Special featuresof RAD Sequencing data: 
implicationsfor genotyping
John W. Davey, Timothée Cezard, PabloFuentesUtrilla, Cathlene Eland, KarimGharbi, 
Mark L. Blaxter
Supporting Information
Additional Methods
Plasmid digestion and shearing
To evaluate the effect of fragmentlength onshearing efficiency (Figure S2) two commercial cloning vectors
of different sequence lengths (pOJ260, 3,469 bp, GenBank GU270843.1, and pcDNA3 RL-GW, 7,981 bp, 
www.dkfz.de/gpcf/749.html), were subjected tothe equivalent shearing conditions usedin RAD-Seq library 
preparation. Cloning vectorswere linearized by digesting 2.5!gof eachvector with 20U ofEcoRI-HF (New 
EnglandBioLabs) in a 50!Lreaction using two replicatesper vector. Digestions werediluted with water toa 
volume of135!L, with130!L transferred to a Covaris microTUBE withAFA fiber forshearing in a Covaris
S2. Shearing conditions used followed the manufacturer’s recommended values for 500 bp target peak 
(www.covarisinc.com, protocolpn_40056). Pre-and post-shearing sample DNAfragmentswere analyzed by 
microfluidic capillary electrophoresis (High Sensitivity DNA Chip Assay in an Agilent 2100 Bioanalyzer, 
Agilent Technologies).
C. elegans libraries for shearing test
TwoadditionalPstI RAD libraries were prepared toinvestigatethe impact ofshearingon readdepth (Figure 
S3). Short RAD adapterswereusedfor these libraries asper Andersen et al. (2012), modified forpaired end
sequencing  and indexing on the Illumina platform (Table S3). Both libraries were prepared with 4.8 !g 
gDNA and enrichedwithdifferent indexing P2 PCR primers (threeper library). The libraries weresubjected
to two different shearing conditions in a Covaris S2 sonicator using TC13 tubes (max. 1.5 mL sample). In 
library “Single”, allDNA was shearedto 10 shearing cycles, one shearing cycle consisting of a shearing step (duty cycle 20%, intensity 5, 200 cycles per burst, cycling time 30s)and a pause step to allow cooling of the 
sample (duty cycle 3%, intensity 1; 50 cycles per burst, cycle time 20s). In library “Mixed”, one half of the 
sample DNA wassheared to 10 cycles and the other half to 20 cycles. After 14 cyclesof PCRenrichment, 
librarieswerenormalizedto15nM,  pooled togetherequimolarly and sequencedinone Illumina MiSeq run 
(150 bp pairedendreads).
Sequence analysis for additionalpublic RADlibraries
Two additional public data sets are shown in Figure S1: EcoRI C. elegans RAD data (Andersen et al. 2012, 
Sequence Read Archive  accession 047839, library ECA17, kindly made available by E. Andersen and J. 
Shapirobefore SRA release) and SbfI stickleback RAD data (Hohenlohe et al. 2010, Sequence Read Archive 
run SRR034312, aligned to G. aculeatusgenome assembly BROAD S1 in Ensembl release66). These datasets
were analysed asfor Figure 3, described in themainMethods section, except that asonly read 1 data was
available for the EcoRI and SbfI samples, our C. elegans PstI data was reanalysed using only read 1 data. 
Libraries were subsampled forcomparisonto 6,516,557 reads, the numberof uniquely mapping readsin the 
selectedstickleback library.
Dataanalysis and graphics tools
Loess regression curvesand all plots generated withggplotversion 0.9.0 (Wickham 2009), except Figure 1, 
produced using IGV v2.1.16 (Thorvaldsdóttir etal.  2012), and the lowerpanelsofFigure6, produced using 
the R gridpackage (Murrell 2005). Figuresand tables were prepared forpublication usingAdobe Illustrator 
v5.1 andApple iWork ’09.Additional Discussion
Test datafor RADanalyses
To test the biases described here, it wasnecessary to developa set of heterozygousRAD loci withknown 
restriction fragmentlengths. Wehave used a RAD library from a Heliconius mapping cross for this purpose. 
This data was the only publicly available data known to us that featured a reference genome (allowing 
measurement of restriction fragment lengths), was highly heterozygous (allowing tests of many special 
features of RAD data as shown in Figure 5), had known, simple genetic structure (making phasing of 
haplotypespossible, essential for testing alleles withvarying restriction fragment lengths) and had paired
endreadsavailable for assessment of RADcontig assembly.
However, this data set has several drawbacks. Firstly,  the H. melpomene reference genome is new and, as
with all recent genome sequences,  has many errors. We have attempted to avoid many of these by only 
including restriction fragments containedwithin a singlecontig, and discarding those within a scaffold but 
spanning multiple contigs. This haslimited theavailable fragment lengths to some extent, with few RAD
loci having fragmentlengths over 10 kb. Secondly, wehaveno externaldataset(forexample, whole genome 
sequencing)to compare the RAD genotypes to, andso thevalidated genotypes areentirely derived fromthe 
RAD data itself. We have confidence in our identification of recombinants in this family, and so in our 
genotypes for eachindividual, making itpossible to have someconfidence in genotypes across lociand in 
our inference of genotypes for missing data. But many loci were excluded in this process; the data set is
intendedtobe free of repetitive loci andloci with questionable genotypeswere excluded.
Thismeans that, while thisdata set was suitable forexploring the effects of restriction fragmentlength and
restriction site heterozygosity, it underestimates the complexity of real RAD data sets, and so was not 
suitable for a fullassessmentof thetools discussed here. Itwould be highly desirable to develop a data setof 
heterozygous locifora species with a reference genomewheregenotypesare already known from other data 
sources. Thisdata set couldbeusedto more thoroughly validateexisting tools andsearchfor other biasesin 
RADdata that may not have been detectedhere. Legendsfor Supporting FiguresandTables
Figure S1 Effect of restriction fragment length in three RAD-Seq data sets. Left panels, relationship of 
restriction fragment length to read depth as per Figure 3. Right panels, histograms showing frequency of 
restriction fragmentlengths for each restrictionenzymein the relevant reference genome, indepedent of the 
RADlibraries. All three librariesshow a tendency for read depth toincrease asrestrictionfragmentlength 
increases, untilfragmentlengths reach 10 kb, a boundary clearlyvisible inA andCbut not reachedin B due 
to lack of long fragmentsproducedby EcoRI. Density ofpointsreflects number of restriction cutsites (PstI, 
13,552;  EcoRI, 45,939; SbfI, 19,354). Heterozygosity of sample has a large influence onread depth profile; the 
wildsamplesshown in C aremore comparable to theHeliconius datashown in Figure 6, top panel, than the 
clonal lab strain shown in A.
Figure S2 Shearing ofplasmids pOJ260(3.5kb, red) and pLUM(9 kb, blue). Toppanel, unsheared; bottom
panel, sheared. Neither of the plasmids is completely sheared, but shearing of the 3.5 kb plasmid is less
complete than the 9 kb plasmid.
Figure S3 Read depth varies with modified shearing conditions. Two C. elegans PstI RAD libraries were 
sequenced. Thefirst was sheared aspertheother C. elegans libraries, using 10 shearing cycles (Single). The 
second was a pool of two libraries, one sheared using 10 cycles, another sheared using 20 cycles (Mixed). 
Additionalshearing increasesreaddepth for RADloci fromshortrestrictionfragments and proportionally 
reducesdepth for RAD loci fromlongerrestriction fragments. Further experimentation with varied shearing 
of additional aliquotsmay permit balancing of readdepth acrossthe range of fragment lengths. 
Figure S4Heliconius allele countscannot be separatedby removing PCRduplicates. The same lociasshown 
in Figure 6, toppanel, are shownwith Sheared Fragment Depth rather than Read Depth. While varianceis
reduced, there isstill an overlap between depth of single copy allelesandtwocopy alleles.
Figure S5 Additional RAD contig assembly comparisons. A, assembly by allele rather than locus avoids
difficulties with heterozygosity. B and C, coverage for all contigs (B) and longest contig (C) for four 
additional assemblers. VelvetKandVelvetOptimisertogetherapproach the quality of VelvetOptimiser alone 
but some failedassembliesremain (redbar on left).Table S1 C. elegans PstI RAD library details. Four replicates for each test of 14, 16, 18, 20, 22 and 24 PCR 
cycles were pooled together using 24 different barcodes. Fragments are unique read1+read2 sequence 
counts, approximating the number of DNA fragments in initial samples after removal of PCR duplicates. 
Proportionscalculatedasdescribedin Materialsandmethods.
Table S2 RAD contig assembly details. URLs are given where no publication exists for an assembly tool. 
While optimized assemblers perform considerably better than unoptimized assemblers, the performance 
cost issubstantial.
Table S3 RAD-Seq adapter and primer sequences. Adapter oligos were ordered from Integrated DNA
Technologies (Ultramer synthesis, TruGrade purification). PCR primers were ordered from Sigma-Aldrich 
(PAGEpurified).Figure S1
A C. elegans N2 lab strain cut with PstI, shearedwith Covarissonication
B C. elegans N2-like wildstrainscut with EcoRI, shearedwith nebulization (Andersen et al. 2012)
C G. aculeatus wildpopulation cut with SbfI, shearedwith Bioruptor sonication (Hohenlohe et al. 2010)
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100 1,000 10,000 50,000
Restriction Fragment Length
Read Depth
Shearing
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Mixed
SingleFigure S4
Mother (AC) Father (BC) AB AC CB CC
0
20
40
60
80
100
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0.4 1 4 10 40 0.4 1 4 10 40 0.4 1 4 10 40 0.4 1 4 10 40 0.4 1 4 10 40 0.4 1 4 10 40
Restriction Fragment Length (kbp)
Shear
ed Fragment Depth
Allele
Copies
●
●
1
2Figure S5
Consensus LOCAS VelvetK + Optimiser SOAPdenovo CLC bio
0
50
100
150
200
250
0 100 200 300 400 0 100 200 300 400 0 100 200 300 400 0 100 200 300 400 0 100 200 300 400
All Contigs Reference Coverage
Fr
equency
Number of
Contigs
0
1
2
3
>3
Consensus LOCAS VelvetK + Optimiser SOAPdenovo CLC bio
0
50
100
150
200
250
0 100 200 300 400 0 100 200 300 400 0 100 200 300 400 0 100 200 300 400 0 100 200 300 400
Longest Contigs Reference Coverage
Fr
equency
Number of
Contigs
0
1
2
3
>3
Consensus Velvet LOCASopt VelvetOptimiser IDBA−UD VelvetK
0
100
200
300
400
0 100 200 300 400 0 100 200 300 400 0 100 200 300 400 0 100 200 300 400 0 100 200 300 400 0 100 200 300 400
All Contigs Reference Coverage
Fr
equency
Number of
Contigs
0
1
2
3
>3
A
B
CTable S1
PCR cycles Replicate Barcode Reads Perfectly Aligned 
Reads
Read 
Proportion
Perfectly Aligned 
Fragments
Fragment 
Proportion
14
1
2
3
4
16
1
2
3
4
18
1
2
3
4
20
1
2
3
4
22
1
2
3
4
24
1
2
3
4
ACCAT 12,326,499 8,379,012 9.59 4,304,560 8.29
CTAGG 3,948,749 2,599,743 2.98 1,845,053 3.55
GGTTC 3,858,053 2,584,773 2.96 1,853,492 3.57
TAGCA 7,245,703 4,820,991 5.52 2,963,020 5.70
AGTCA 3,074,799 2,049,193 2.35 1,380,758 2.66
CGCGC 6,425,852 4,289,467 4.91 2,536,337 4.88
GATCG 4,402,052 2,911,895 3.33 1,852,804 3.57
TTAAT 3,322,388 2,199,939 2.52 1,443,247 2.78
ACTGC 11,077,912 7,058,494 8.08 3,594,563 6.92
CGTAT 1,500,152 936,050 1.07 757,827 1.46
GTACA 3,840,509 2,394,621 2.74 1,669,857 3.21
TGTGG 2,296,297 1,434,825 1.64 1,101,822 2.12
ATCGA 2,030,189 1,292,540 1.48 1,093,084 2.10
CAGTC 4,576,312 2,880,937 3.30 2,141,614 4.12
GCATT 4,161,704 2,623,745 3.00 1,984,393 3.82
TCGAG 4,659,716 2,946,855 3.37 2,181,666 4.20
ATATC 7,350,846 4,693,005 5.37 2,731,100 5.26
CACAG 2,705,832 1,704,638 1.95 1,196,171 2.30
GGCCT 3,640,740 2,307,852 2.64 1,546,958 2.98
TCAGA 7,603,924 4,849,554 5.55 2,778,670 5.35
AGAGT 11,324,477 7,565,595 8.66 3,696,726 7.12
CCAAC 16,077,734 10,842,389 12.41 4,640,207 8.93
GACTA 2,367,316 1,545,522 1.77 1,058,500 2.04
TTCCG 3,711,388 2,453,537 2.81 1,589,064 3.06Table S2
Assembler Version Reference Parameters Mean time per assembly (s)
By locus By allele
CLCbio
IDBA-UD
LOCAS
LOCASOpt
SOAPdenovo
Velvet
VelvetK
VelvetOptimiser
VelvetK + 
VelvetOptimiser
4.06beta.67189 http://www.clcbio.com Default 0.46 0.25
1.0.9 Peng et al.2012 Default 1.49 0.63
0.1.7 Klein et al.2011 Default 2.85 0.48
0.5 Willing et al.2011 kmer range 11-15
required overlap length 21-35 bases,
error rate 0.02-0.04
87.20 18.75
1.05 Li, Zhu et al.2010 Default 0.26 0.13
1.2.05 Zerbino et al.2008 kmer 29 0.74 0.24
6 June 2012 http://www.vicbioinformatics.com/software.velvetk.shtml use --best option
then run Velvet with kmer chosen by VelvetK
0.91 0.63
2.2.0 http://www.vicbioinformatics.com/software.velvetoptimiser.shtml kmer range 19-99
optimise for longest contig
170.58 170.62
As above As above VelvetOptimiser kmer range +/-10 from VelvetK estimate 46.57 46.40
Table S3
Name Sequence Notes
Short P1 Adapter Top
Short P1 Adapter Bottom
Short P2 Adapter Top
Short P2 Adapter Bottom
Forward amplification primer
Reverse amplification primer
ACACTCTTTCCCTACACGACGCTCTTCCGATCTXXXXXXXXTGC*A XXXXXXXX= 8bp barcode sequence;
* = phosphothiorate bond
/5Phos/YYYYYYYYAGATCGGAAGAGCGTCGTGTAGGGAAAGAGTGT YYYYYYYY= 8bp barcode sequence, reverse
complement; 5Phos = added phosphate
/5Phos/GATCGGAAGAGCGGTTCAGCAGGTCCATTC 5Phos = added phosphate
GAGATCGGTCTCGGCATTCCTGCTGAACCGCTCTTCCGATCT*T * = phosphothiorate bond
AATGATACGGCGACCACCGAGATCTACACTCTTTCCCTACACGACGCTC*T * = phosphothiorate bond
Sequence provided by Michael A Quail (mq1@sanger.ac.uk; Wellcome Trust Sanger Institute, Hinxton, Cambridge, CB10 1SA, UK), available on request