327. Bolívar, P., Mugal, C.F., Rossi, M., Nater, A., Wang, M., Dutoit, L. & Ellegren, H. (2018) Biased inference of selection due to GC-biased gene conversion and the rate of protein evolution in flycatchers when accounting for it. Molecular Biology and Evolution, in press.

326. Yazdi, H.P. & Ellegren, H. (2018) A genetic map of the ostrich Z chromosome and the role of inversions in avian sex chromosome evolution. Genome Biology and Evolution, in press.

325. Dutoit, L., Mugal, C.F., Bolivar, P., Wang, M., Nadachowska-Brzyska, K., Smeds, L., Papoli, H., Gustafsson, L. & Ellegren, H. (2018) Sex-biased gene expression, sexual antagonism and levels of genetic diversity in the collared flycatcher (Ficedula albicollis) genome. Molecular Ecology, in press.

324. Ekblom, R., Brechlin, B., Persson, J., Smeds, L., Johansson, M., Magnusson, J., Flagstad, Ø. & Ellegren, H. (2018). Genome sequencing and conservation genomics in the Scandinavian wolverine (Gulo gulo) population. Conservation Biology, in press.

323. Craig, R., Suh, A., Wang, M. & Ellegren, H. (2018) Natural selection beyond genes: Identification and analyses of evolutionarily conserved elements in the genome of the collared flycatcher (Ficedula albicollis). Molecular Ecology 27:476-492.

322. Suh, A., Smeds, L. & Ellegren, H. (2018) Abundant recent activity of retrovirus-like retrotransposons within and among flycatcher species implies a rich source of structural variation in songbird genomes. Molecular Ecology 27:99-111.

321. Kardos, M., Åkesson, M., Fountain, T., Flagstad, Ø., Liberg, O., Olason, P., Sand, H., Wabakken, P., Wikenros, C. & Ellegren, H. (2018) Genomic consequences of intensive inbreeding in an isolated wolf population. Nature Ecology & Evolution 2:124-131.


320. Kawakami, T., Mugal, C.M., Suh, A., Nater, A., Burri, R., Smeds, L. & Ellegren, H. (2017) Whole-genome patterns of linkage disequilibrium across flycatcher populations clarify the causes and consequences of fine-scale recombination rate variation in birds. Molecular Ecology 26: 4158-4172.

319. Ellegren, H. & Wolf, J.B (2017) Parallelism in genomic landscapes of differentiation, conserved genomic features and the role of linked selection. Journal of Evolutionary Biology 30: 1516-1518.

318. Vijay, N., Weissensteiner, M., Burri, R., Kawakami, T., Ellegren, H. & Wolf, J.B.W. (2017) Genomewide patterns of variation in genetic diversity are shared among populations, species and higher-order taxa. Molecular Ecology 26: 4284-4295.

317. Wang, M., Uebbing, S., & Ellegren, H. (2017) Bayesian inference of allele-specific gene expression indicates abundant cis-regulatory variation in natural flycatcher populations. Genome Biology and Evolution 9: 1266-1279.

316. Silva, C.N.S, McFarlane, S.E., Hagen, I.J., Rönnegård, L., Billing, A.M., Kvalnes, T., Kemppainen, P., Rønning, B., Ringsby, T.H., Sæther, B-E., Qvarnström, A., Ellegren, H., Jensen, H, & Husby, A. (2017) Insights into the genetic architecture of morphological traits in two passerine bird species. Heredity 119: 197-205.

315. Dutoit, L., Vijay, N., Mugal, C.F., Bossu, C.M., Burri, R., Wolf, J. & Ellegren, H. (2017) Co-variation in levels of nucleotide diversity in homologous regions of the avian genome long after completion of lineage sorting. Proceedings of the Royal Society of London Series B: Biological Sciences 284: 20162756.

314. Kardos, M., Qvarnström, A., & Ellegren, H. (2017) Inferring individual inbreeding and demographic history from segments of identity by descent in Ficedula flycatcher genome sequences. Genetics 205: 1319-1334.

313. Knief, U., Schielzeth, H., Backström, N., Hemmrich-Stanisak, G., Wittig, M., Franke, A., Griffith, S.C., Ellegren, H., Kempenaers, B. & Forstmeier, W. (2017) Association mapping of morphological traits in wild and captive zebra finches: reliable within but not between populations. Molecular Ecology 26: 1285-1305.

312. Dutoit, L. Burri, R., Nater, A., Mugal, C.F. & Ellegren, H. (2017) Genomic distribution and estimation of nucleotide diversity in natural populations: perspectives from the collared flycatcher (Ficedula albicollis) genome. Molecular Ecology Resources 17:586-597.

311. Wolf, J.B. & Ellegren, H. (2017) Making sense of genomic islands of differentiation in light of speciation. Nature Reviews Genetics  18:87-100.


310. Kardos, M., Taylor, H.R., Ellegren, H., Luikart, G. & Allendorf, F.W. (2016) Genomics advances the study of inbreeding depression in the wild. Evolutionary Applications 9:1205-1218.

309. Smeds, L., Qvarnström, A. & Ellegren, H. (2016) Direct estimate of the rate of germline mutation in a bird. Genome Research 26:1211-128.

308. Smeds, L., Mugal, C.F., Qvarnström, A. & Ellegren, H. (2016) High-resolution mapping of crossover and non-crossover recombination events by whole-genome re-sequencing of an avian pedigree. PLoS Genetics 12:e1006044.

307. Ellegren, H. & Galtier, N. (2016) Determinants of genetic diversity. Nature Reviews Genetics 17:422-433.

306. Uebbing, S., Künstner, A., Mäkinen, H., Backström, N., Bolivar, P., Burri, R., Dutoit, L., Mugal, C.F., Nater, A., Aken, B., Flicek, P., Martin, F.J., Searle, S.M.J. & Ellegren, H. (2016) Divergence in gene expression within and between two closely related flycatcher species. Molecular Ecology 25:2015-2028.

305. Figuet, E., Nabholz, B., Bonneau, M., Mas Carrio, E., Nadachowska-Brzyska, K., Ellegren, H. & Galiter, N. (2016) Life history traits, protein evolution and the nearly-neutral theory in amniotes. Molecular Biology and Evolution 33:1517-1527.

304. Nadachowska-Brzyska, K., Burri, R., Smeds, L. & Ellegren, H. (2016) PSMC-analysis of effective population sizes in molecular ecology and its application to black-and-white Ficedula flycatchers. Molecular Ecology 25:1058-1072.

303. Rönnegård, L., McFarlane, E.S., Husby, A., Kawakami, T., Ellegren, H. & Qvarnström, A. (2016) Increasing the power of genome wide association studies in natural populations using repeated measures - evaluation and implementation. Methods in Ecology and Evolution 7:792-799.

302. Bischof, R, Gregersen, E.R., Brøseth, H., Ellegren, H. & Flagstad, Ø. (2016) Non-invasive sampling reveals intra-sex territoriality in wolverines. Ecology and Evolution 6:1527-1536.

301. Kardos, M., Husby, A., McFarlane, S.E., Qvarnström, A. & Ellegren, H. (2016) Whole genome resequencing of extreme phenotypes in collared flycatchers highlights the difficulty of detecting quantitative trait loci in natural populations. Molecular Ecology Resources 16:216-227.

300. Bolivar, P., Mugal, C.F. Nater, A. & Ellegren, H. (2016) Recombination rate variation modulates gene sequence evolution mainly via GC-biased gene conversion, not Hill-Robertson interference, in an avian system. Molecular Biology and Evolution 33:216-227.


299. Mugal, C.F., Weber, C.C. & Ellegren, H. (2015) GC-biased gene conversion links the recombination landscape and demography to genomic base composition. BioEssays 37:1317-1326. 

298. Burri, R., Nater, A., Kawakami, T., Mugal, C.F., Olason, P.I., Smeds, L., Suh, A., Dutoit, L., Bureš, S., Garamszegi, L.Z., Hogner, S., Moreno, J., Qvarnström, A., Ružić, M., Sæther, S.A., Sætre, G.P., Török, J. & Ellegren, H. (2015) Linked selection and recombination rate variation drive the evolution of the genomic landscape of differentiation across the speciation continuum of Ficedula flycatchers. Genome Research 25:1656-1665.

297. Schmid, M., Smith, J., Burt, D.W., Aken, B.L., Antin, P.B., Archibald, A.L., Ashwell, C., Blackshear, P.J., Boschiero, C., Brown, C.T., Burgess, S.C., Cheng, H.H., Chow, W., Coble, D.J., Cooksey, A., Crooijmans, R.P.M.A., Damas, J., Davis, R.V.N., de Konig, D.-J., Delany, M.E., Derrien, T., Desta, T.T., Dunn, I.C., Dunn, M. Ellegren, H. et al. (79 additional co-authors). (2015) Third report on chicken genes and chromosomes. Cytogenetics and Genome Research 145:78-179.

296. Gervasi, V., Brøseth, H., Nilse, E.B., Ellegren, H., Flagstad, Ø. & Linnell, J.D.C. Compensatory immigration counteracts contrasting conservation strategies of wolverines (Gulo gulo) within Scandinavia. Biological Conservation 191:632-639.

295. Suh, A., Smeds, L. & Ellegren, H. (2015) The dynamics of incomplete lineage sorting across the ancient adaptive radiation of neoavian birds. PLoS Biology 13:e1002224.

294. Nater, A., Burri, R., Kawakami, T., Smeds, L. & Ellegren, H. (2015) Resolving evolutionary relationships in closely related species with whole-genome sequencing data. Systematic Biology 64:1000-1017.

293. Uebbing, S., Konzer, A., Xu, L., Backström, N., Brunström, B., Bergquist, J. & Ellegren, H. (2015) Quantitative mass spectrometry reveals partial translational regulation for dosage compensation in chicken. Molecular Biology and Evolution 32:2716-2725.

292. Knief, U., Schielzeth, H., Ellegren, H., Kempenaers, B. & Forstmeier, W. (2015) A prezygotic transmission distorter acting equally in female and male zebra finches Taeniopygia guttata. Molecular Ecology 24:3846-3859.

291. Smeds, L., Warmuth, V., Bolivar, P., Uebbing, S., Burri, R., Suh, A., Nater, A., Bureš, S., Garamszegi, L.Z., Hogner, S., Moreno, J., Qvarnström, A., Ružić, M., Sæther, S.A., Sætre, G.P., Török, J. & Ellegren H. (2015) Evolutionary analysis of the female-specific avian W chromosome. Nature Communications 6:7330.

290. Nadachowska-Brzyska, K., Li, C., Smeds, L., Zhang, G. & Ellegren, H. (2015) Temporal dynamics of avian populations during Pleistocene revealed by whole-genome sequences. Current Biology 25:1375-1380.

289. Husby, A., Kawakami, T., Rönnegård, L., Smeds, L., Ellegren, H. & Qvarnström, A. (2015) Genome-wide association mapping in a wild avian population identifies a link between genetic and phenotypic variation in a life-history trait. Proceedings of the Royal Society of London, Series B 282:20150156.

288. Mugal, C.D., Arndt, P.F, Holm, L. & Ellegren, H. (2015) Evolutionary consequences of DNA methylation on the GC content in vertebrate genomes. G3 5:441-447.

287. Jarvis, E.D., Mirarab, S., Aberer, A.J., Li, B., Houde, P., Li, C., Ho, S.Y., Faircloth, B.C., Nabholz, B., Howard, J.T., Suh, A., Weber, C.C., da Fonseca, R.R., Alfaro-Núñez, A., Narula, N., Liu, L., Burt, D., Ellegren, H., Edwards, S.V., Stamatakis, A., Mindell, D.P., Cracraft, J., Braun, E.L., Warnow, T., Jun, W., Gilbert, M.T. & Zhang G; Avian Phylogenomics Consortium. (2015) Phylogenomic analyses data of the avian phylogenomics project. Gigascience 4:4.


286. Jarvis, E.D. et al. (Avian Phylogenomics Consortiuum) (2014) Whole-genome analyses resolve early branches in the tree of life of modern birds. Science 346: 1320-1331.

285. Li, C. et al. (40 co-authors) (2014) Two Antarctic penguin genomes reveal insights into their evolutionary history and molecular changes related to the Antarctic environment. Gigascience 3:27.

284. Suh, A., Weber, C.C., Kehlmaier, C., Braun, E.L., Green, R.E., Fritz, U., Ray, D.A. & Ellegren, H. (2014) Crocodilian and turtle paleoviruses suggest Early Mesozoic coexistence of amniotes and Hepadnaviridae. PLoS Genetics, 10:e1004559.

283. Weber, C.C.,Boussau, B., Romiguier, J., Jarvis, E.D. & Ellegren, H. (2014) Evidence for GC-biased gene conversion as a driver of between-lineage differences in avian base composition. Genome Biology 15:549.

282. Weber, C.C., Nabholz, B., Romiguier, J. & Ellegren, H. (2014) Kr/Kc but not dN/dS correlates positively with body mass in birds, raising implications for inferring lineage-specific selection. Genome Biology 15:542.

281. Smeds, L., Kawakami, T., Burri, R., Bolivar, P., Husby, A., Qvarnström, A., Uebbing, S. & Ellegren, H. (2014) Genomic identification and characterization of the pseudoautosomal region in highly differentiated avian sex chromosomes. Nature Communications 5:5448.

280. Ekblom, R., Smeds, L. & Ellegren, H. (2014) Patterns of sequencing coverage bias revealed by ultra-deep sequencing of vertebrate mitochondria. BMC Genomics 15:467.

279. Kawakami, T., Smeds, L., Backström, N., Husby, A., Qvarnström, A., Mugal, C.F., Olason, P. & Ellegren, H. (2014) A high-density linkage map enables a second-generation collared flycatcher genome assembly and reveals the patterns of avian recombination rate variation and chromosomal evolution. Molecular Ecology 23:4035-4058.

278. Kawakami, T., Backström, N., Burri, R., Husby, A., Olason, P., Rice, A.M., Ålund, M., Qvarnström, A. & Ellegren, H. (2014) Estimation of linkage disequilibrium and interspecific gene flow in Ficedula flycatchers by a newly developed 50k SNP array. Molecular Ecology Resources 14:1248-1260.

277. Yazdi, H.P. & Ellegren, H. (2014) Old but not (so) degenerated: slow evolution of largely homomorphic sex chromosomes in ratites. Molecular Biology and Evolution 31:1444-1453.

276. Ellegren, H. (2014) Genome sequencing and population genomics in non-model organisms. Trends in Ecology and Evolution 29:51-63.


275. Nadachowska-Brzyska, K., Burri, R., Olason, P.I., Kawakami, T., Smeds, L. & Ellegren, H. (2013) Demographic divergence history of pied flycatcher and collared flycatcher inferred from whole-genome re-sequencing data. PLoS Genetics 9:e1003942

274. Uebbing, S., Künstner, A., Mäkinen, H. & Ellegren, H. (2013) Transcriptome sequencing reveals the character of incomplete dosage compensation across multiple tissues in flycatchers. Genome Biology and Evolution 5:1555-1566.

273. Ellegren, H. (2013) The evolutionary genomics of birds. Annual Reviews of Ecology, Evolution and Systematics 44:239-259.

272. Poelstra, J.W., Ellegren, H. & Wolf, J.B. (2013) An extensive candidate gene approach to speciation: diversity, divergence and linkage disequilibrium in candidate pigmentation genes across the European crow hybrid zone. Heredity 111:467-473.

271. Mugal, C.F., Arndt, P.F. & Ellegren, H. (2013) Twisted signatures of GC-biased gene conversion embedded in an evolutionary stable karyotype. Molecular Biology and Evolution 30:1700-1712.

270. Corl, A. & Ellegren, H. (2013) Sampling strategies for species trees: The effects on phylogenetic inference of the number of genes, number of individuals, and whether loci are mitochondrial, sex-linked, or autosomal. Molecular Phylogenetics and Evolution 67: 358-366.

269. Mugal, C.F., Nabholz, B. & Ellegren, H. (2013) Genome-wide analysis in chicken reveals that local levels of genetic diversity are mainly governed by the rate of recombination. BMC Genomics 14:86.

268. Adolfsson, S. & Ellegren, H. (2013) Lack of dosage compensation accompanies the arrested stage of sex chromosome evolution in ostriches. Molecular Biology and Evolution 30:806-810.

267. Backström, N., Saetre, G.P. & Ellegren, H. (2013) Inferring the demographic history of European Ficedula flycatcher populations. BMC Evolutionary Biology 13:2.

266. Parsch, J. & Ellegren, H. (2013) The evolutionary consequences of sex-biased gene expression. Nature Reviews Genetics 14:83-87.

265. Nabholz, B., Ellegren, H. & Wolf, J.B.W (2013) High levels of gene expression explain the strong evolutionary constraint of mitochondrial protein-coding genes. Molecular Biology and Evolution 30:272-284.


264. Ellegren, H., Smeds, L., Burri, R., Olason, P.I., Backström, N., Kawakami, T., Künstner, A., Mäkinen, H., Nadachowska-Brzysja, K., Qvarnström, A., Uebbing, S. & Wolf, J.B.W. (2012) The genomic landscape of species divergence in Ficedula flycatchers. Nature 491:756-760.

263. Knief, U., Schielzeth, H., Kempenaers, B., Ellegren, H. & Forstmeier, W. (2012) QTL and quantitative genetic analysis of beak morphology reveals patterns of standing genetic variation in the zebra finch. Molecular Ecology 21:3704-3717.

262. Nam, K. & Ellegren, H. (2012). Recombination drives vertebrate genome contraction. PLoS Genetics 8: e1002680.

261. Forstmeier, W., Schielzeth, H., Mueller, J.C., Ellegren, H. & Kempenaers, B. (2012) Heterozygosity-fitness correlations in zebra finches: microsatellite markers can be better than their reputation. Molecular Ecology 21: 3237-3249.

260. Corl, A. & Ellegren, H. (2012) The genomic signature of sexual selection in the genetic diversity of the sex chromosomes and autosomes. Evolution 66: 2138-2149.

259. Schielzeth, H., Forstmeier, W., Kempenaers, B. & Ellegren, H. (2012) QTL linkage mapping of wing length in zebra finch using genome-wide SNP markers. Molecular Ecology 21: 329-339.

258. Schielzeth, H., Kempenaers, B., Ellegren, H. & Forstmeier, W. (2012) QTL linkage mapping of zebra finch beak colour shows an oligogenic control of a sexually selected trait. Evolution 66: 18-30.


257. Künstner, A., Nabholz, B. & Ellegren, H. (2011) Significant selective constraint at 4-fold degenerate sites in the avian genome and its consequence for detection of positively selected genes. Genome Biology and Evolution 3: 1381-1389.

256. Ellegren, H. (2011) Emergence of male-biased genes on the chicken Z-chromosome: Contrasts between male and female heterogametic systems. Genome Research 21:2082-2086.

255. Mugal, C.F. & Ellegren, H. (2011) Substitution rate variation at human CpG sites correlates with non-CpG divergence, methylation level and GC content. Genome Biology 12:R58.

254. Künstner, A., Nabholz, B. & Ellegren, H. (2011). Evolutionary constraint in flanking regions of avian genes. Molecular Biology and Evolution 28:2481-2489.

253. Nabholz, B., Künstner, A., Wang, R., Jarvis, E.D. & Ellegren, H. (2011) Dynamic evolution of base composition: causes and consequences in avian phylogenomics. Molecular Biology and Evolution 28:2197-2210.

252. Ellegren, H. (2011) Sex chromosome evolution: recent progress and the influence of male and female heterogamety. Nature Reviews Genetics 12:157-166.

251. Rice, A.M., Rudh, A., Ellegren, H. & Qvarnström, A. (2011) A guide to the genomics of ecological speciation in natural animal populations. Ecological Letters 14:9-18.

250. Ellegren, H. (2011) The singing genome. Heredity 106:533-534.


249. Nabholz, B., Jarvis, E.D. & Ellegren, H. (2010) Obtaining mtDNA genomes from next-generation transcriptome sequencing: A case study of the basal Passerida (Aves: Passeriformes) phylogeny. Molecular Phylogenetics and Evolution 57: 466-470.

248. Backström, N., Lindell, J., Zhang, Y., Palkopoulou, E., Qvarnström, A., Saetre, G.P. & Ellegren, H. (2010) A high-density scan of the Z-chromosome in Ficedula flycatchers reveals candidate loci for diversifying selection. Evolution 64: 3461-3475.

247. Backström, N., Palkopoulou, E., Qvarnström, A. & Ellegren, H. (2010) No evidence for Z-chromosome rearrangements between the pied flycatcher and the collared flycatcher as judged by gene-based comparative genetic maps. Molecular Ecology 19: 3394-3405.

246. Axelsson, E., Albrechtsen, A., As P. van, Li, L., Megens, H.J., Vereijken, A.L.J., Crooijmans, R.P.M.A., Groenen, M.A.M., Ellegren, H., Willerslev, E. & Nielsen, R. (2010) Segregation distortion in chicken and the evolutionary consequences of female meiotic drive in birds. Heredity 105: 290-298.

245. Forstmeier, W. & Ellegren, H. (2010) Trisomy and triploidy are sources of embryo mortality in the zebra finch. Proceedings of the Royal Society of London, Series B 277: 2655-2660.

244. Nam, K., Mugal, C., Nabholz, B., Schielzeth, H., Wolf, J.B., Backström, N., Künstner, A., Balakrishnan, C.N., Heger, A., Ponting, C.P., Clayton, D.F. & Ellegren, H. (2010) Molecular evolution of genes in avian genomes. Genome Biology 11: R68.

243. Mank, J.E., Nam, K., Brunström, B. & Ellegren, H. (2010) Ontogenetic complexity of sexual dimorphism and sex-specific selection. Molecular Biology and Evolution 27: 1570-1578.

242. Qvarnström, A., Rice, A.M. & Ellegren, H. (2010) Speciation in Ficedula flycatchers. Philosophical Transactions of the Royal Society of London B: Biological Sciences 365: 1841-1852.

241. Vali, U., Dombrovski, V., Treinys, R., Bergmanis, U., Daroczi, S.J., Dravecky, M., Ivanovski, V., Lontkowski, J., Maciorowski, G., Meyburg, B.U., Mizera, T., Zeitz, R. & Ellegren, H. (2010) Widespread hybridization between the Greater Spotted Eagle Aquila clanga and the Lesser Spotted Eagle Aquila pomarina (Aves: Accipitriformes) in Europe. Biological Journal of the Linnean Society 100: 725-736.

240. Lira, J., Linderholm, A., Olaria, C., Brandström Durling, M., Gilbert, T.P., Ellegren, H., Willerslev, E., Lidén, K., Arsuaga, J.L. & Götherström, A. (2010) Ancient DNA reveals traces of Iberian Neolithic and Bronze Age lineages in modern Iberian horses. Molecular Ecology 19: 64-78.

239. Völker, M., Backström, N., Skinner, B.M., Langley, E.J., Bunzey, S.K., Ellegren, H. & Griffin, D.K. (2010) Copy number variation, chromosome rearrangement, and their association with recombination during avian evolution. Genome Research 20: 503-511.

238. Warren, W.C., Clayton, D.F., Ellegren, H., Arnold, A.P., Hillier, L.W., Künstner, A., Searle, S., White, S., Vilella, A.J., Fairley, S., Heger, A., Kong, L., Ponting, C.P., Jarvis, E.D., Mello, C.V., Minx, P., Lovell, P., Velho, T.A., Ferris, M., Balakrishnan, C.N., Sinha, S., Blatti, C., London, S.E., Li, Y., Lin, Y.C., George, J., Sweedler, J., Southey, B., Gunaratne, P., Watson, M., Nam, K., Backström, N., Smeds, L., Nabholz, B., Itoh, Y., Whitney, O., Pfenning, A.R., Howard, J., Völker, M., Skinner, B.M., Griffin, D.K., Ye, L., McLaren, W.M., Flicek, P., Quesada, V., Velasco, G., Lopez-Otin, C., Puente, X.S., Olender, T., Lancet, D., Smit, A.F., Hubley, R., Konkel, M.K., Walker, J.A., Batzer, M.A., Gu, W., Pollock, D.D., Chen, L., Cheng, Z., Eichler, E.E., Stapley, J., Slate, J., Ekblom, R., Birkhead, T., Burke, T., Burt, D., Scharff, C., Adam, I., Richard, H., Sultan, M., Soldatov, A., Lehrach, H., Edwards, S.V., Yang, S.P., Li, X., Graves, T., Fulton, L., Nelson, J., Chinwalla, A., Hou, S., Mardis, E.R. & Wilson, R.K. (2010) The genome of a songbird. Nature 464: 757-762.

237. Mugal, C.F., Wolf, J.B.W., von Grünberg H.H. & Ellegren, H. (2010) Conservation of neutral substitution rate and substitutional asymmetries in mammalian genes. Genome Biology and Evolution 2010: 19-28.

236. Künstner, A., Wolf, J.B.W., Backström, N., Whitney, O., Balakrishnan, C., Day, L., Edwards, S.V., Janes, D.E., Schlinger, B.A., Wilson, R.K., Jarvis, E.D., Warren, W.C. & Ellegren, H. (2010) Comparative genomics based on massive parallel transcriptome sequencing reveals patterns of substitution and selection across 10 bird species. Molecular Ecology 19 Suppl 1: 266-276.

235. Backström, N., Forstmeier, W., Schielzeth, H., Mellenius, H., Nam, K., Bolund, E., Webster, M.T., Öst, T., Schneider, M., Kempenaers, B. & Ellegren, H. (2010) The recombination landscape of the zebra finch Taeniopygia guttata genome. Genome Research 20: 485-495.

234. Ellegren, H. (2010) Evolutionary stasis – the stable chromosomes of birds. Trends in Ecology and Evolution 25: 283-291.

233. Mank, J.E., Nam, K. & Ellegren, H. (2010) Faster-Z evolution is predominantly due to genetic drift. Molecular Biology and Evolution 27: 661-670.

232. Brøseth, H., Flagstad, Ø., Wärdig, C., Johansson, M. & Ellegren, H. (2010) Large-scale noninvasive genetic monitoring of wolverines using scats reveals density dependent adult survival. Biological Conservation 143: 113-120.

231. Hansson, B., Ljungqvist, M., Dawson, D.A., Mueller, J.C., Olano-Marin, J., Ellegren, H. & Nilsson, J.Å. (2010) Avian genome evolution: insights from a linkage map of the blue tit (Cyanistes caeruleus). Heredity 104: 67-78.


230. Ellegren, H. (2009) Sex determination: two copies for one cock. Current Biology 19: R909-910.

229. Wolf, J.B.W., Künstner, A., Nam, K., Jakobsson, M. & Ellegren, H. (2009) Non-linear dynamics of non-synonymous (dN) and synonymous (dS) substitution rates affects inference of selection. Genome Biology and Evolution 1: 308-319.

228. Mank, J.E. & Ellegren, H. (2009) Sex-bias in gene expression is not the same as dosage compensation. Heredity 103: 434.

227. Ellegren, H. (2009) Is genetic diversity really higher in large populations? Journal of Biology 8: 41.

226. Ellegren, H. (2009) The different levels of genetic diversity in sex chromosomes and autosomes. Trends in Genetics 25: 278-284.

225. Hagenblad, J., Olsson, M., Parker, H.G., Ostrander, E.A. & Ellegren, H. (2009) Population genomics of the inbred Scandinavian wolf. Molecular Ecology 18: 1341-1351.

224. Mank, J.E. & Ellegren, H. (2009) Are sex-biased genes more dispensable? Biology Letters 23: 409-412.

223. Mank, J.E. & Ellegren, H. (2009) Sex-linkage of sexually antagonistic genes is predicted by female, but not male, effects in birds. Evolution 63: 1464-1472.

222. Axelsson, E. & Ellegren, H. (2009) Quantification of adaptive evolution of genes expressed in avian brain and the population size effect on the efficacy of selection. Molecular Biology and Evolution 26: 1073-1079.

221. Ellegren, H. (2009) A selection model of molecular evolution incorporating the effective population size. Evolution 63: 301-305.

220. Mank, J.E. & Ellegren, H. (2009) All dosage compensation is local: Gene-by-gene regulation of sex-biased gene expression on the chicken Z chromosome. Heredity 102: 312-320.

219. Ellegren, H. (2009) Genomic evidence for a large-Z effect. Proceedings of the Royal Society of London, Series B 276: 361-366.


218. Ellegren, H. (2008) Comparative genomics and the study of evolution by natural selection. Molecular Ecology 17: 4586-4596.

217. Berlin, S., Qu, L. & Ellegren, H. (2008) Adaptive evolution of gamete-recognition proteins in birds. Journal of Molecular Evolution 67: 488-496.

216. Brandström, M., Bradshaw, A.T., Gemmell, N.J. & Ellegren, H. (2008) The relationship between microsatellite polymorphism and recombination hotspots in the human genome. Molecular Biology and Evolution 25: 2579-2587.

215. Nam, K. & Ellegren, H. (2008) The chicken (Gallus gallus) Z chromosome contains at least three non-linear evolutionary strata. Genetics 180: 1131-1136.

214. Berlin, S., Qu, L. Yang, N. & Ellegren, H. (2008) Positive diversifying selection in avian Mx genes. Immunogenetics 60: 689-697.

213. Väli, U., Einarsson, A., Waitz, L. & Ellegren, H. (2008) To what extent do microsatellite markers reflect genome-wide genetic diversity in natural populations? Molecular Ecology 17: 3808-3817.

212. Ellegren, H. (2008) Platypus genome suggests a recent origin for the human X. Current Biology 18: R557-559.

211. Backström, N., Karaiskou, N., Leder, E.H., Gustafsson, L. Primmer, C.R., Qvarnström, A. & Ellegren, H. (2008) A gene-based genetic linkage map of the collared flycatcher (Ficedula albicollis) reveals extensive synteny and gene order conservation during 100 million years of avian evolution. Genetics 179: 1479-1495.

210. Brandström, M. & Ellegren, H. (2008) Genome-wide analysis of microsatellite polymorphism in chicken circumventing the ascertainment bias. Genome Research 18: 881-887.

209. Mank, J.E., Hultin-Rosenberg, L., Webster, M.T. & Ellegren, H. (2008) The unique genomic properties of sex-biased genes: insights from avian microarray data. BMC Genomics 9: 148.

208. Axelsson., E., Hultin-Rosenberg, L., Brandström, M., Zwahlén, M., Clayton, D.F. & Ellegren, H. (2008) Natural selection in avian protein-coding genes expressed in brain. Molecular Ecology 17: 3008-3017.

207. Ellegren, H. & Sheldon, B.C. (2008) Genetic basis for fitness differences in natural populations. Nature 452: 169-175.

206. Ellegren, H. (2008) Sequencing goes 454 and takes large-scale genomics into the wild. Molecular Ecology 17: 1629-1631.

205. Väli, U., Brandström, M., Johansson, M. & Ellegren, H. (2008) Insertion-deletion polymorphisms (indels) as genetic markers in natural populations. BMC Genetics 9: 8.

204. Hefti-Gautschi, B., Pfunder, M., Jenni, L. Keller, V. & Ellegren, H. (2008) Identification of conservation units in the European Merganser merganser based on nuclear and mitochondrial DNA markers. Conservation Genetics 10: 87-99.

203. Sundqvist, A.K., Ellegren, H. & Vilà, C. (2008) Wolf or dog? Genetic identification of predator from saliva collected around bite wounds on prey. Conservation Genetics 9: 1275-1279.

202. Backström, N., Fagerberg, S. & Ellegren, H. (2008) Genomics of wild bird populations: a gene-based set of references markers spread across the avian genome. Molecular Ecology 17: 964-980.

201. Mank, J.E., Hultin-Rosenberg, L., Zwahlén, M. & Ellegren, H. (2008) Pleiotropic constraint hampers the resolution of sexual antagonism in vertebrate gene expression. American Naturalist 171: 35-43.


200. Ellegren, H. & Parsch, J. (2007) The evolution of sex-biased genes and sex-biased gene expression. Nature Reviews Genetics 8: 689-698.

199. Mank, J.E., Hultin-Rosenberg, L., Axelsson, E. & Ellegren, H. (2007) Rapid evolution of female-biased, but not male-biased, genes expressed in the avian brain. Molecular Biology and Evolution 24: 2698-2706.

198. Ellegren, H., Hultin-Rosenberg, L., Brunström, B., Dencker, L., Kultima, K. & Scholtz, B. (2007) Faced with inequality: chicken does not have a general dosage compensation of sex-linked genes. BMC Biology 5: 40.

197. Brandström, M. & Ellegren, H. (2007) The genomic landscape of short insertion and deletion polymorphisms in the chicken (Gallus gallus) genome: a high frequency of deletions in tandem duplicates. Genetics 176: 1691-1701.

196. Mank. J.M. & Ellegren, H. (2007) Parallel divergence and degradation of the avian W sex chromosome. Trends in Ecology and Evolution 22: 389-391.

195. Ellegren, H. (2007) A dinosaur’s view on genome size evolution. Current Biology 17; R470-472.

194. Engen, S., Ringsby, T.H., Lande, R., Jensen, H., Sæther, B.E., Lillegard, M. & Ellegren, H. (2007) Effective size of fluctuating populations with overlapping generations: extension of Wright’s formula for two sexes. Evolution 61: 1873-1885.

193. Mank, J.M., Axelsson, E. & Ellegren, H. (2007) Fast-X on Z: rapid evolution of sex-linked genes in birds. Genome Research 17: 618-624.

192. Ellegren, H. (2007) Avian molecular evolutionary genomics. Cytogenetics and Genome Research 117: 120-130.

191. Ellegren, H. (2007) Characteristics, causes and evolutionary consequences of male-biased mutation. Proceedings of the Royal Society London, Series B 274: 1-10.

190. Kaiser, V.B., van Tuinen, M. & Ellegren, H. (2007) The phylogenetic position of the chicken revisited: insertion events of CR1 retrotransposable elements elucidate the order of speciation events in galliform species (Aves). Molecular Biology and Evolution 24: 338-347.

189. Hedmark, E. & Ellegren, H. (2007) DNA-based monitoring of two newly founded Scandinavian wolverine populations. Conservation Genetics 8: 843-852.

188. Hedmark, E., Persson, J. & Ellegren, H. (2007) Paternity and mating system in wolverines. Wildlife Biology 13: 13-30.


187. Kaiser, V.B. & Ellegren, H. (2006) Non-random distribution of genes with sex-biased expression in the chicken genome. Evolution 60: 1945-1951.

186. Backström, N., Brandström, M., Gustafsson, L., Qvarnström, A., Cheng, H. & Ellegren, H. (2006) Genetic mapping in a natural population of collared flycatchers (Ficedula albicollis): Conserved synteny but gene order rearrangements on the avian Z chromosome. Genetics 174: 377-386.

185. Backström, N., Qvarnström, A., Gustafsson, L. & Ellegren, H. (2006) Levels of linkage disequilibrium (LD) in a wild bird population. Biology Letters 2: 435-438.

184. Hailer, F., Helander, B., Folkestad, A.O., Ganusevich, S.A., Garstad, S., Hauff, P., Koren, C., Nygård, T., Volke, V., Vilà, C. & Ellegren, H. (2006) Bottlenecked but long-lived: high genetic diversity retained in white-tailed eagles upon recovery from population decline. Biology Letters 2: 316-319.

183. Webster, M.T., Axelsson, E. & Ellegren, H. (2006) Strong regional biases in nucleotide substitution in the chicken genome. Molecular Biology and Evolution 23: 1203-1216.

182. Sundqvist, A.K., Björnerfeldt, S., Leonard, J.A., Hailer, F., Hedhammar, Å., Ellegren, H. & Vilà, C. (2006) Unequal contribution of sexes in the origin of dog breeds. Genetics 172: 2345-2350.

181. Berlin, S., Brandström, M., Backström, N., Axelsson, E., Smith, N.G.C. & Ellegren, H. (2006) Substitution rate heterogeneity and the male mutation bias. Journal of Molecular Evolution 62: 226-233.

180. Berlin, S. & Ellegren, H. (2006) Accumulation of deleterious mutations on the female-specific W chromosome in birds. Journal of Molecular Evolution 62: 66-72.

179. Seddon, J.M., Sundqvist, A.K. & Ellegren, H. (2006) Genetic identification of immigrants in the Scandinavian wolf population. Conservation Genetics 7: 225-230.

178. Hedmark, E. & Ellegren, H. (2006) A test of the multiplex pre-amplification approach in microsatellite genotyping of wolverine faecal DNA. Conservation Genetics 7: 289-293.


177. Ellegren, H. (2005) The dog has its day. Nature 438: 745-746.

176. Ellegren, H. (2005) Natural selection in the evolution of human and chimps. Current Biology 15: R919-922.

175. Götherström, A., Anderung, C., Hellborg, L. Elburg, R., Smith, C. & Ellegren, H. (2005) Cattle domestication in the Near East was followed by hybridization with aurochs bulls in Europe. Proceedings of the Royal Society London, Series B 272: 2345-2350.

174. Hedmark, E. & Ellegren, H. (2005) Successful genotyping of DNA isolated from claws left on tanned carnivore hides. International Journal of Legal Medicine 119: 370-373.

173. Backström, N., Ceplitis, H. & Ellegren, H. (2005) Gene conversion drives the evolution of HINTW, a multi-copy gene family on the avian W chromosome. Molecular Biology and Evolution 22: 1992-1999.

172. Anderung, C., Bouwman, A., Persson, P., Carretero, J. M., Ortega, A. I., Arasuaga, J. L. Elburg, R., Smith, C. Ellegren, H. & Götherström, A. (2005) Genetics of Iberian Bronze Age cattle remains indicate prehistoric contacts over the Straits of Gibraltar. Proceedings of the National Academy of Sciences USA 102: 8431-8435.

171. Webster, M.T., Smith, N.G.C., Hultin-Rosenberg, L., Arndt, P.F. & Ellegren, H. Male-driven biased gene conversion governs the evolution of base composition in human Alu repeats. Molecular Biology and Evolution 22: 1468-1474.

170. Berggren, K.T., Ellegren, H., Hewitt, G.M. & Seddon, J.M. (2005) Understanding the phylogeographic patterns of European hedgehogs, Erinaceus concolor and E. europaeus using the MHC. Heredity 95: 84-90.

169. Ellegren, H. (2005) The avian genome uncovered. Trends in Ecology and Evolution 20: 180-186.

168. Vilà, C., Seddon, J.M. & Ellegren, H. (2005) Genes of domestic mammals augmented by backcrossing with wild ancestors. Trends in Genetics 21: 214-218.

167. Seddon, J. M., Parker, H.G., Ostrander, E.A. & Ellegren, H. (2005) SNPs in ecological and conservation studies: a test in the Scandinavian wolf population. Molecular Ecology 14: 503-511.

166. Axelsson, E., Webster, M.T., Smith, N.G., Burt, D.W. & Ellegren, H. (2005) Comparison of the chicken and turkey genomes reveals a higher rate of nucleotide divergence on microchromosomes than macrochromosomes. Genome Research 15: 120-125.


165. International Chicken Polymorphism Map Consortium. (2004) A genetic variation map for chicken with 2.8 million single nucleotide polymorphisms. Nature 432: 717-722.

164. International Chicken Genome Sequencing Consortium. (2004) Sequencing and comparative analysis of the chicken genome. Nature 432: 695-716.

163. Berlin, S. & Ellegren, H. (2004) Chicken W: a genetically uniform chromosome in a highly variable genome. Proceedings of the National Academy of Sciences USA 101: 15967-15969.

162. Seddon, J.M. & Ellegren, H. (2004) A temporal analysis shows MHC loci in the Scandinavian wolf population are consistent with neutral evolution. Proceedings of the Royal Society London, Series B 271: 2283-2291.

161. Smith, N.G., Brandström, M. & Ellegren, H. (2004) Evidence for turnover of functional noncoding DNA in mammalian genome evolution. Genomics 84: 806-813.

160. Webster, M.T., Smith, N.G.C., Lercher, M.J. & Ellegren, H. (2004) Gene expression, synteny, and local similarity in human noncoding mutation rates. Molecular Biology and Evolution 21: 1820-1830.

159. Saetre, P., Lindberg, J., Leonard, J., Olsson, K., Pettersson, U., Ellegren, H., Bergström, T., Vila, C. & Jazin, E. (2004) From wild wolf to domestic dog: gene expression changes in the brain. Brain Research Molecular Brain Research 126: 198-206.

158. Ellegren, H. (2004) Microsatellites: simple sequences with complex evolution. Nature Reviews Genetics 5: 435-445.

157. Sundström, H., Webster, M. T. & Ellegren, H. (2004) Reduced variation on the chicken Z chromosome. Genetics 167: 377-385.

156. Lindgren, G., Backström, N., Swinburne, J., Hellborg, L., Einarsson, A., Sandberg, K., Vilà, C., Binns, M. & Ellegren, H. (2004) Limited number of patrilines in horse domestication. Nature Genetics 36: 335-336.

155. Axelsson, E., Smith, N. G. C., Sundström, H., Berlin, S. & Ellegren, H. (2004) Male-biased mutation rate and divergence in autosomal, Z-linked and W-linked introns of chicken and turkey. Molecular Biology and Evolution 21: 1538-1547.

154. Jensen, H., Saether, B.E., Ringsby, T.H., Tufto, J., Griffith, S.C. & Ellegren, H. Lifetime reproductive success in relation to morphology in house sparrow (Passer domesticus). Journal of Animal Ecology 73: 599-611.

153. Hedmark, E., Flagstad Ø., Segerström P., Persson, J., Landa, A. & Ellegren, H. (2004) DNA-based individual and sex identification from wolverine (Gulo gulo) faeces and urine. Conservation Genetics 5: 405-410.

152. Hellborg, L., & Ellegren, H. (2004) Low levels of nucleotide diversity in mammalian Y chromosomes. Molecular Biology and Evolution 21: 158-163.

151. Flagstad, Ø., Hedmark, E., Landa, A., Brøseth, H., Persson, J., Andersen, R., Segerström, P. & Ellegren, H. (2004) Colonization history and non-invasive monitoring of a re-established wolverine (Gulo gulo) population. Conservation Biology 18: 676-688.

150. Ceplitis, H. & Ellegren, H. (2004) Adaptive molecular evolution in HINTW, a female-specific gene in birds. Molecular Biology and Evolution 21: 249-254.

149. Brohede, J., Arnheim, N. & Ellegren, H. (2004) Single molecule analysis of the hypermutable tetranucleotide repeat locus D21S1245 through sperm genotyping: a heterogeneous pattern of mutation but no clear male age effect. Molecular Biology and Evolution 21: 58-64.

148. Berlin, S., Smith, N.G.C. & Ellegren, H. (2004) Do avian mitochondria recombine? Journal of Molecular Evolution 58: 163-167.

147. Lawson-Handley, L., Ceplitis, H. & Ellegren, H. (2004) Evolutionary strata on the chicken Z chromosome: Implications to sex chromosome evolution. Genetics 167: 367-376.


146. Jensen, H., Saether, B.E., Ringsby, T.H., Tufto, J., Griffith, S.C. & Ellegren, H. (2003) Sexual variation in heritability and genetic correlations of morphological traits in house sparrow (Passer domesticus). Journal of Evolutionary Biology 16: 1296-1307.

145. Ellegren, H., Smith, N.G.C. & Webster, M.T. (2003) Mutation rate variation in the mammalian genome. Current Opinion in Genetics and Development 13: 562-568.

144. Brohede, J., Møller, A.P. & Ellegren, H. (2003) Individual variation in microsatellite mutation rate in barn swallows. Mutation Research 545: 73-80.

143. Rueness, E. K., Jorde, P. E., Hellborg, L., Stenseth, N. C., Ellegren, H. & Jacobsen, K. S. (2003) Cryptic population structure in a large, mobile mammalian predator: the Scandinavian lynx. Molecular Ecology 12: 2623-2633.

142. Rueness, E. K., Stenseth, N. C., O’Donoghue, M., Boutin, S., Ellegren, H. & Jakobsen, K. S. (2003) The phylogeography of the Canadian lynx: coinciding genetic and ecological spatial structuring. Nature 425: 69-72.

141. Bartosch-Härlid, A., Berlin, S., Smith, N.G.C., Møller, A.P. & Ellegren, H. (2003) Life history and the male mutation bias. Evolution 57: 2398-2406.

140. Ellegren, H. & Fridolfsson, A.K. (2003) Sex-specific mutation rates in salmonid fish. Journal of Molecular Evolution 56: 458-463.

139. Sundström, H., Webster, M.T. & Ellegren, H. (2003) Is the rate of insertion and deletion mutation male-biased? - Molecular evolutionary analysis of avian and primate sex chromosome sequences. Genetics 164: 259-268.

138. Flagstad, Ø., Walker, C.W., Vilà, C., Sundqvist, A.K., Fernholm, B., Hufthammar, A.K., Wiig, Ø., Kojola, I. & Ellegren, H. (2003) Two centuries of the Scandinavian wolf population: patterns of genetic variability and migration during an era of publication decline. Molecular Ecology 12: 869-880.

137. Ellegren, H. (2003) Levels of polymorphism on the sex-limited chromosome – a clue to Y from W? BioEssays 25: 163-167.

136. Vilà, C., Walker, C., Sundqvist, A.K., Flagstad, Ø., Andersone, Z., Casulli, A., Kojola, I., Valdmann, H., Halverson, J. & Ellegren, H. (2003) Combined use of maternal, paternal and bi-parental genetic markers for the identification of wolf-dog hybrids. Heredity 90: 17-24.

135. Smith, N.G.C., Webster, M.T. & Ellegren, H. (2003) A low rate of simultaneous double nucleotide mutations in primates. Molecular Biology and Evolution 20: 47-53.

134. Webster, M.T., Smith, N.G.C. & Ellegren, H. (2003) Compositional evolution of non-coding DNA in the primate genome. Molecular Biology and Evolution 20: 278-286.

133. Vila, C., Sundqvist, A.K., Flagstad, Ø., Seddon, J.M., Bjornerfeldt, S., Kojola, I., Sand, H., Wabakken, P. & Ellegren, H. (2003) Rescue of a severely bottlenecked wolf (Canis lupus) population by a single immigrant. Proceedings of the Royal Society of London Series B 270: 91-98.

132. Hellborg, L. & Ellegren, H. (2003) Y chromosome conserved anchored tagged sites (YCATS) for the analysis of mammalian male-specific sequences. Molecular Ecology 12: 283-291.


131. Hurst, L.D. & Ellegren, H. (2002) Mystery of the mutagenic male. Nature 420: 365-366.

130. Ellegren, H. (2002) Mismatch repair and mutation biases in microsatellite DNA. Trends in Genetics 18: 552.

129. Seddon, J.M. & Ellegren, H. (2002) MHC class II genes in European wolves: a comparison with dogs. Immunogenetics 54: 590-600.

128. Hipkiss, T., Hörnfeldt, B., Lundmark, A., Norbäck, M. & Ellegren, H. (2002) Sex ratio and age structure of nomadic Tengmalm’s owls: a molecular approach. Journal of Avian Biology 33: 107-110.

127. Ellegren, H. (2002) It took many mares to form the domestic horse. Trends in Genetics 18:500-501.

126. Smith, N.G.C., Webster, M.T. & Ellegren, H. (2002) Deterministic mutation rate variation in the human genome. Genome Research 12: 1350-1356.

125. Webster, M., Smith, N.G.C. & Ellegren, H. (2002) Microsatellite evolution inferred from human-chimpanzee genomic sequence alignment at the Mb-scale level. Proceedings of the National Academy of Sciences USA 99: 8748-8753.

124. Ellegren, H. (2002) Human mutation – blame (mostly) men. Nature Genetics 31: 9-10.

123. Brohede, J., Primmer C.R., Møller A.P. & Ellegren H. (2002) The rate and pattern of germline mutation differ between individual microsatellite loci. Nucleic Acids Research 30: 1997-2003.

122. Ellegren, H. (2002) Microsatellite evolution: a battle between replication slippage and point mutations. Trends in Genetics 18: 70.

121. Ellegren, H. (2002) Dosage compensation: do birds do it as well? Trends in Genetics 18: 25-28.

120. Hellborg, L., Walker, C.W., Rueness, E.K., Stacy, J.E., Kojola, I., Valdmann, H., Vilá, C., Zimmermann, B., Jakobsen, K.S. & Ellegren, H. (2002) Differentiation and levels of genetic variation in northern European lynx (Lynx lynx) populations revealed by microsatellites and mitochondrial DNA analysis. Conservation Genetics 3: 97-111.


119. Montell, H., Fridolfsson, A.K. & Ellegren, H. (2001) Contrasting levels of nucleotide diversity on the avian Z and W sex chromosomes. Molecular Biology and Evolution 18: 2010-2016.

118. Berlin, S. & Ellegren, H. (2001) Clonal co-transmission of mitochondrial DNA and the female-specific avian W chromosome. Nature 413: 37-38.

117. Sundqvist, A.K., Ellegren, H., Olivier, M. & Vilá, C. (2001) Y chromosome haplotyping in Scandinavian wolves (Canis lupus) based on microsatellite markers. Molecular Ecology 10: 1959-1966.

116. Hedrick, P., Fredrickson, R. & Ellegren, H. (2001) Evaluation of d2, a microsatellite measure of inbreeding and outbreeding, in wolves with a known pedigree. Evolution 6:1256-1260.

115. Lindgren, G, Swinburne, J.E., Breen, M., Mariat, D., Sandberg, K., Guerin, G., Ellegren, H., & Binns, M.M. (2001) Physical anchorage and orientation of equine linkage groups by FISH mapping BAC clones containing microsatellite markers. Animal Genetics 32: 37-39.

114. Lindgren, G., Breen, M., Godard, S., Bowling, A., Murray, J., Scavone, M., Skow, L., Sandberg, K., Guerin, G., Binns, M. & Ellegren, H. (2001) Mapping of 13 horse genes by fluorescence in-situ hybridization (FISH) and somatic cell hybrid analysis. Chromosome Research 9: 53-59.

113. Ellegren, H. (2001) Hens, cocks and avian sex determination - A quest for genes on Z or W? EMBO Reports 2: 192-196.

112. Vila, C., Ellegren, H,, Götherström, A., Leonard, J.A. & Wayne, R.K. (2001) Tales from the DNA of domestic horses – Reply. Science 292: 218-219.

111. Vilà, C., Leonard, J.A., Götherström, A., Marklund, S., Sandberg, K., Lidén, K., Wayne, R.K. & Ellegren, H. (2001) Widespread origins of domestic horse lineages. Science 291: 474-477.

110. Saetre, G.P., Borge, T., Lindell, J., Moum, T., Primmer, C.R., Sheldon, B.C., Haavie, J., Johnsen, A. & Ellegren. H. (2001) Speciation, introgressive hybridization and nonlinear rate of molecular evolution in flycatchers. Molecular Ecology 10: 737-749.

109. Ellegren, H. & Carmichael, A. (2001) Multiple and independent cessation of recombination between avian sex chromosomes. Genetics 158: 325-331.

108. Walker, C.W., Vilà, C., Landa, A., Lindén, M. & Ellegren, H. (2001) Genetic variation and structure in Scandinavian wolverine (Gulo gulo) populations. Molecular Ecology 10: 53-62.


107. Ellegren, H. (2000) Microsatellite mutations in the germline: implications for evolutionary inference. Trends in Genetics 16: 551-558.

106. Carmichael, A.N., Fridolfsson, A.K. & Ellegren, H. (2000) Male-biased mutation rates revealed from Z- and W-chromosome linked ATP synthase a-subunit (ATP5A1) sequences in birds. Journal of Molecular Evolution 50: 443-447.

105. Fridolfsson, A.K. & Ellegren, H. (2000) Molecular evolution of the avian CHD1 genes on the Z and W sex chromosomes. Genetics 155: 1903-1912.

104. Hörnfeldt, B, Hipkiss, T, Fridolfsson, AK, Eklund, U & Ellegren, H. (2000) Sex ratio and fledging success of supplementary-fed Tengmalm's owl broods. Molecular Ecology 9: 187-192.

103. Ellegren, H. (2000) Heterogeneous mutation processes in human microsatellite DNA sequences. Nature Genetics 24: 400-402.

102. Ellegren, H. (2000) Evolution of the avian sex chromosomes and their role in sex determination. Trends in Ecology and Evolution 15: 188-192.

101. Berlin, S., Merila, J. & Ellegren, H. (2000) Isolation and characterization of polymorphic microsatellite loci in the common frog, Rana temporaria. Molecular Ecology 9: 1938-1939.


100. Vila, C., Amorim, I.R., Leonard, J.A., Posada, D., Castroviejo, J., Petrucci-Fonseca, F., Crandall, K.A., Ellegren, H. & Wayne, R.K. (1999) Mitochondrial DNA phylogeography and population history of the grey wolf Canis lupus. Molecular Ecology 8: 2089-2103.

99. Guérin, G. et al. (33 additional authors) (1999) Report of the international equine gene mapping work shop: male linkage map. Animal Genetics 30: 341-354. 98. Ellegren, H. (1999) Inbreeding and relatedness in Scandinavian grey wolves. Hereditas 130: 239-244.

97. Ellegren, H., Carlsson, A. & Stenberg, I. (1999) Genetic structure and variability of white-backed woodpecker (Dendrocopus leucotos) populations in northern Europe. Hereditas 130: 291-299.

96. Saino, N., Ellegren, H. & Møller, A.P. (1999) No evidence for adjustment of sex allocation in relation to paternal ornamentation and paternity in barn swallows. Molecular Ecology 8: 399-406.

95. Brohede, J. & Ellegren, H. (1999) Microsatellite evolution: polarity of substitutions within repeats and neutrality of flanking sequences. Proceedings of the Royal Society London, Series B 266: 825-833.

94. Saino, N., Primmer, C., Ellegren, H. & Møller, A.P. (1999) Breeding asynchrony affects certainty of paternity in the barn swallow (Hirundo rustica). Behavioural Ecology and Sociobiology 45: 211-218.

93. Fridolfsson, A.K. & Ellegren, H. (1999) A simple and universal method for sexing non-ratite birds. Journal of Avian Biology 30: 116-121.

92. Lindgren, G., Persson, H. & Ellegren, H. (1999) Five equine dinucleotide microsatellite loci HTG17, HTG20, HTG21, HTG28 and HTG31. Animal Genetics 30: 70-71.


91. Johnsen, A., Lifjeld, J.T., Rohde, P.A. & Ellegren, H. (1998) Sexual conflict over fertilizations: female bluethroats escape male paternity guards. Behavioural Ecology and Sociobiology 43: 401-408.

90. Sheldon, B.C. & Ellegren, H. (1998) Sexual selection resulting from extra-pair paternity in collared flycatchers. Animal Behaviour 57:285-298.

89. Lidén, K., Götherström, A. & Ellegren, H. (1998) "Alia vero gens ibi moratur Suehans, quae velud Thyringi equis utuntur eximiis" or the excellent horses in Sveland. Laborativ Arkeologi 10-11: 59-64.

88. Hurst, L.D. & Ellegren, H. (1998) Sex differences in the mutation rate. Trends in Genetics 14: 446-452.

87. Sheldon, B.C. & Ellegren, H. (1998) Paternal effort related to experimentally manipulated paternity of male collared flycatchers. Proceedings of the Royal Society London, Series B 265: 1737-1742.

86. Sheldon, B.C., Merilä, J., Lindgren, G. & Ellegren, H. (1998) Gender and environmental sensitivity in nestling collared flycatchers. Ecology 79: 1939-1948.

85. Knott, S.A., Marklund, L., Haley, C.S., Andersson, K., Davies, W., Ellegren, H., Fredholm, M., Hoyheim, B., Hansson, I., Lundström, K., Möller, M. & Andersson, L. (1998) Multiple marker mapping of quantitative trait loci in an outbred cross between wild boar and large white pigs. Genetics 149: 1069-1080.

84. Lindgren, G., Sandberg, K., Persson, H., Marklund, S., Breen, M., Sandgren, B., Carlstén, J. & Ellegren, H. (1998) A primary male autosomal linkage map of the horse genome. Genome Research 8: 951-966.

83. Fridolfsson, A.K., Cheng, H., Copeland, N.G., Jenkins, N.A., Liu, H.C., Raudsepp, T., Woodage, T., Chowdhary, B., Halverson, J. & Ellegren, H. (1998) Evolution of the avian sex chromosomes from an ancestral pair of autosomes. Proceedings of the National Academy of Sciences USA 95: 8147-8152.

82. Merilä, J., Sheldon, B.C. & Ellegren, H. (1998) Quantitative genetics of sexual dimorphism in the collared flycatcher Ficedula albicollis. Evolution 52: 870-876.

81. Primmer, C.R., Saino, N., Møller, A.P. & Ellegren, H. (1998) Unravelling the process of microsatellite evolution through analysis of germline mutations in barn swallows. Molecular Biology and Evolution 15: 1047-1054.

80. Primmer, C.R. & Ellegren, H. (1998) Patterns of molecular evolution at avian microsatellite loci. Molecular Biology and Evolution 15: 997-1008.

79. Lifjeld, J.T., Slagsvold, T. & Ellegren, H. (1998) Experimentally reduced paternity affects paternal effort and reproductive success in pied flycatchers. Animal Behaviour 55: 319-329.


78. Ellegren, H. & Fridolfsson, A.K. (1997) Male-driven evolution of DNA sequences in birds. Nature Genetics 17:182-184.

77. Ellegren, H., Lindgren, G., Primmer, C.R. & Møller, A.P. (1997) Fitness loss and germline mutations in barn swallows breeding in Chernobyl. Nature 389: 393-396.

76. Hasler-Rapacz, J.H., Ellegren, H., Fridolfsson, A.K., Kirkpatrick, B., Kirk, S., Andersson, L. & Rapacz, J. (1997) Positional candidate cloning of a gene for familial hypercholesterolemia in swine. American Journal of Medical Genetics 76:379-386.

75. Fridolfsson, A.K., Hori, T., Winterø, A.K., Fredholm, M., Yerle, M., Robic, A., Andersson, L. & Ellegren, H. (1997) Expansion of the pig comparative map by expressed sequenced tags (EST) mapping. Mammalian Genome 8: 907-912.

74. Merilä, J., Sheldon, B.C. & Ellegren, H. (1997) Antagonistic natural selection revealed by molecular sex identification of nestling collared flycatchers. Molecular Ecology 6: 1167-1175.

73. Ellegren H. & Sheldon B.C. (1997) Flow cytometry for sexing birds – Reply. Trends in Ecology and Evolution 12: 489-490.

72. Ellegren, H., Moore, S., Robinson, N., Byrne, K., Ward, W. & Sheldon, B.C. (1997) Microsatellite evolution - a reciprocal study of repeat lengths at homologous loci in cattle and sheep. Molecular Biology and Evolution 14: 854-860.

71. Primmer, C.R., Raudsepp, T., Chowdhary, B. & Ellegren, H. (1997) Low frequency of microsatellites in the avian genome. Genome Research 7: 471-482.

70. Ellegren, H. & Sheldon, B. (1997) New tools for sex identification and the study of sex allocation in birds. Trends in Ecology and Evolution 12: 255-259.

69. Breen, M., Lindgren, G., Binns, M.M., Norman, J., Irvin, Z., Bell, K., Sandberg, K. & Ellegren, H. (1997) Genetical and physical assignments of equine microsatellite - first integration of anchored markers in horse genome mapping. Mammalian Genome 8: 267-273.

68. Saino, N., Primmer, C.R., Ellegren, H. & Møller, A.P. (1997) An experimental study of paternity and tail ornamentation in the barn swallow. Evolution 51: 562-570.

67. Lifjeld, J.T., Slagsvold, T. & Ellegren, H. (1997) Experimental mate switching in pied flycatchers: male copulatory access and fertilization success. Animal Behaviour 53: 1225-1232.

66. Lifjeld, J.T., Slagsvold, T., Dale, S. & Ellegren, H. (1997) A sexually selected paradox in the pied flycatcher: attractive males are cuckolded. Auk 114: 112-115.

65. Sheldon, B.C., Merilä, J., Qvarnström, A. & Ellegren, H. (1997) Female genetic benefit from extra-pair copulation predicted by relative size of male secondary sexual character. Proceedings of the Royal Society London, Series B 264: 297-302.


64. Ellegren, H. (1996) First gene on the avian W chromosome provides a tag for universal sexing of non-ratite birds. Proceedings of the Royal Society London, Series B 263: 1635-1641.

63. Ellegren, H., Savolainen, P. & Rosén, B. (1996) The genetical history of an isolated population of the endangered grey wolf Canis lupus: a study using nuclear and mitochondrial polymorphisms. Philosophical Transactions of the Royal Society London, Series B 351: 1661-1669.

62. Sheldon, B.C. & Ellegren, H. (1996) Offspring sex ratio and paternity in collared flycatcher. Proceedings of the Royal Society London, Series B 263: 1017-1021.

61. Primmer, C.R., Møller, A.P. & Ellegren, H. (1996) New microsatellites from the pied flycatcher and the swallow genomes. Hereditas 124: 281-283.

60. Ellegren, H., Gustafsson, L. & Sheldon, B.C. (1996) Sex ratio adjustment in relation to paternal attractiveness in a wild bird population. Proceedings of the National Academy of Sciences USA 93: 11723-11728.

59. Primmer, C.R., Saino, N., Møller, A.P. & Ellegren, H. (1996) Directional evolution in germline microsatellite mutations. Nature Genetics 13: 391-393.

58. Marklund, L., Johansson Möller, M., Høyheim, B., Davies, W., Fredholm, M., Juneja, R.K., Mariani, P., Coppieters, W., Ellegren, H. & Andersson, L. (1996) A comprehensive linkage map of the pig based on a wild pig - Large White intercross. Animal Genetics 27: 255-269.

57. Mariani, P., Johansson Möller, M., Høyheim, B., Marklund, L., Davies, W., Ellegren, H. & Andersson, L. (1996) Assignment of the extension (E) coat color locus to pig chromosome 6. Journal of Heredity 87: 272-276.

56. Primmer, C., Møller, A.P., & Ellegren, H. (1996) A wide-range survey of microsatellite cross-species amplification in birds. Molecular Ecology 5: 365-378.

55. Ellegren, H., Mikko, S., Wallin, K. & Andersson, L. (1996) Limited polymorphism at major histocompatibility complex (MHC) loci in the Swedish moose. Molecular Ecology 5: 3-10.


54. Larsen, N.J., Ellegren, H., Bräuner-Nielsen, P. & Andersson, L. (1995) Genetic variation at the growth hormone locus in a wild pig intercross; test of association to phenotypic traits and linkage to the blood group D locus. Theoretical and Applied Genetics 91: 1074-1077.

53. Ellegren, H., Primmer, C.R. & Sheldon, B.C. (1995) Microsatellite evolution: directionality or bias? Nature Genetics 11: 360-362.

52. Ellegren, H., Lifjeld, J.T., Slagsvold, T. & Primmer, C.R. (1995) Handicapped males and extrapair paternity in pied flycatchers: a study using microsatellite markers. Molecular Ecology 4: 441-447.

51. Paszek, A.A. et al. (a total of 21 authors) (1995) First international workshop on porcine chromosome 6. Animal Genetics 26: 377-401.

50. Ellegren, H. & Basu, T. (1995) Filling the gaps in the porcine linkage map: isolation of microsatellites from chromosome 18 using flow-sorting and SINE-PCR. Cytogenetics and Cell Genetics 71: 370-373.

49. Edfors-Lilja, I., Gustafsson, U., Duval-Iflah, Y., Ellegren, H., Johansson, M., Juneja, R.K., Marklund, L. & Andersson, L. (1995) The porcine intestinal receptor for Escherichia coli K88ab, K88ac; Regional localization on chromosome 13 and influence in IgG response to the K88 antigen. Animal Genetics 26: 237-242.

48. Primmer, C., Møller, A.P., & Ellegren, H. (1995) Resolving genetic relationships with microsatellite markers: A parentage testing system for the swallow Hirundo rustica. Molecular Ecology 4: 493-498.

47. Ellegren, H. (1995) Mutation rates at porcine microsatellite loci. Mammalian Genome 6: 376-377.

46. Archibald, A. et al. (a total of 50 authors). (1995) The PiGMaP consortium linkage map of the pig (Sus scrofa). Mammalian Genome 6: 157-175.

45. Johansson, M., Ellegren, H. & Andersson, L. (1995) Comparative mapping reveals extensive linkage conservation but also gene order rearrangements between the pig and human genomes. Genomics 25: 682-690.


44. Johansson, M., Ellegren, H., Marklund, L., Coppieters, W. & Andersson, L. (1994) Linkage maps of porcine chromosomes 3, 6 and 9 in pigs based on 31 polymorphic markers. Mammalian Genome 5: 785-790.

43. Ellegren, H., Chowdhary, B.P., Fredholm, M., Høyheim, B., Johansson, M., Bräuner-Nielsen, P., Thomsen, P.D. & Andersson, L. (1994) A physically anchored linkage map of pig chromosome 1 uncovers sex- and position-specific recombination rates. Genomics 24: 342-350.

42. Andersson, L., Edfors-Lilja, I., Ellegren, H., Johansson, M. & Marklund, L. (1994) Mapping trait loci by crossbreeding genetically divergent populations of domestic animals. Animal Biotechnology 5: 225-231.

41. Ellegren, H., Chowdhary, B.P., Johansson, M., Marklund, L., Fredholm, M., Gustavsson, I. & Andersson, L. (1994) A primary linkage map of the porcine genome reveals a low rate of genetic recombination in the pig. Genetics 137: 1089-1100.

40. Ellegren, H., Chowdhary, B., Johansson, M. & Andersson, L. (1994) Integrating the porcine physical and linkage map using cosmid-derived markers. Animal Genetics 25: 155-164.

39. Ellegren, H., Johansson, M., Hartman, G. & Andersson, L. (1994) DNA fingerprinting with the human 33.6 minisatellite probe identifies sex in beavers Castor fiber. Molecular Ecology 3: 273-274.

38. Andersson, L., Haley, C.S., Ellegren, H., Knott, S.A., Johansson, M., Andersson, K., Andersson-Eklund, L., Edfors-Lilja, I., Fredholm, M., Hansson, I., Håkansson, J. & Lundström, K. (1994) Genetic mapping of quantitative trait loci for growth and fatness in pigs. Science 263: 1771-1774.

37. Chowdhary, B.P., Ellegren, H., Johansson, M., Andersson, L. & Gustavsson, I. (1994) In situ hybridization mapping of the growth hormone receptor (GHR) gene assigns a linkage group (C9, FSA, GHR and S0105) to chromosome 16 in pigs. Mammalian Genome 5: 160-162.

36. Ellegren, H. (1994) Linkage mapping of the APOA1 gene to pig chromosome 9. Mammalian Genome 5: 58-59.

35. Marklund, S., Ellegren, H., Eriksson, S., Sandberg, K. & Andersson, L. (1994) Parentage testing and linkage analysis in the horse using a set of highly polymorphic microsatellites. Animal Genetics 25: 19-23.


34. Ellegren, H. (1993) Speed of migration and migratory flight lengths of passerine birds ringed during autumn migration in Sweden. Ornis Scandinavica 24: 220-228.

33. Holmgren, N., Ellegren, H. & Pettersson, J. (1993) Stopover length, body mass and fuel deposition rate in autumn migrating adult Dunlins Calidris alpine: evaluating the effects of moulting status and age. Ardea 81: 9-20.

32. Holmgren, N., Ellegren, H. & Pettersson, J. (1993) The adaptation of moult pattern in migratory Dunlins Calidris alpine. Ornis Scandinavica 24: 21-27.

31. Ellegren, H. (1993) Abundant (A)n•(T)n mononucleotide repeats in the pig genome: linkage mapping of the porcine APOB, FSA, ALOX12, PEPN and RLN loci. Animal Genetics 24: 367-372.

30. Edfors-Lilja, I., Ellegren, H., Winterø, A.K., Ruohonen-Lehto, M., Fredholm, M., Gustavsson, U., Juneja, R.K. & Andersson, L. (1993) A large linkage group on pig chromosome 7 including the MHC class I, class II (DQB), and class III (TNFB) genes. Immunogenetics 38: 363-366.

29. Ellegren, H., Hartmann, G., Johansson, M. & Andersson, L. (1993) Major histocompatibility complex monomorphism and low levels of DNA fingerprinting variability in a reintroduced and rapidly expanding population of beavers. Proceedings of the National Academy of Sciences (USA) 90: 8150-8153.

28. Chowdhary, B.P., Johansson, M., Chaudhary, R., Ellegren, H., Gu, F., Andersson, L. & Gustavsson, I. (1993) In situ hybridization mapping and RFLP analysis of the porcine albumin (Alb) and transferrin (Tf) genes. Animal Genetics 24: 85-90.

27. Johansson, M., Chowdhary, B., Gu, F., Ellegren, H., Gustavsson, I. & Andersson, L. (1993) Genetic analysis of the gene for porcine submaxillary gland mucin (MUC): physical assignment of the MUC and interferon gamma (IFNG) genes to chromosome 5. Journal of Heredity 84: 259-262.

25. Ellegren, H. (1993) Variable SINE 3' poly(A) sequences in the pig genome: an abundant class of genetic markers. Mammalian Genome 4: 429-434.

26. Ellegren, H. (1993) Genomic DNA from museum bird feathers. In: Ancient DNA (B. Herrman & S. Hummel, Eds.), pp. 207-213, Springer Verlag, New York.

24. Ellegren, H., Fredholm, M., Edfors-Lilja, I., Winterø, A.K. & Andersson, L. (1993) Conserved synteny between pig chromosome 8 and human chromosome 4 but rearranged and distorted linkage maps. Genomics 17: 599-603.

23. Ellegren, H., Davies, C.J. & Andersson, L. (1993) Strong association between polymorphisms in an intronic microsatellite and in the coding sequence of the BoLA-DRB3 gene. Implications for microsatellite stability and PCR-based DRB3 typing. Animal Genetics 24: 269-276.

22. Ellegren, H., Johansson, M., Chowdhary, B.P., Marklund, S., Brunnberg, L., Bräuner-Nielsen, P., Edfors-Lilja, I., Gustavsson, I., Juneja, R.K. & Andersson, L. (1993) Assignment of 20 microsatellite markers to the porcine linkage map. Genomics 16: 431-439.

21. Lagercrantz, U., Ellegren, H. & Andersson, L. (1993) The abundance of various polymorphic simple repeat motifs differ between plants and vertebrates. Nucleic Acids Research 21: 1111-1115.


20. Ellegren, H. (1992) Effects of age and sex on the fat-free body mass in autumn migrating Bluethroats Luscinia s. svecica. Ardea 80: 255-259.

19. Ellegren, H. & Fransson, T. (1992) Fat loads and estimated flight ranges in four Sylvia warblers analysed on autumn migration at Gotland, SE Sweden. Ringing & Migration 13: 1-12.

18. Johansson, M., Ellegren, H., Marklund, L., Gustavsson, U., Ringmar-Cederberg, E., Andersson, K., Edfors-Lilja, I. & Andersson, L. (1992) The gene for dominant white color in the pig is closely linked to ALB and PDGFRA on chromosome 8. Genomics 14: 965-969.

17. Ellegren, H. (1992) Polymerase chain reaction (PCR) analysis of avian microsatellites - a new approach to genetic studies in birds. Auk 109: 886-895.

16. Ellegren, H. (1992) Population genetics of museum birds by hypervariable microsatellite typing of genomic DNA prepared from museum feathers. Ancient DNA Newsletter 1: 10.

15. Johansson, M., Ellegren, H. & Andersson, L. (1992) Cloning of porcine microsatellites. Journal of Heredity 83: 196-198.

14. Ellegren, H., Johansson, M., Sandberg, K. & Andersson, L. (1992) Cloning of highly polymorphic microsatellites in the horse. Animal Genetics 23: 133-142.

13. Ellegren, H., Andersson, L., Johansson, M. & Sandberg, K. (1992) DNA fingerprinting in horses using a simple (TG)n probe and its application to population comparisons. Animal Genetics 23: 1-9.

12. Mariani, P., Johansson, M., Ellegren, H., Harbitz, I., Juneja, R.K. & Andersson, L. (1992) Multiple RFLPs in the porcine calcium release channel gene (CRC): assignment to the halothane (HAL) linkage group. Animal Genetics 23: 257-262.


11. Ellegren, H. (1991) DNA typing of museum birds. Nature 354: 113.

10. Ellegren, H. (1991) Stopover ecology in autumn migrating Bluethroats Luscinia s. svecica in relation to age and sex. Ornis Scandinavica 22: 340-348.

9. Ellegren, H. & Wallin, K. (1991) Autumn migrating Bluethroats Luscinia s. svecica orient in a east-southeastern direction at Gävle, East Sweden. Ornis Svecica 1: 47-50.

8. Ellegren, H., Andersson, L. & Wallin, K. (1991) DNA polymorphism in the moose (A. alces) revealed by the polynucleotide probe (TC)n. Journal of Heredity 82: 429-431.

7. Ellegren, H. (1991) Fingerprinting birds DNA with a polynucleotide probe (TG)n. Auk 108: 956-958.

6. Mariani, P., Johansson, M., Ellegren, H. & Andersson, L. (1991) A TaqI RFLP at the porcine platelet-derived growth factor ß-receptor locus (PDGFRB). Animal Genetics 22: 446.

5. Mariani, P., Johansson, M., Ellegren, H. & Andersson, L. (1991) TaqI and PvuII RFLPs at the porcine plasminogen activator, urokinase, locus (PLAU). Animal Genetics 22: 447.

4. Kågedahl, L., Engström, B., Ellegren, H., Lieber, A.K., Lundström, H., Persson, A. & Schenning, M. (1991) Chemical, physical and chromatographic properties of Superdex 75 prep grade and Superdex 200 prep grade gel filtration media. Journal of Chromatography 537: 17-32.


3. Ellegren, H. (1990) Timing of autumn migration in Bluethroats Luscinia s. svecica depends on timing of breeding. Ornis Fennica 67: 13-17.

2. Ellegren, H. (1990) Autumn migration speed in Scandinavian Bluethroats Luscinia s. svecica. Ringing & Migration 11: 121-131.


1. Ellegren, H. & Låås, T. (1989) Size-exclusion chromatography of DNA restriction fragments; fragment length determinations and a comparison with the behaviour of proteins in size-exclusion chromatography. Journal of Chromatography 467: 217-226.