Standard Event System

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The "Standard Event System" (SES) to Study Vertebrate Embryos was developed in 2009 to establish a common language in comparative embryology.[1] Homologous developmental characters are defined therein and should be recognisable in all vertebrate embryos. The SES includes a protocol on how to describe and depict vertebrate embryonic characters. The SES was initially developed for external developmental characters of organogenesis, particularly for turtle embryos. However, it is expandable both taxonomically and in regard to anatomical or molecular characters. This article should act as an overview on the species staged with SES and document the expansions of this system. New entries need to be validated based on the citation of scientific publications. The guideline on how to establish new SES-characters and to describe species can be found in the original paper of Werneburg (2009).[1]

SES-staged species[edit]

Overview on the vertebrate species staged with SES. "Original" means that the SES-staging was not derived from an other staging system.

Major taxon Species Reference Original
Lissamphibia Ambystoma mexicanum [2] no
Testudines Emydura subglobosa [3] yes
Testudines Graptemys nigrinoda [2] yes
Testudines Apalone spinifera, Caretta caretta, Carettochelys insculpta, Chelonia mydas, Chelydra serpentina, Chrysemys picta, Dermochelys coriacea, Eretmochelys imbricata, Lepidochelys olivacea, Natator depressus, Testudo hermanni, Trachemys scripta [2] no
Crocodilia Alligator mississippiensis [2] no
Squamata Lacerta vivipara [2] no
Squamata Bothropoides jararaca [4] yes
Squamata Calyptommatus sinebranchiatus [5] yes
Squamata Nothobachia ablephara [5] yes
Aves Gallus gallus [2] no
Mammalia Tachyglossus aculeatus [6] yes/no
Mammalia Dasypus hybridus, Didelphis virginiana [2] no
Mammalia Echinops telfairi, Tenrec ecaudatus [7] yes
Mammalia Mus musculus [7] no


New SES-characters are continuously descripted in new publications. Currently, characters of organogenesis are described for Vertebrata, Gnathostomata, Tetrapoda, Amniota, Sauropsida, Squamata, Mammalia, and Monotremata.


  1. ^ a b Werneburg (2009). A Standard System to Study Vertebrate Embryos. PLoS ONE 4(6): e5887.
  2. ^ a b c d e f g Werneburg I and Sánchez-Villagra MS (2009). Timing of organogenesis support basal position of turtles in the amniote tree of life. BMC Evolutionary Biology, 9:82
  3. ^ Werneburg I, Hugi J, Müller J, Sánchez-Villagra MS (2009). Embryogenesis and ossification of Emydura subglobosa (Testudines, Pleurodira, Chelidae) and patterns of turtle development. Developmental Dynamics, Volume 238, Issue 11 Pages 2770-2786
  4. ^ Polachowski KM and Werneburg I (2013). Late embryos and bony skull development in Bothropoides jararaca (Serpentes, Viperidae). Zoology
  5. ^ a b Roscito and Rodriges (2012). Embryonic development of the fossorial gymnophthalmid lizards Nothobachia ablephara and Calyptommatus sinebranchiatus. Zoology 115:302-318
  6. ^ Werneburg I and Sánchez-Villagra MR (2011). The early development of the echidna, Tachyglossus aculeatus (Mammalia: Monotremata) and patterns of mammalian development. Acta Zoologica. 82(1)
  7. ^ a b Werneburg I, Tzika AC, Hautier L, Asher RJ, Milinkovitch MC, Sánchez-Villagra MR (2013). Development and embryonic staging in non-model organisms: the case of an afrotherian mammal. The Journal of Anatomy 222:2-18

External links[edit]