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Count opposite homozygous (OH) loci between parent-offspring pairs and Mendelian errors (ME) between parent-parent-offspring trios, and calculate the parental log-likelihood ratios (LLR).


  Pedigree = NULL,
  GenoM = NULL,
  CalcLLR = TRUE,
  LifeHistData = NULL,
  AgePrior = FALSE,
  SeqList = NULL,
  Err = 1e-04,
  ErrFlavour = "version2.0",
  Tassign = 0.5,
  Tfilter = -2,
  Complex = "full",
  Herm = "no",
  quiet = FALSE



dataframe with columns id-dam-sire. May include non-genotyped individuals, which will be treated as dummy individuals. If provided, any pedigree in SeqList is ignored.


numeric matrix with genotype data: One row per individual, and one column per SNP, coded as 0, 1, 2 or -9 (missing). See also GenoConvert.


calculate log-likelihood ratios for all assigned parents (genotyped + dummy/non-genotyped; parent vs. otherwise related). If FALSE, only number of mismatching SNPs are counted (OH & ME), and parameters LifeHistData, AgePrior, Err, Tassign, and Complex are ignored. Note also that calculating likelihood ratios is much more time consuming than counting OH & ME.


data.frame with up to 6 columns:


max. 30 characters long


1 = female, 2 = male, 3 = unknown, 4 = hermaphrodite, other numbers or NA = unknown


birth or hatching year, integer, with missing values as NA or any negative number.


minimum birth year, only used if BirthYear is missing


maximum birth year, only used if BirthYear is missing


Last year in which individual could have had offspring. Can e.g. in mammals be the year before death for females, and year after death for males.

"Birth year" may be in any arbitrary discrete time unit relevant to the species (day, month, decade), as long as parents are never born in the same time unit as their offspring, and only integers are used. Individuals do not need to be in the same order as in `GenoM', nor do all genotyped individuals need to be included.


logical (TRUE/FALSE) whether to estimate the ageprior from Pedigree and LifeHistData, or a matrix as generated by MakeAgePrior and included in the sequoia output. The AgePrior affects which relationships are considered possible: only those where \(P(A|R) / P(A) > 0\). When TRUE, MakeAgePrior is called using its default values. When FALSE, all relationships are considered possible for all age differences, except that parent-offspring pairs cannot have age difference zero, and grand-parental pairs have an age difference of at least two.


list with output from sequoia. If input parameter Pedigree=NULL, SeqList$Pedigree will be used if present, and SeqList$PedigreePar otherwise. If SeqList$Specs is present, input parameters with the same name as its items are ignored, except 'CalcLLR' and 'AgePriors=FALSE'. The list elements `LifeHist', `AgePriors', and `ErrM' are also used if present, and override the corresponding input parameters.


estimated genotyping error rate, as a single number, or a length 3 vector with P(hom|hom), P(het|hom), P(hom|het), or a 3x3 matrix. See details below. The error rate is presumed constant across SNPs, and missingness is presumed random with respect to actual genotype. Using Err >5% is not recommended, and Err >10% strongly discouraged.


function that takes Err (single number) as input, and returns a length 3 vector or 3x3 matrix, or choose from inbuilt options 'version2.9', 'version2.0', 'version1.3', or 'version1.1', referring to the sequoia version in which they were the default. Ignored if Err is a vector or matrix. See ErrToM for details.


minimum LLR required for acceptance of proposed relationship, relative to next most likely relationship. Higher values result in more conservative assignments. Must be zero or positive.


threshold log10-likelihood ratio (LLR) between a proposed relationship versus unrelated, to select candidate relatives. Typically a negative value, related to the fact that unconditional likelihoods are calculated during the filtering steps. More negative values may decrease non-assignment, but will increase computational time.


Breeding system complexity. Either "full" (default), "simp" (simplified, no explicit consideration of inbred relationships), "mono" (monogamous).


Hermaphrodites, either "no", "A" (distinguish between dam and sire role, default if at least 1 individual with sex=4), or "B" (no distinction between dam and sire role). Both of the latter deal with selfing.


logical, suppress messages


The Pedigree dataframe with additional columns:


Log10-Likelihood Ratio (LLR) of this female being the mother, versus the next most likely relationship between the focal individual and this female (see Details for relationships considered)


idem, for male parent


LLR for the parental pair, versus the next most likely configuration between the three individuals (with one or neither parent assigned)


Number of loci at which the offspring and mother are opposite homozygotes


idem, for father


Number of Mendelian errors between the offspring and the parent pair, includes OH as well as e.g. parents being opposing homozygotes, but the offspring not being a heterozygote. The offspring being OH with both parents is counted as 2 errors.

Number of SNPs scored (non-missing) for the focal individual

Number of SNPs scored (non-missing) for both individual and dam

Number of SNPs scored for both individual and sire


Sex in LifeHistData, or inferred Sex when assigned as part of parent-pair


mode of birth year probability distribution


lower limit of 95% highest density region of birth year probability distribution


higher limit

The columns 'LLRdam', 'LLRsire' and 'LLRpair' are only included when CalcLLR=TRUE. When a parent or parent-pair is incompatible with the lifehistory data or presumed genotyping error rate, the error value '777' may be given.

The columns 'Sexx', 'BY.est', 'BY.lo' and 'BY.hi' are only included when LifeHistData is provided, and at least one genotyped individual has an unknown birth year or unknown sex.


Any individual in Pedigree that does not occur in GenoM is substituted by a dummy individual; these can be recognised by the value 0' in columns '' and `` in the output. For non-genotyped individuals the parental log-likelihood ratio can be calculated if they have at least one genotyped offspring (see also getAssignCat).

The birth years in LifeHistData and the AgePrior are not used in the calculation and do not affect the value of the likelihoods for the various relationships, but they _are_ used during some filtering steps, and may therefore affect the likelihood _ratio_. The default (AgePrior=FALSE) assumes all age-relationship combinations are possible, which may mean that some additional alternatives are considered compared to the sequoia default, resulting in somewhat lower LLR values.

A negative LLR for A's parent B indicates either that B is not truely the parent of A, or that B's parents are incorrect. The latter may cause B's presumed true, unobserved genotype to divert from its observed genotype, with downstream consequences for its offspring. In rare cases it may also be due to 'weird', non-implemented double or triple relationships between A and B.

See also

SummarySeq for visualisation of OH & LLR distributions; CalcPairLL for the likelihoods underlying the LLR, GenoConvert to read in various genotype data formats, CheckGeno; PedPolish to check and 'polish' the pedigree; getAssignCat to find which id-parent pairs are both genotyped or can be substituted by dummy individuals; sequoia for pedigree reconstruction.


# count Mendelian errors in an existing pedigree
Ped.OH <- CalcOHLLR(Pedigree = Ped_HSg5, GenoM = SimGeno_example,
                    CalcLLR = FALSE)
#> Counting Mendelian errors ...
#>        id    dam   sire OHdam OHsire MEpair
#> 50 a01010 a00008 b00016     0      0      1     200         199          198
#> 51 b01011 a00008 b00016     0      0      1     200         199          198
#> 52 b01012 a00008 b00016     0      0      1     198         197          196
#> 53 b01013 a00011 b00001     0      0      0     199         199          199
#> 54 b01014 a00011 b00001     0      0      0     198         198          198
#> 55 b01015 a00011 b00001     0      0      1     198         198          198
# view histograms
SummarySeq(Ped.OH, Panels="OH")

# Parent likelihood ratios in an existing pedigree, including for
# non-genotyped parents
Ped.LLR <- CalcOHLLR(Pedigree = Ped_HSg5, GenoM = SimGeno_example,
                    CalcLLR = TRUE, LifeHistData=LH_HSg5, AgePrior=TRUE)
#> Ageprior: Flat 0/1, discrete generations, MaxAgeParent = 1,1
#> Counting Mendelian errors ...
#> Counting opposing homozygous loci between all individuals ...
#> Calculating parental LLR ...
SummarySeq(Ped.LLR, Panels="LLR")

if (FALSE) {
# likelihood ratios change with presumed genotyping error rate:
Ped.LLR.B <- CalcOHLLR(Pedigree = Ped_HSg5, GenoM = SimGeno_example,
                    CalcLLR = TRUE, LifeHistData=LH_HSg5, AgePrior=TRUE,
                    Err = 0.005)
SummarySeq(Ped.LLR.B, Panels="LLR")

# run sequoia with CalcLLR=FALSE, and add OH + LLR later:
SeqOUT <- sequoia(Geno_griffin, LH_griffin, CalcLLR=FALSE,quiet=TRUE,Plot=FALSE)
PedA <- CalcOHLLR(Pedigree = SeqOUT[["Pedigree"]][, 1:3], GenoM = Genotypes,
  LifeHistData = LH_griffin, AgePrior = TRUE, Complex = "full")
SummarySeq(PedA, Panels=c("LLR", "OH"))