Standard Event System

The "Standard Event System" (SES) to Study Vertebrate Embryos was developed in 2009 to establish a common language in comparative embryology.Werneburg (2009). A Standard System to Study Vertebrate Embryos. PLoS ONE 4(6): e5887. http://www.plosone.org/article/info%3Adoi%2F10.1371%2Fjournal.pone.0005887 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). SES-characters are used to reconstruct ancestral developmental sequences in evolution such as that of the last common ancestor of placental mammals.Werneburg and Spiekman (2016). Mammalian embryology and organogenesis. From gametes to weaning. In: Zachos F., Asher R. (eds.). Mammalia. Series: The Handbook of Zoology / Handbuch der Zoologie. De Gruyter, Berlin Also the plasticity of developmental characters can be documented and analysed.

SES-staged species

Overview on the vertebrate species staged with SES. Major taxon 1 Major taxon 2 Species Reference Lissamphibia Caudata Ambystoma mexicanum 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 http://www.biomedcentral.com/1471-2148/9/82 Testudines Pleurodira Emydura subglobosa 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 http://onlinelibrary.wiley.com/doi/10.1002/dvdy.22104/abstract Testudines Cryptodira Graptemys nigrinoda Testudines Cryptodira Kinosternum scorpionoides Nunes Silva R and Sobral Sampaio F (2014). Immunoreactivity of Mel1a-like melatonin receptor and NRH: Quinone reductase enzyme (QR2) in testudine whole embryo and in developing whole retinas Trends in Developmental Biology 8:39-46. Testudines Cryptodira Apalone spinifera, Caretta caretta, Carettochelys insculpta, Chelonia mydas, Chelydra serpentina, Chrysemys picta, Dermochelys coriacea, Eretmochelys imbricata, Lepidochelys olivacea, Natator depressus, Pelodiscus sinensis, Testudo hermanni, Trachemys scripta Crocodilia Alligatoridae Alligator mississippiensis Squamata Lacertidae Zootoca vivipara Squamata Serpentes Bothropoides jararaca Polachowski KM and Werneburg I (2013). Late embryos and bony skull development in Bothropoides jararaca (Serpentes, Viperidae). Zoology Squamata Gymnophthalmidae Calyptommatus sinebranchiatus, Nothobachia ablephara Roscito and Rodriges (2012). Embryonic development of the fossorial gymnophthalmid lizards Nothobachia ablephara and Calyptommatus sinebranchiatus. Zoology 115:302-318 Squamata Varanidae Varanus panoptes Werneburg et al. (2015). Bony skull development in the Argus monitor (Squamata, Varanidae, Varanus panoptes) with comments on developmental timing and adult anatomy. Zoology 118(4):255-280 Squamata Iguanidae Pogona vitticeps Ollonen, J., Da Silva, F.O., Mahlow, K. and Di-Poï, N., 2018. Skull development, ossification pattern, and adult shape in the emerging lizard model organism Pogona vitticeps: a comparative analysis with other squamates. Frontiers in physiology, 9, p.278. Aves Galliformes Gallus gallus Mammalia Monotremata Tachyglossus aculeatus 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) http://onlinelibrary.wiley.com/doi/10.1111/j.1463-6395.2009.00447.x/abstract Mammalia Monotremata Ornithorhynchus anatinus Werneburg I, Laurin M, Koyabu D, Sánchez-Villagra (2016). Evolution of organogenesis and the origin of altriciality in mammals. Evolution and Development Mammalia Marsupialia Didelphis virginiana Mammalia Marsupialia Monodelphis domestica, Sminthopsis macroura, Dasyurus hallucatus, Dasyurus viverrinus, Phascolarctos cinereus, Petauroides volans, Trichosurus vulpecula, Bettongia gaimardi, Macropus eugenii Mammalia Marsupialia Caenolestes fuliginosus González B, Soria-Escobar AM, Rojas-Díaz V, Pustovrh MC, Monsalve LS, Rougier GW (2020). The embryo of the silky shrew opossum, Caenolestes fuliginosus (Tomes, 1863): First description of the embryo of Paucituberculata. Journal of Morphology 2020: 1-12 Mammalia Xenarthra Dasypus hybridus Mammalia Xenarthra Bradypus tridactylus Mammalia Afrotheria Echinops telfairi, Tenrec ecaudatus 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 Mammalia Afrotheria Elephas maximus, Loxodonta africana Mammalia Laurasiatheria Talpa occidentalis, Talpa europaea, Erinaceus europaeus, Erinaceus roumanicus, Suncus murinus, Crocidura russula, Neomys fodiens, Sorex araneus, Pecari tajacu, Sus scrofa domesticus, Rangifer tarandus, Capreolus capreolus, Bos primigenius taurus, Capra aegagrus hircus, Ovis aries, Phocoena phocoena, Delphinus delphis, Stenella longirostris, Stenella attenuata, Hipposideros armiger, Hipposideros pratti, Molossus rufus, Myotis lucifugus, Myotis myotis, Pipistrellus abramus, Scotophilus kuhlii, Miniopterus natalensis, Miniopterus schreibersii, Carollia perspicillata, Equus ferus caballus, Manis javanica, Felis catus, Mustela putorius, Canis lupus familiaris Mammalia Laurasiatheria Artibeus fimbriatus, Artibeus lituratus, Artibeus obscurus, Carollia perspicillata, Eumops patagonicus, Hipposideros armiger, Hipposideros pratti, Miniopterus natalensis, Miniopterus schreibersii, Molossus rufus, Myotis albescens, Myotis lucifugus, Myotis myotis, Pipistrellus abramus, Rhinolophus ferrumequinum, Rousettus aegyptiacus, Rousettus amplexicaudatus, Scotophilus kuhlii, Vespertilio sinensis, Mus musculus Taro Nojiri, Dai Fukui, Ingmar Werneburg, Takashi Saitoh, Hideki Endo, Daisuke Koyabu (2021). Embryonic staging of bats with special reference to Vespertilio sinensis and its cochlear development. Development Dynamics, DOI10.1002/dvdy.325 Mammalia Boreoeutheria Oryctolagus cuniculus, Spermophilus citellus, Cavia porcellus, Cricetulus barabensis, Mesocricetus auratus, Myodes glareolus, Meriones unguiculatus, Acomys dimidiatus, Apodemus sylvaticus, Apodemus flavicollis, Apodemus agrarius, Rattus norvegicus, Tupaia javanica, Tupaia belangeri, Galago senegalensis, Nycticebus coucang, Tarsius tarsier, Callithrix jacchus, Erythrocebus patas, Papio cynocephalus, Macaca mulatta, Homo sapiens Mammalia Boreoeutheria Mus musculus

SES-characters

New SES-characters are continuously described in new publications. Currently, characters of organogenesis are described for Vertebrata (V), Gnathostomata (G), Tetrapoda (T), Amniota (A), Sauropsida (S), Squamata (SQ), Mammalia (M), and Monotremata (MO). In total, 166 SES-characters are currently defined. Character complex Character Character description Reference Illustration A. Egg (V01). A1. Egg lay (V01a). Most authors begin to create their staging tables shortly after or around egg lay time. B. Blastula (V02). B1. Blastoporus (V02a). The blastoporus begins to form as a lip and is later on visible as a clear depression in the blastula.