Comparative Developmental Biology

Publications Hejnol Group

Research Articles 




Reviews and Book Chapters











  • Grande C, Martín-Durán JM, Kenny NJ, Truchado-García M, Hejnol A (2014). Evolution, divergence and loss of the Nodal signalling pathway: new data and a synthesis across the Bilateria. Int J Dev Biol 58:521-532 
  • Bekkouche N, Kristensen RM, Hejnol A, Sørensen, MV, Worsaae K (2014).  Detailed reconstruction of the musculature in Limnognathia maerski (Micrognathozoa) and comparison with other GnathiferaFront Zool 11:71 - link 
  • Børve A, Hejnol A (2014).  Development and juvenile anatomy of the nemertodermatid Meara stichopi (Bock) Westblad 1949 (Acoelomorpha). Front Zool 11:50
  • Schnitzler CE, Simmons DK, Pang K, Martindale MQ and Baxevanis AD (2014).  Expression of multiple Sox genes through embryonic development in the ctenophore Mnemiopsis leidyi is spatially restricted to zones of cell proliferation EvoDevo 5:15  


  • Ryan JF, Pang K, Schnitzler CE, Nguyen AD, Moreland RT, Simmons DK, Koch BJ, Francis WR, Havlak P, NISC Comparative Sequencing Program, Smith SA, Putnam NH, Haddock SHD, Dunn CW, Wolfsberg TG, Mullikin JC, Martindale MQ, Baxevanis AD (2013). The Genome of the Ctenophore Mnemiopsis leidyi and Its Implications for Cell Type Evolution Science 342:1242592 
  • Flot JF, Hespeels B, Li X, Noel B, Arkhipova I, Danchin EGJ, Hejnol A, Henrissat B, Koszul R, Aury JM, Barbe V, Barthélémey R-M, Bast J, Bazykin GA, Chabrol O, Couloux A, Da Rocha M, Da Silva C, Gladyshev E, Gouret P, Hallatschek O, Hecox-Lea B, Lebadie K, Lejeune B, Piskurek O, Poulain J, Rodriguez F, Ryan JF, Vakhrusheva OA, Wajnberg E, Wirth B, Yushenova I, Kellis M, Kondrashov AS, Mark Welch DB, Pontarotti P, Weissenbach J, Wincker P, Jaillon O, Van Doninck K (2013).  Genomic evidence for ameiotic evolution in the bdelloid rotifer Adineta vagaNature 500:453-457 
  • Martín-Durán JM, Mendoza A, Sebé-Pedrós A, Ruiz-Trillo I, Hejnol A (2013).  A broad genomic survey reveals multiple origins and frequent losses in the evolution of respiratory hemerythrins and hemocyanins Genome Biol Evol 5:1435-1442 - abstract 
  • Chiodin M, Børve A, Berezhikov E, Ladurner P, Martinez P, Hejnol A (2013). Mesodermal gene expression in the acoel Isodiametra pulchra indicates a low number of mesodermal cell types and the endomesodermal origin of the gonads. PLoS ONE 8(2):e55499 


  • Schnitzler CE, Pang K, Powers ML, Reitzel AM, Ryan JF, Simmons D, Tada T, Park M, Gupta J, Brooks SY, Blakesley RW, Yokoyama S, Haddock SHD, Martindale MQ and Baxevanis AD (2012).  Genomic organization, evolution, and expression of photoprotein and opsin genes in Mnemiopsis leidyi: a new view of ctenophore photocytes BMC Biol 10:107 - abstract 
  • DuBuc TQ, Ryan JF, Shinzato C, Satoh N, Martindale MQ (2012). Coral Comparative Genomics Reveal Expanded Hox Cluster in the Cnidarian–Bilaterian AncestorIntegr Comp Biol 52(6):835-841 
  • Martín-Durán JM, Janssen R, Wennberg S, Budd GE, Hejnol A (2012). Deuterostomic Development in the Protostome Priapulus caudatus.  Curr Biol 22:2161-2166 - abstract 
  • Maxwell EK, Ryan JF, Schnitzler CE, Browne WE and Andreas D Baxevanis (2012).  MicroRNAs and essential components of the microRNA processing machinery are not encoded in the genome of the ctenophore Mnemiopsis leidyi BMC Genomics 13:714 - abstract 
  • Santagata S, Resh C, Hejnol A, Martindale MQ, Passamaneck, YJ (2012).  Development of the larval anterior neurogenic domains of Terebratalia transversa (Brachiopoda) provides insights into the diversification of larval apical organs and the spiralian nervous systemEvoDevo 3:3  


  • Fuchs J, Martindale MQ and Hejnol A (2011) Gene expression in bryozoan larvae suggest a fundamental importance of pre-patterned blastemic cells in the bryozoan life-cycle EvoDevo 2:13 
  • Passamaneck YJ, Furchheim N, Hejnol A, Martindale MQ, Lüter C (2011). Ciliary photoreceptors in the cerebral eyes of a protostome larva EvoDevo 2:6 - abstract


  • Hejnol A, Martindale MQ (2009).  Coordinated spatial and temporal expression of Hox genes during embryogenesis in the acoel Convolutriloba longifissura. BMC Biol 7(1):65 - abstract


Reviews and Book Chapters

  • Annunziata R, Andrikou C, Perillo M (2019). Development and evolution of gut structures: from molecules to function. Cell and Tissue Research doi 10.1007/s00441-019-03093-9 
  • Hejnol A (2017). Ladders, trees, complexity and other metaphors in evolutionary thinking. (In "Arts of Living on a Damaged Planet: Ghosts and Monsters of the Anthropocene", eds. Tsing AL, Swanson HA, Gan E, Brubandt N, Minnesota University Press, Minnesota, pp. 110–117) 
  • Hejnol A, Vellutini BC (2017) Larval Evolution: I'll tail you later... Curr Biol 29 R21-R23  
  • Hejnol A, Pang K (2016).  Xenacoelomorpha’s significance for understanding bilaterian evolution Curr Opin Dev Gen 39:48-54 
  • Hejnol A, Dunn CW (2016).  Animal Evolution: Are Phyla Real? Curr Biol 26:R424-R426 
  • Arnone MI, Hejnol A (2015).  Genomics going wild: Marine sampling for studies of evolution and development Mar Genomics, doi:10.1016/j.margen.2015.11.003 
  • Hejnol A, Lowe CJ (2015).  Embracing the comparative approach: how robust phylogenies and broader developmental sampling impacts the understanding of nervous system evolution Phil Trans R Soc B 370:20150045 
  • Adell T, Martín-Durán JM, Salo E, Cebria F (2015).  Platyhelminthes. Evolutionary Developmental Biology of Invertebrates Volume 2, ed. Wanninger A, Springer Verlag, pp. 21-40. 
  • Hejnol A, Rentzsch F (2015).  Primer: Neural Nets. Curr Biol 25:R782-R786 
  • Hejnol A (2015). Acoelomorpha. Structure and Evolution of Invertebrate Nervous Systems eds. Schmidt-Rhaesa A, Hartzsch S, Purschke G, Oxford University Press, Oxford, pp. 56-61. 
  • Hejnol A (2015). Acoelomorpha and Xenoturbellida. Evolutionary Developmental Biology of InvertebratesVolume 1, ed. Wanninger A., Springer Verlag, pp. 203-14. 
  • Hejnol A (2015). Cycloneuralia. Evolutionary Developmental Biology of Invertebrates Volume 3, ed. Wanninger A, Springer Verlag, pp. 1-13. 
  • Hejnol A (2015). Gnathifera. Evolutionary Developmental Biology of Invertebrates Volume 2, ed. Wanninger A, Springer Verlag, pp. 1-12. 
  • Hejnol A (2015). Gastrotricha. Evolutionary Developmental Biology of Invertebrates Volume 2, ed. Wanninger A, Springer Verlag, pp. 13-19. 
  • Hejnol A, Martín-Durán JM (2015). Getting to the bottom of anal evolution. Zool Anz, doi: 10.1016/j.jcz.2015.02.006 - link 
  • Hejnol A, Lowe C (2014).  Animal Evolution: Stiff or Squishy Notochord Origins? Curr Biol 24:R1131-R1133 
  • Dunn CW, Giribet G, Edgecombe GD, Hejnol A (2014). Animal Phylogeny and Its Evolutionary Implications. Annu Rev Ecol Evol Syst 45:371-395 
  • Hejnol A (2014). Evolutionary Biology: Excitation about jelly nerves. Nature, 510, doi:10.1038/nature13340, Nature Podcast Not so Simple 
  • Martín-Durán JM, Hejnol A (2014).  Priapulida (penis worms): Implications of embryonic development. 306-309 In: McGraw-Hill 2014 Yearbook of Science & Technology, McGraw-Hill Education, New York 
  • Hejnol A (2012). Evolutionary Biology: Muscle's dual origins. Nature 487:181-182. 
  • Martín-Durán JM, Egger B (2012).  Developmental diversity in free-living flatworms. EvoDevo, 3:7 - abstract 
  • Edgecombe GD, Giribet G, Dunn CW, Hejnol A, Kristensen RM, Neves RC, Rouse GW, Worsaae K, Sørensen MV. (2011) Higher-level metazoan relationships: recent progress and remaining questions. Org Divers EvolVolume 11, Number 2, 151-172. 
  • Hejnol A (2011).  Mark Q. Martindale: Shedding new light on developmental diversity Int J Dev Biol 55:237-242 
  • Hejnol A (2010).  A Twist in Time--The Evolution of Spiral Cleavage in the Light of Animal Phylogeny Integr Comp Biol 50:695-706, doi: 10.1093/icb/icq103


Earlier Publications

  • Hejnol A, Obst M, Stamatakis A, Ott M, Rouse GW, Edgecombe GD, Martinez P, Baguñà J, Bailly X, Jondelius U, Wiens M, Müller, WEG, Seaver E, Wheeler WC, Martindale MQ, Giribet G, Dunn CW (2009). Assessing the root of bilaterian animals with scalable phylogenomic methods. Proc R Soc B 276:4261-4270
  • Hejnol A and Martindale MQ (2009). The mouth, the anus and the blastopore - open questions about questionable openings. Animal Evolution: Genomes, Fossils and Trees, eds. Littlewood D.T.J. & Telford, M.J., Oxford University Press, pp. 33-40. (pdf) 
  • Martindale MQ and Hejnol A (2009).  A developmental perspective: changes in the position of the blastopore during bilaterian evolution.  Dev Cell 17:162-174. 
  • Bourlat SJ and Hejnol A (2009).  Quick Guide: Acoels.  Curr Biol 19:R279-R280 
  • Hejnol A and Martindale MQ (2009).  The mouth, the anus and the blastopore - open questions about questionable openings. In: “Animal Evolution: Genomes, Fossils and Trees”, eds. Littlewood D.T.J. & Telford, M.J., Oxford University Press, pp. 33-40. 
  • Giribet G, Dunn CW, Edgecombe GD, Hejnol A, Martindale MQ, Rouse GW (2009).  Assembling the Spiralian Tree of Life. In: “Animal Evolution: Genomes, Fossils and Trees”, eds. Littlewood D.T.J. & Telford, M.J., Oxford University Press, pp. 52-64. 
  • Hejnol A and Martindale MQ (2008).  Acoel development indicates the independent evolution of the bilaterian mouth and anus.  Nature, 456, 382-386. (listen to the nature podcast
  • Dunn CW, Hejnol A, Matus DQ, Pang K, Browne WE, Smith SA, Seaver E, Rouse GW, Obst M, Edgecombe GD, Sørensen MV, Haddock SHD, Schmidt-Rhaesa A, Okuso A, Kristensen RM, Wheeler WC, Martindale MQ, Giribet G (2008).  Broad phylogenomic sampling improves resolution of the Animal Tree of Life.  Nature, 452, 745-749. 
  • Hejnol A and Martindale MQ (2008).  Acoel development supports a simple planula-like urbilaterian.  Phil Trans R Soc B, 363:1493-1501. 
  • Henry JQ, Hejnol A, Perry KJ, Martindale MQ (2007).  Homology of ciliary bands in spiralian trochophores.  Integ Comp Biol, 47:865-871. 
  • Hejnol A, Martindale MQ, Henry JQ (2007).  High-resolution fate map of the snail Crepidula fornicata: the origins of ciliary bands, nervous system, and muscular elements.  Dev Biol 305:63-76. 
  • Hejnol A, Schnabel R, Scholtz G (2006).  A 4D-microscopic analysis of the germ band in Porcellio scaber (Malacostraca, Peracarida) – developmental and phylogenetic implications.  Dev Genes Evol 216:755-767 
  • Matus DQ, Copley RR, Dunn CW, Hejnol A, Eccleston H, Halanych KH, Martindale MQ, Telford MJ (2006).  Broad taxon and gene sampling indicate that chaetognaths are protostomes.  Curr Biol, 16:R575-R576 
  • Schnabel R, Bischoff M, Hintze A, Schulz A-K, Hejnol A, Meinhardt H, Hutter H (2006).  Global cell sorting in the C. elegans embryo defines a new mechanism for pattern formation.  Dev Biol 294:418-431 
  • Hejnol A and Schnabel R (2006).  What a couple of dimensions can do for you: Comparative developmental studies using 4D-microscopy – examples from tardigrade development.  Integ Comp Biol 46:151–161. 
  • Hejnol A and Schnabel R (2005).  The eutardigrade Thulinia stephaniae has an indeterminate development and the potential to regulate early blastomere ablations.  Development 132:1349-1361. 
  • Hejnol, A., Scholtz, G. (2004) Clonal analysis of Distal-less and engrailed expression patterns during early morphogenesis of uniramous and biramous crustacean limbs. Dev Genes Evol 214:473-485. 
  • Hejnol, A., Pfannenstiel, H.-D. (1998) Myosin and actin are necessary for polar lobe formation and resorption in Ilyanassa obsoleta embryos. Dev Genes Evol 208:229-233.