development versions

To install this package from github, make sure you have the following:

For Linux users, make sure that the function getOption("unzip") returns "unzip" or "internal". If it doesn't, then run options(unzip = "internal").

Once you have devtools and a C compiler installed, you can use the install_github() function to install the current version from github.

Stable version

This release will contain bug fixes and new, documented, and stable features that will be included in future releases. Note: if you don't have LaTeX installed, you should set build_vignettes = FALSE.

devtools::install_github(repo = "grunwaldlab/poppr", build_vignettes = TRUE)

Unstable/Development versions

All new features in testing will be released on different branches. These features will be in various stages of development and may or may not be documented. Install with caution. The below command would install features on the the branch called "devel". Note that these branches might be out of date from the master branch.

devtools::install_github(repo = "grunwaldlab/[email protected]", build_vignettes = TRUE)

Help / Documentation

R documentation

To access a descriptive index of help files in poppr, type in your console:



A few vignettes have been written for poppr:

Title Command
Algorightms and Equations vignette("algo", "poppr")
Data import and manipulation vignette("poppr_manual", "poppr")
Multilocus Genotype Analysis vignette("mlg", "poppr")

User Group

Users who have any questions/comments/suggestions regarding any version of poppr (stable or development) should direct their comments to the Poppr google group


In Spring of 2014, Dr. Niklaus J. Grünwald, Dr. Sydney E. Everhart and Zhian N. Kamvar wrote a primer for population genetic analysis in R located at


Please note that this project is released with a Contributor Code of Conduct. By participating in this project you agree to abide by its terms. If you wish to contribute code to poppr, please fork the repository and create a pull request with your added feature.


poppr 2.8.2


poppr 2.8.1



  • aboot() documentation was updated to add the citation and make clear its purpose and limitations.


  • DOIs have been hyperlinked to instead of (#188, @katrinleinweber)

poppr 2.8.0


  • win.ia() now has more consistent behavior with chromosome structure and will no longer result in an integer overflow. (see Thanks to @MarisaMiller for the detailed bug report.
  • plot_filter_stats() will plot stats if supplied a list of thresholds.


  • win.ia() may result in slightly different results because of two changes:
    1. The windows will now always start at position one on any given chromosome. This will result in some windows at the beginning of chromosomes having a value of NA if the first variant starts beyond the first window.
    2. Windows are now calculated for each chromosome independently. The previous version first concatenated chromosomes with at least a window-sized gap between the chromosomes, but failed to ensure that the window always started at the beginning of the chromosome. This version fixes that issue. (see


  • The chromosome_buffer argument for win.ia() has been permanently set to TRUE and deprecated as it is no longer used.


  • poppr.amova() will now handle genlight/snpclone objects. See for details.

  • bitwise.dist() now has two new options: euclidean and scale_missing. When both of these are set to TRUE, the distance measured will be Euclidean scaled for the amount of missing data in each comparison. This matches the output of base R's dist() function at a fraction of time and memory. See for details.

  • make_haplotypes() is now a generic defined for both genind and genlight.

  • genind2genalex() will no longer write to "genalex.csv" by default. Instead, it will warn the user and write to a temporary file. See for details.

  • genind2genalex() now has an overwrite parameter set to FALSE to prevent accidental overwriting of files.

  • win.ia() has a new argument name_window, which will give each element in the result the designation of the terminal position of that window. Thanks to @MarisaMiller for the suggestion!

  • pair.ia() can now calculate p-values via permutations. (See for details)


  • cutoff_predictor() was added to the MLG vignette


poppr 2.7.1


  • Missing documentation for poppr.amova() has been added.
  • Polysat is now listed in imports.

poppr 2.7.0


  • make_haplotypes() will split your data into pseudo-haplotypes for use in AMOVA-like analyses. This was a previously internal function, but has been promoted to a user-facing function in this version.

  • as.genambig() will convert genind/genclone objects to Polysat's "genambig" class. Note that polysat must be installed for this to work.


  • AMOVA will now default to using euclidean distance. This affects all calculations made with within = FALSE or filter = TRUE without a user-supplied distance. This will not have affect those with haploid or diploid data using within = TRUE. The dissimilarity distance is equivalent to a squared euclidean distance for haploid genotypes, but not for any higher ploidy. Those using filter = TRUE without specifying a distance should use a euclidean threshold. This should not be an issue for those who simply want to group isolates with missing data, however as a zero distance is the same for euclidean and dissimilarity. Thanks goes to Patrick Meirmans for alerting me to this error.


  • AMOVA will now calculate within-individual variance for polyploid data.


  • printing of AMOVA will now better handle any changes in methods from pegas or ade4.

poppr 2.6.1


poppr 2.6.0


  • The new function boot.ia() is conceptually similar to resample.ia(), except it resamples with replacement.


  • The function resample.ia() now can resample individuals weighted by their Psex value.
  • The minimum spanning networks will now scale nodes by area instead of radius. This gives a more accurate picture of the differences between MLGs. See for details.
  • A legend for samples/node is now added to all minimum spanning networks. See for details.
  • The imsn() option for node size scale has been changed to a slider.



  • The minimum version of igraph has been set to 1.0.0.


  • The MSN is now plotted last in plot_poppr_msn() so additional legends can be added if necessary.

poppr 2.5.0


  • Identified in, Bruvo's distance will now consider all possible combinations of ordered alleles in the calculation under the genome addition and loss models for missing data. This will affect those who have polyploid data that contain more than one missing allele at any genotype

    To facilitate comparison, the global option old.bruvo.model, has been created. By default it is set to FALSE, indicating that poppr should use the ordered allele combinations. If the user wants to use the method considering unorderd allele combinations, they can set options(old.bruvo.model = TRUE)

    It must be repeated that this does not affect haploid or diploid comparisons, those that use the infinite alleles model, or those who do not have more than one missing allele at any genotype.


  • The warning for a short repeat length vector for Bruvo's distance is deprecated and will become an error in the future
  • jack.ia is deprecated in favor of resample.ia for clarity.



  • The internal plotting function for mlg.table now uses tidy evaluation for dplyr versions > 0.5.0
  • The package reshape2 was removed from imports and replaced with base functions (see for details)


  • Due to the migration to dplyr version 0.7.0, poppr now imports the "!!" operator from the rlang package

poppr 2.4.1


poppr 2.4.0


  • jack.ia() will randomly jackknife your sample to a specified n (default is the number of MLG), and calculate the index of association over multiple iterations, giving a distribution of possible values at a given sample size.


  • The function mlg.table() gains new parameters, "color" and "background". The "color" parameter will create a single barplot with colors representing populations while the "background" parameter will create a background plot showing the abundance of MLGs across populations within the facets.
  • The function win.ia() will now take into consideration chromosomal coordinates when constructing windows. It has additionally acquired a new parameter chromosome_buffer, which allows the user to specify whether or not the window should be limited to within chromosomes.


  • calculation of MLGs for snpclone and genlight objects will be performed via distance-based methods by default. This is in contrast to the previous behavior where individuals were assumed to have unique genotypes. see for details.
  • An error will be thrown when attempting to use mlg.crosspop() with an object that has < 2 populations.
  • genotype_curve() will now remove monomorphic loci before calculation by default as these loci misleadingly influence the shape of the curve. This will change the shape of the curve if you have monomorphic loci. This change IS optional via the drop and dropna parameters, but it is not recommended to change these parameters.
  • The calculation for psex() has changed to be more accurate when using method = "multiple". It also gains the ability to use several values of G, one for each population. Documentation for psex() has also been improved. For details of the change, see



  • The documentation for bitwise.dist() clarifies the role of the differences_only flag (see
  • Interruptions in C code is now handled gracefully via R_CheckUserInterrupt(). The benefit is that long-running calculations are interrupted near instantly, but at the cost of a few more milliseconds of computation time. (see
  • Bruvo's distance now has complete tests for recursion as of commit 4e4fa40d16

poppr 2.3.0



  • There is now a plot parameter for the genotype curve to enable or suppress plotting.
  • Progress bars are now automatically suppressed when running non-interactively. to turn them on when running non-interactively, use options(poppr.debug = TRUE).
  • The progress bar for ia() and poppr() will now show estimated time. This is from dplyr's progress_estimated().


  • The hist argument in the ia() is deprecated in favor of plot.
  • The x axis for the genotype_curve() plot is now numeric, allowing you to fit a smoothing function over the points without having to use the hack geom_smooth(aes(group = 1)). This is thanks to Kara Woo for pointing this out on twitter (
  • The "show" method for genclone objects now delimits populations and strata by a comma, avoiding confusion with multi-word population names. Thanks to @knausb for the fix in
  • Documentation for poppr.amova now contains a note about significance testing with the ade4 function randtest.amova.


poppr 2.2.1




poppr 2.2.0


  • incomp() will check your data to see if there are any incomparable samples.


  • Threshold argument added to filter_stats() (see
  • The pipe operator (%>%) is now exported from magrittr to make chaining commands easier.
  • mlg.filter() can now return multiple statistics.
  • The user can now control the size of the labels in the index of association plots with the labsize and linesize arguments in the plot method.
  • private_alleles() gains a drop argument.
  • recode_polyploids() can now take haplodiploid data.


  • An issue that caused errors in the imsn() code output was fixed (see
  • Caught bug where the mlg.filter() assignment method was using nei.dist() instead of diss.dist() when no distance was specified.
  • A bug that resulted in significantly negative values from bitwise.ia() with large sample sizes was fixed. Spotted by @knausb (see
  • Fixed issue where plot_filter_stats() wasn't displaying the full range of MLGs
  • Color vectors are now correctly parsed when passed to msn functions (see for details. Long color vectors are now accepted, albiet with warning.
  • An issue where mll.reset() did not reset non-MLG class objects in the mlg slot was fixed.


  • Documentation for mlg.filter() was clarified and updated with more examples.
  • The vignette "Migration from poppr version 1" has been removed.
  • Previously deprecated *hierarchy() functions have been removed.

poppr 2.1.1


  • imsn() now has collapsible side panels
  • nmll() and mll() will now handle genind and genlight objects
  • rraf() now gives options for minor allele correction encompassed in the internal function rare_allele_correction(). This extends also to pgen() and psex(), which must correct minor allele frequencies by default. See for details.


  • mlg.filter() now defaults to using diss.dist()
  • default threshold for filter_stats() is now 1e+6
  • mlg.filter() now returns a list instead of a pairlist
  • the "hist" argument for poppr() is now deprecated in favor of "plot"
  • documentation improvements
  • the show method for snpclone objects now looks distinct from genlight
  • genlight objects no longer get passed to missingno() in filter_stats()
  • filter_stats() now returns invisibly when plot = TRUE; see for details.


poppr 2.1.0


  • win.ia and samp.ia gain a significant speedup thanks to Jonah Brooks implementing the code in C.
  • The internal code for the genotype_curve has been implemented in C for a 10x increase in speed.


  • poppr.msn, bruvo.msn, and plot_poppr_msn gain the ability to take character vectors for color palettes. See issue #55 ( for details.
  • plot_poppr_msn returns the modified graph.
  • All functions related to Bruvo's distance can now take a named vector of repeat lengths in any order. See issue #61 ( for details.
  • aboot gains the argument strata so that you can automatically convert genind to genpop.
  • genotype_curve can now take in loci objects from pegas.
  • You can now specify the maximum number of loci to analyze in genotype_curve.
  • filter_stats can now optionally plot a histogram in the background.
  • bruvo.dist can now optionally return distance matrices by locus. This is addresed in issue #60 (
  • aboot can now handle matrices as previously specified in the documentation.
  • aboot can now take custom functions to calculate distance for genlight objects.
  • poppr.amova can now perform amova using the pegas implementation.


  • rrmlg will calculate round-robin multilocus genotypes for each locus.
  • rraf will calculate round-robin allele frequencies for each locus.
  • pgen will calculate the probabilities of observed genotypes.
  • psex will calculate the probability that an observed genotype will be observed more than once by chance.


  • because we're through being cool.


  • Documentation for genclone and snpclone classes are more coherent.
  • Accessors added for internal MLG objects (for developers).
  • The genotype accumulation curve displays the iteration and locus number instead of a progress bar.
  • The genotype accumulation plot is now scaled from 0 to the number of observed mlgs.
  • Documentation for poppr.amova no longer references the "hierarchy" slot.


  • Single locus data sets can now be read in with read.genalex. This was brought up in issue #58 (Thanks to Nick Wong for spotting it).
  • A bug in informloci where the MAF argument wasn't being applied to P/A data has been fixed.
  • Printing of genclone objects with mixed ploidies previously reported erroneously due to a sorting error. This has been fixed.
  • Edge case where a missing cell in the genalex matrix was interpreted as a literal "NA" was fixed.
  • msn functions now return nodes that are correctly named. See Issue #66 ( for details.

poppr 2.0.2


  • Definition of Hexp was fixed. It originally was mis-calculated, inflating the metric. It is now correctly calculated and documented. More information at issue #47

poppr 2.0.1


  • Memory leak with Bruvo's distance was fixed by @JonahBrooks in 90facb4 (issue #40)
  • Cutoff field now works for distances other than dissimilarity in imsn (issue #41)
  • Switching between data sets no longer shows an error in imsn
  • read.genealex can now correctly import missing data for diploids (issue #42)


  • Startup message now tells you if poppr was compiled with OMP support.

poppr 2.0.0


  • poppr has moved to version 2.0 due to adegenet's recent update. The hierarchy slot introduced in version 1.1 is now being moved to adegenet and renamed strata. For maximum backwards compatibility, all of the hierarchy methods still exist, but they are deprecated and will print a warning with the proper function to use. If you were accessing the hierarchy slot without using the *hierarchy() methods, your code will fail as the hierarchy slot now should only contain a formula object.


  • poppr now imports elements from dplyr and shiny
  • As required as of 2015-06-29, poppr now explicitly imports: stats, graphics, grDevices, and utils


  • poppr now suggests the cowplot, poweRlaw, and polysat packages.


  • A data set called Pram containing SSR genotypes from the Sudden Oak Death pathogen Phytophthora ramorum (Kamvar, 2015)


  • refreshing!


  • The default plot for the index of association will now be a single histogram. The user has the option to visualize the standardized index of association (index = "rbarD", default) or the classic index of association (index = "Ia"). If the user uses the function ia with the argument valuereturn = TRUE, then the resulting object can be plotted with the plot function.
  • The function poppr will now plot all populations in a single faceted plot instead of one plot per population.
  • aboot and bruvo.boot will now be able to utilize any function to generate trees (suggested in issue #18).
  • The mlg slot in the genclone object can now optionally hold an MLG class object. This object will contain different definitions of multilocus genotypes, allowing the user to switch between observed, custom, and mlgs defined given a genetic distance threshold.
  • minimum spanning network functions gain the ability to include reticulations using the option include.ties
  • minimum spanning network functions gain the ability to collapse multilocus genotypes by genetic distance with the option threshold.
  • poppr.amova gains the ability to filter multilocus genotypes before calculation.
  • informloci gains the argument "MAF", which allows the specification of a minor allele frequency cutoff in addition to the cutoff argument. Examples have been updated.
  • aboot can now take genlight objects.
  • poppr.msn and plot_poppr_msn can now take genlight objects.
  • plot_poppr_msn now gives users the option to exclude the legends.
  • Default plotting for mlg.table will no longer produce one plot per population. It will now produce a single ggplot object for all populations. Note that the bars are no longer colored by count.
  • poppr will no longer calculate "Hexp". Instead, Simpson's index will be calumniated, but the old index can be retrieved by using (N/(N - 1))*lambda.
  • poppr can now take any statistic that can be calculated from a table of multilocus genotype counts.
  • genind2genalex gains the ability to selectively write different strata.


  • mlg.filter will contract multilocus genotypes given a genetic distance and threshold using one of three algorithms. It can report statistics such as the multilocus genotypes returned, the number of samples within each multilocus genotype, the thresholds at which multilocus genotypes were collapsed, and the genetic distance matrix that represents the new multilocus genotypes.
  • filter_stats will show you graphical output of all the algorithms in mlg.filter.
  • cutoff_predictor will predict the cutoff threshold from mlg.filter.
  • bitwise.dist can efficiently calculate absolute genetic distance for genlight objects.
  • mll "multilocus lineages" is a new replacement for mlg.vector which gains the functionality of selecting the multilocus genotype definition from the mlg slot.
  • nmll counts the number of multilocus lineages
  • mll.custom allows the user to define custom multilocus genotypes.
  • mll.levels allows the user to edit the names of custom multilocus genotypes.
  • poppr_has_parallel will return TRUE if poppr was built with OpenMP parallel library.
  • win.ia calculates windows of \bar{r}_d along genlight chromosomes.
  • samp.ia calculates \bar{r}_d for genlight object by randomly sampling a user-defined number of SNPs.
  • test_replen will test repeat lengths of microsatellite markers for consistency.
  • fix_replen will fix inconsistent repeat lengths for microsatellite markers.
  • diversity_stats returns a matrix containing diversity statistics. Defaults to 4 found in poppr, but can be extended to any statistic that can be calculated on a vector of MLG counts.
  • diversity_boot will bootstrap a MLG matrix over the statistics specified for get_stats. Can also perform rarefaction bootstrap.
  • diversity_ci will calculate and plot confidence intervals for bootstrap resampling of an MLG matrix. This includes rarefaction to the smallest sample size.
  • imsn provides an interactive shiny interface for construction of minimum spanning networks.
  • pair.ia will calculate the index of association for pairs of loci and plot heatmaps.


  • snpclone is an extension of the genlight object that acts very much like genclone. It contains an mlg slot.
  • MLG is an internal class that lives inside the mlg slot of snpclone and genclone objects. It allows the user to easily switch between multilocus genotype definitions.


  • I was going too fast to count.

poppr 1.1.5


  • Fixed internal bug for fix_negative_branch when only one branch had a negative edge.
  • Fixed bug in diss.dist where a single locus would return an error.
  • Fixed bug in poppr.amova where a single locus would return an error due to repool_haplotypes.
  • Fixed bug from the future! mlg.table will now return a matrix all the time (Fix #25).

poppr 1.1.4


  • Fixed an internal bug that fails only on Windows OS.

poppr 1.1.3


  • new arguments to plot_poppr_msn to allow for easier manipulation of node sizes and of labeling
  • read.genalex can now take read text connections as input. Addresses issue #8
  • users can now specify cutoff for missing values in aboot


  • Fixed issue where monomorphic loci would cause an error in recode_polyploids
  • Fixed logical error that would cause The infinite alleles model of Bruvo's distance to inflate the distance. (Found by Michael Metzger. Addresses issue #5).
  • AMOVA can now take subset genclone objects (Addresses issue #7).
  • in mlg.table, the mlgsub argument will now subset by name instead of index (fixed in #7).
  • Fixed issue for neighbor-joining trees where the internal function to fix negative branch lengths was accidentally shuffling the corrected branches. Addresses issue #11.
  • diss.dist can now be used with aboot


  • info_table will print a discrete scale as opposed to colorbar when type = "ploidy"
  • attempted to make model choices for Bruvo's distance more clear in the documentation

poppr 1.1.2


  • Fixed memory allocation bug (Further addresses issue #2).
  • Memory allocated in C function bruvo_dist is now properly freed.

poppr 1.1.1


  • Fixed bug where the loss and add options for Bruvo's distance were switched.
  • Fixed illegal memory access error by UBSAN. Made memory management of internal C functions more sane. (Addresses issue #2).
  • Fixed directional quotes and em-dashes produced by Mavericks (Addresses issue #3).

poppr 1.1.0


  • Polyploids with ambiguous genotypes are now supported in poppr. See documentation for recode_polyploids for details.
  • Calculations of Bruvo's distance now features correction for partial missing data utilizing genome addition and genome loss models as presented in Bruvo et al. 2004.
  • diss.dist now has options to return raw distances and a matrix instead of a dist object.
  • read.genalex now has the option to import as a genclone object. This is the default action.
  • poppr.all will be able to analyze lists of genind or genclone objects.
  • ia now has the argument valuereturn which will return the sampled data.
  • [bruvo,poppr].msn functions now give the user the choice to show the graph.
  • bruvo.boot has a cleaner plot style.


  • The genclone object is a new extension of the genind object from adegenet. This object contains slots containing population hierarchies and multilocus genotype definitions and will work with all analyses in adegenet and poppr.


  • [get,set,name,split,add]hierarchy - functions that will manipulate the hierarchy slot in a genclone object utilizing hierarchical formulae as arguments for simplification.
  • setpop will set the population of a genclone object utilizing model formulae regarding the hierarchy slot.
  • as.genclone will automatically convert genind objects to genclone objects.
  • is.genclone checks the validity of genclone objects.
  • poppr.amova will run amova on any hierarchical level. This also includes the feature to run amova on clone censored data sets. It utilizes the ade4 version of amova.
  • info_table will calculate missing data per population per locus or ploidy per individual per locus and gives the user the option to visualize this as a heatmap.
  • locus_table will calculate diversity and evenness statistics over all loci in a genind or genclone object.
  • *.dist functions will calculate Nei's distance, Rogers' Distance, Edwards' Distance, Reynolds' Distance, and Provestis' Distance.
  • aboot will allow the user to create bootstrapped dendrograms for ANY distance that can be calculated on genind or genpop objects.
  • plot_poppr_msn will plot minimum spanning networks produced with poppr.
  • private_alleles will give information about the presence of private alleles within a genind or genclone object.
  • recode_polyploids will take in a polyploid genind/genclone object (with missing alleles coded as extra zero-value allele) and recode them to have frequencies relative to the observed number of alleles.
  • genotype_curve will create a genotype accumulation curve for increasing number of loci.
  • will return a list indicating the samples belonging to a specific multilocus genotype.


  • Pinf - a data set of 86 isolates from different populations of the late blight pathogen, Phytophthora infestans. Provided by Erica Goss
  • monpop - a large data set of 694 Monilinia fructicola isolates from a single orchard over three years. Provided by Sydney E. Everhart


  • Not really.


  • poppr no longer depends on pegas.
  • ade4 and reshape2 are now explicitly required.


  • default shuffling algorithm has been implemented in C to increase speed.
  • output of the mlg functions are now represented as integers to decrease their size in memory.
  • mlg.matrix is now calculated faster utilizing R's internal tabulating capabilities.
  • The function poppr will no longer return rounded results, but rather is printed with three significant digits.


  • Added unit tests.
  • The poppr user manual has been shortened to only include instructions on data manipulation.
  • A new vignette, "Algorithms and Equations" gives algorithmic details for calculations performed in poppr.

poppr 1.0.7


  • Updated README to include link to poppr google group.


  • Made last bug fix more stable (corrected on ape side).

poppr 1.0.6


  • Fixed bug for users who have downloaded ape version 3.1 or higher where bruvo.boot would throw an error.


  • Updated citation information.

poppr 1.0.5


  • The default shuffling algorithm for calculating the index of association has changed from multilocus-style sampling to permutation of alleles. All of the 4 methods are available, but new assignments are as follows: Method 1: permute alleles, Method 2: parametric bootstrap, Method 3: non-parametric bootstrap, Method 4: Multilocus-style sampling. Previously, Multilocus was 1 and the rest followed in the same order. There should be no compatibility issues with this change. Functions affected: ia, poppr shufflepop


  • Bootstrapping algorithm for bruvo.boot function was not shuffling the repeat lengths for each locus resulting in potentially erroneous bootstrap support values. This has been fixed by implementing an internal S4 class that will allow direct bootstrapping of the data and repeat lengths together.
  • An occasional error, "INTEGER() can only be applied to a 'integer', not a 'NULL'" in bruvo.boot or bruvo.dist fixed.


  • Changes to bruvo.boot allow for ever so slightly faster bootstrapping.


  • Permutations for I_A and \bar{r}_d are now visualized as a progress bar as opposed to dots.

poppr 1.0.4


  • A previous error where bootstrap values greater than 100 were reported from bruvo.boot on UPGMA trees has been fixed.
  • Fixed correction of negative branch lengths using Kuhner and Felsenstein (1994) normalization for NJ trees.


  • github repository for poppr has changed from to

poppr 1.0.3


  • Optimized internal sampling function to run up to 2x faster.
  • Utilized rmultinom function to increase speed of bootstrap sampling methods for shufflepop and ia.


  • Function informloci will remove phylogenetically uninformative loci.


  • Now importing specific functions from igraph and ape due to dependency issues.
  • Removed igraph, ape, ggplot2, and phangorn form "Dependencies", but keeping them in "Imports".


  • read.genalex will no longer insert an "X" in front of loci with numeric names.

poppr 1.0.2


  • Fixed bug in diss.dist function that would return an inflated distance for haploids.


  • Added explanation for the index of association in poppr_manual.
  • Expanded installation section to include installation instructions from github.


  • internal permutation algorithm no longer lists permutations in reverse order

poppr 1.0.1


  • Algorithm for the index of association was updated to increase speed.


  • Removed unnecessary rounding factor for missing data in bruvo.dist.
  • Corrected handling of duplicate entries for read.genind.
  • Input values that are not multiples of the specified repeat length for Bruvo's distance are now rounded (as opposed to being forced as integers).


  • Vignette updated for aesthetics and to reflect algorithmic changes.

poppr 1.0.0


  • Poppr has been confirmed to work on Linux, Mac, and Windows systems with R 3.0.0.
  • Vignette poppr_manual now has cross-references to different sections.
  • Vignette poppr_manual is quicker loading.


  • removed alpha channel from plot for resampled values of I_A and \bar{r}_d due to warnings.

poppr 0.4.1


  • getfile has a new argument, "combine", which will automatically add the path to the list of files, so they can be read without switching working directory.
  • information printed to screen from missingno and mlg.crosspop will now be wrapped to 80 characters.


  • poppr will now be able to correctly recognize GenAlEx files with both geographic and regional data.
  • calculation of the index of association on P/A data with missing values will no longer return an error.

poppr 0.4


  • mistake in Bruvo's distance where it did not correctly check for ploidy level was fixed.
  • read.genalex will be able to correctly distinguish between SNP and AFLP data.
  • read.genalex can now correctly recognize regional formatting without an extra column.


  • read.genalex will now be able to take in a file that is formatted with both regional and geographic data.
  • genind2genalex can now export xy coordinates into the GenAlEx format.
  • poppr_manual vignette now contains images of example GenAlEx files.


  • rootrot2.csv is an example of a GenAlEx file formatted with regional data.


  • function for guessing repeat lengths for Bruvo's distance moved into internal file.
  • redundancy in read.genalex was removed.
  • changed instructions in README

poppr 0.3.1


  • read.genalex will now give a warning whenever the input file is not comma delimited.

poppr 0.3


  • poppr.msn will draw a minimum spanning network for any distance matrix derived from your data set.


  • vignette now has sections describing poppr.msn, diss.dist, greycurve, and a section discussing how to export graphics.


  • The graphs output by poppr and ia will now display \bar{r}_d instead of \bar{r}_D.
  • bruvo.boot now has a dedicated quiet argument.

poppr 0.2.2


  • index of association distributions will now feature a rug plot at the bottom as a better way to visualize the distribution of the index of association from the shuffled data sets.

poppr 0.2.1


  • diss.dist will produce a distance matrix based on discreet distances.
  • greycurve will produce a grey scale adjusted to user-supplied parameters. This will be useful for future minimum spanning network functions.


  • bruvo.msn can now adjust the edge grey level to be weighted toward either closely or distantly weighted individuals.
  • bruvo.msn will now return a list giving the user the graph with all of the color, label, and weight properties so that they can plot it themselves. The legend arguments are also returned.


  • fixed shufflepop so that it will now shuffle PA markers with a specific method
  • fixed warning message mistakes in clonecorrect function.

poppr 0.2


  • Added NEWS file and will now be incrementing version number (3/15/2013)

poppr 0.1

  • First development version of poppr (2012 - 3/2013)

Reference manual

It appears you don't have a PDF plugin for this browser. You can click here to download the reference manual.


2.9.3 by Zhian N. Kamvar, 4 months ago,,

Report a bug at

Browse source code at

Authors: Zhian N. Kamvar [cre, aut] , Javier F. Tabima [aut] , Sydney E. Everhart [ctb, dtc] , Jonah C. Brooks [aut] , Stacy A. Krueger-Hadfield [ctb] , Erik Sotka [ctb] , Brian J. Knaus [ctb] , Patrick G. Meirmans [ctb] , Frédéric D. Chevalier [ctb] , David Folarin [aut] , Niklaus J. Grünwald [ths]

Documentation:   PDF Manual  

GPL-2 | GPL-3 license

Imports stats, graphics, grDevices, utils, vegan, ggplot2, ape, igraph, methods, ade4, pegas, polysat, dplyr, rlang, boot, shiny, magrittr, progressr

Depends on adegenet

Suggests testthat, knitr, rmarkdown, poweRlaw, cowplot, RClone

Suggested by AlleleShift, adegenet, dartR, vcfR.

See at CRAN