vestnik-bio-msuВестник Московского университета. Серия 16. БиологияVestnik Moskovskogo universiteta. Seriya 16. Biologiya0137-0952Lomonosov Moscow State University, School of Biology10.55959/MSU0137-0952-16-80-4-4vestnik-bio-msu-1574Research ArticleОБЗОРREVIEWEctopharyngeata и Endopharyngeata вместо Protostomia и Deuterostomia?Ectopharyngeata and Endopharyngeata instead of Protostomia and Deuterostomia?https://orcid.org/0000-0003-1637-2354МалаховВ. В.MalakhovV. V.

Малахов Владимир Васильевич – докт. биол. наук, проф., акад. РАН, зав. кафедрой

119234, г. Москва, Ленинские горы, д. 1, стр. 12

Тел.: 8-495-939-27-76

Leninskye gory, 1–12, Moscow, 119234

vmalakhov@inbox.ruhttps://orcid.org/0000-0003-4533-7769ЕжоваО. В.EzhovaO. V.

Ежова Ольга Владимировна – канд. биол. наук, доцент кафедры

119234, г. Москва, Ленинские горы, д. 1, стр. 12

Тел.: 8-495-939-27-76

Leninskye gory, 1–12, Moscow, 119234

olga_ejova@mail.ruhttps://orcid.org/0000-0002-4854-2787ГанцевичМ. М.GantsevichM. M.

Ганцевич Михаил Маркович – канд. биол. наук, ст. науч. сотр. кафедры

119234, г. Москва, Ленинские горы, д. 1, стр. 12

Тел.: 8-495-939-27-76

Leninskye gory, 1–12, Moscow, 119234

mgantsevich@gmail.comКафедра зоологии беспозвоночных, биологический факультет, Московский государственный университет имени М.В. ЛомоносоваРоссияDepartment of Invertebrate Zoology, Faculty of Biology, M.V. Lomonosov Moscow State UniversityRussian Federation202518022026804219228Copyright © Малахов В.В., Ежова О.В., Ганцевич М.М., 20262026Малахов В.В., Ежова О.В., Ганцевич М.М.Malakhov V.V., Ezhova O.V., Gantsevich M.M.Данная работа распространяется под лицензией Creative Commons Attribution 4.0.This work is licensed under a Creative Commons Attribution 4.0 License.https://vestnik-bio-msu.elpub.ru/jour/article/view/1574

Использование молекулярно-генетических методов привело к глубокой перестройке системы и филогении трехслойных Bilateria (=Triploblastica). Тем не менее, в большинстве филогенетических деревьев сохраняются две основные клады, совпадающие с Protostomia (первичноротыми) и Deuterostomia (вторичноротыми), установленными К. Гроббеном (Grobben, 1908) более 100 лет назад. Для обеих клад Triploblastica первичным способом закладки сквозного кишечника является амфистомия. Однако анализ показывает, что только происхождение рта и ануса не позволяет провести различие между этими двумя основными кладами. Все группы, которые были отнесены К. Гроббеном к Protostomia, характеризуются эктодермальной глоткой, поэтому эту кладу правильнее было бы назвать Ectopharyngeata. На основе модификации кутикулы глотки у многих эктофарингеальных развиваются различные структуры, предназначенные для измельчения пищи. Группы, отнесенные К. Гроббеном к Deuterostomia, имеют энтодермальную глотку, поэтому правильнее было бы называть эту кладу Endopharyngeata. Синапоморфия Endopharyngeata заключается в развитии жаберных щелей в стенках энтодермальной глотки и подразделении глотки на две зоны – дорсальную жаберную и вентральную пищеварительную. В процессе эволюции хордовых произошла инверсия сторон тела, и та часть пищеварительной трубки, которая гомологична пищеварительной зоне глотки Hemichordata, стала дорсальной хордой, а жаберная зона глотки оказалась вентральной. Таким образом, разделение Triploblastica на две клады полностью сохраняет свое значение в современной биологии, хотя и основано на ином наборе морфологических признаков, чем тот, который рассматривал К. Гроббен.

The use of molecular phylogenetic methods has caused a profound rearrangement of the system and phylogeny of the triploblastic Bilateria (=Triploblastica). Nevertheless, in most phylogenetic trees, two main clades remain, coinciding with Protostomia and Deuterostomia, established by K. Grobben (1908) more than 100 years ago. For both clades of Triploblastica, amphistomy is the primary way of forming a through intestine. However, the analysis shows that only the origin of the mouth and anus does not allow us to distinguish between the two main clades. All the groups that were referred to Grobben’s Protostomia are characterized by an ectodermal pharynx, therefore this clade would be more correctly called Ectopharyngeata. Based on the modification of the pharyngeal cuticle, many ectopharyngeates develop various structures designed to grind food. The groups attributed by K. Grobben to Deuterostomia have an endodermal pharynx, therefore it would be more correct to call this clade Endopharyngeata. The synapomorphy of Endopharyngeata is the development of gill slits in the walls of the endodermal pharynx and the division of the pharynx into two zones – the dorsal branchial zone, and ventral digestive zone. During the evolution of the chordates, the sides of the body were inverted. As a result, that part of the digestive tube, which is homologous to the hemichordate digestive zone of the pharynx, became the dorsal notochord and the branchial zone of the pharynx turned out to be ventral. Thus, the division of Triploblastica into two clades entirely retains its significance in modern biology, although it is based on a different set of morphological features than that considered by K. Grobben.

BilateriaTriploblasticaамфистомияэктодермальная глоткаэнтодермальная глоткажаберные щелиэволюцияфилогенияBilateriaTriploblasticaamphystomyectodermal pharynxendodermal pharynxgill slitsevolutionphylogenyРабота выполнена при финансовой поддержке Российского научного фонда (проект № 23-14- 00047) и проведена без использования животных и без привлечения людей в качестве испытуемых.This study was supported by the Russian Science Foundation, project no. 23-14-00047.ReferencesGrobben K. Die systematische Einteilung des Tier­reiches. Verh. kaiser.-könig. zool.-bot. Ges. Wien. 1908;58(10):491–511.Grobben K. Die systematische Einteilung des Tier­reiches. Verh. kaiser.-könig. zool.-bot. Ges. Wien. 1908;58(10):491–511.Malakhov V.V. A new system of Bilateria. Her. Russ. Acad. Sci. 2010;80(3):29–41. DOI: 10.1134/S1019331610010041Malakhov V.V. A new system of Bilateria. Her. Russ. Acad. Sci. 2010;80(3):29–41. DOI: 10.1134/S1019331610010041Malakhov V.V. A revolution in zoology: New concepts of the metazoan system and phylogeny. Her. Russ. Acad. Sci. 2013;83(2):123–127. DOI: 10.1134/S1019331613020044Malakhov V.V. A revolution in zoology: New concepts of the metazoan system and phylogeny. Her. Russ. Acad. Sci. 2013;83(2):123–127. DOI: 10.1134/S1019331613020044Halanych K.M., Bacheller J.D., Aguinaldo A.M., Liva S.M., Hillis D.M., Lake J.A. Evidence from 18S ribosomal DNA that the lophophorates are protostome animals. Science. 1995;267(5204):1641–1643. DOI: 10.1126/science.7886451Halanych K.M., Bacheller J.D., Aguinaldo A.M., Liva S.M., Hillis D.M., Lake J.A. Evidence from 18S ribosomal DNA that the lophophorates are protostome animals. Science. 1995;267(5204):1641–1643. DOI: 10.1126/science.7886451Halanych K.M., Dahlgren T.G., McHugh D. Unseg­mented annelids? Possible origins of four lophotrochozoan worm taxa. Integr. Comp. Biol. 2002;42(3):678–684. DOI: 10.1093/icb/42.3.678Halanych K.M., Dahlgren T.G., McHugh D. Unseg­mented annelids? Possible origins of four lophotrochozoan worm taxa. Integr. Comp. Biol. 2002;42(3):678–684. DOI: 10.1093/icb/42.3.678Aguinaldo A.M., Turbeville J.M., Linford L.S., Rive­ra M.C., Garey J.R., Raff R.A., Lake J.A. Evidence for a clade of nematodes, arthropods and other moulting animals. Nature. 1997;387(6632):489–493. DOI: 10.1038/387489a0Aguinaldo A.M., Turbeville J.M., Linford L.S., Rive­ra M.C., Garey J.R., Raff R.A., Lake J.A. Evidence for a clade of nematodes, arthropods and other moulting animals. Nature. 1997;387(6632):489–493. DOI: 10.1038/387489a0de Rosa R., Grenier J.K., Andreeva T., Cook C.E., Adoutte A., Akam M., Carroll S.B., Balavoine G. Hox genes in brachiopods and priapulids and protostome evolution. Nature. 1999;399(6738):772–776. DOI: 10.1038/21631de Rosa R., Grenier J.K., Andreeva T., Cook C.E., Adoutte A., Akam M., Carroll S.B., Balavoine G. Hox genes in brachiopods and priapulids and protostome evolution. Nature. 1999;399(6738):772–776. DOI: 10.1038/21631Halanych K.M. The new view of animal phylogeny. Annu. Rev. Ecol. Evol. Syst. 2004;35:229–256. DOI: 10.1146/annurev.ecolsys.35.112202.130124Halanych K.M. The new view of animal phylogeny. Annu. Rev. Ecol. Evol. Syst. 2004;35:229–256. DOI: 10.1146/annurev.ecolsys.35.112202.130124Dunn C.W., Hejnol A., Matus D.Q. et al. Broad phy­logenomic sampling improves resolution of the animal tree of life. Nature. 2008;452(7178):745–750. DOI: 10.1038/nature06614Dunn C.W., Hejnol A., Matus D.Q. et al. Broad phy­logenomic sampling improves resolution of the animal tree of life. Nature. 2008;452(7178):745–750. DOI: 10.1038/nature06614Dunn C.W., Giribet G., Edgecombe G.D., Hejnol A. Animal phylogeny and its evolutionary implications. Annu. Rev. Ecol. Evol. Syst. 2014;45:71–395. DOI: 10.1146/annurevecolsys-120213-091627Dunn C.W., Giribet G., Edgecombe G.D., Hejnol A. Animal phylogeny and its evolutionary implications. Annu. Rev. Ecol. Evol. Syst. 2014;45:71–395. DOI: 10.1146/annurevecolsys-120213-091627Giribet G. New animal phylogeny: future challenges for animal phylogeny in the age of phylogenomics. Annu. Rev. Ecol. Evol. Syst. 2015;45:371–395. DOI: 10.1007/s13127-015-0236-4Giribet G. New animal phylogeny: future challenges for animal phylogeny in the age of phylogenomics. Annu. Rev. Ecol. Evol. Syst. 2015;45:371–395. DOI: 10.1007/s13127-015-0236-4Telford M., Budd G., Philippe H. Phylogenomic in­sights into animal evolution. Curr. Biol. 2015;25(19):876–887. DOI: 10.1016/j.cub.2015.07.060Telford M., Budd G., Philippe H. Phylogenomic in­sights into animal evolution. Curr. Biol. 2015;25(19):876–887. DOI: 10.1016/j.cub.2015.07.060Laumer C.E., Fernández R., Lemer S., Com­bosch D., Kocot K.M., Riesgo A., Andrade S.C.S., Ster­rer W., Sørensen M.V., Giribet G. Revisiting metazoan phy­logeny with genomic sampling of all phyla. Proc. Biol. Sci. 2019;286(1906):20190831. DOI: 10.1098/rspb.2019.0831Laumer C.E., Fernández R., Lemer S., Com­bosch D., Kocot K.M., Riesgo A., Andrade S.C.S., Ster­rer W., Sørensen M.V., Giribet G. Revisiting metazoan phy­logeny with genomic sampling of all phyla. Proc. Biol. Sci. 2019;286(1906):20190831. DOI: 10.1098/rspb.2019.0831Малахов В.В. Проблемы построения общей системы многоклеточных. Современная эволюционная морфология. Под ред. Э.И. Воробьевой. Киев: Наукова думка; 1991:195–213.Малахов В.В. Проблемы построения общей системы многоклеточных. Современная эволюционная морфология. Под ред. Э.И. Воробьевой. Киев: Наукова думка; 1991:195–213.Hejnol A., Martindale M.Q. The mouth, the anus, and the blastopore – open questions about questionable openings. Ani­mal Evolution: Genomes, Fossils, and Trees. Eds. Telford M. J., and Littlewood D.T.J. Oxford: Oxford Academic; 2009:33–40.Hejnol A., Martindale M.Q. The mouth, the anus, and the blastopore – open questions about questionable openings. Ani­mal Evolution: Genomes, Fossils, and Trees. Eds. Telford M. J., and Littlewood D.T.J. Oxford: Oxford Academic; 2009:33–40.Nielsen C. How to make a protostome. Invertebrate Systematics. 2012;26(1):25–40. DOI: 10.1071/IS11041Nielsen C. How to make a protostome. Invertebrate Systematics. 2012;26(1):25–40. DOI: 10.1071/IS11041Nielsen C. Evolution of deuterostomy and origin of the chordates. Biol. Rev. Camb. Philos. Soc. 2015;92(1):316–325. DOI: 10.1111/brv.12229Nielsen C. Evolution of deuterostomy and origin of the chordates. Biol. Rev. Camb. Philos. Soc. 2015;92(1):316–325. DOI: 10.1111/brv.12229Nielsen C., Brunet T., Arendt D. Evolution of the bila­terian mouth and anus. Nat. Ecol. Evol. 2018;2(9):1358–1376. DOI: 10.1038/s41559-018-0641-0Nielsen C., Brunet T., Arendt D. Evolution of the bila­terian mouth and anus. Nat. Ecol. Evol. 2018;2(9):1358–1376. DOI: 10.1038/s41559-018-0641-0Nielsen C. Blastopore Fate: Amphistomy, Protosto­my or Deuterostomy. eLS. Chichester: John Wiley & Sons Ltd.; 2019:1–8. DOI: 10.1002/9780470015902.a0027481Nielsen C. Blastopore Fate: Amphistomy, Protosto­my or Deuterostomy. eLS. Chichester: John Wiley & Sons Ltd.; 2019:1–8. DOI: 10.1002/9780470015902.a0027481Ivanova-Kazas O.M. The secondary mouth and its phylogenetic significance. Russ. J. Mar. Biol. 2015;41(1):83–93. DOI: 10.1134/S1063074015020054.Ivanova-Kazas O.M. The secondary mouth and its phylogenetic significance. Russ. J. Mar. Biol. 2015;41(1):83–93. DOI: 10.1134/S1063074015020054.Martín-Durán J.M., Passamaneck Y.J., Martin­dale M.Q., Hejnol A. The developmental basis for the recur­rent evolution of deuterostomy and protostomy. Nat. Ecol. Evol. 2016;1(1):5. DOI: 10.1038/s41559-016-0005Martín-Durán J.M., Passamaneck Y.J., Martin­dale M.Q., Hejnol A. The developmental basis for the recur­rent evolution of deuterostomy and protostomy. Nat. Ecol. Evol. 2016;1(1):5. DOI: 10.1038/s41559-016-0005Tagawa K., Nishino A., Humphreys T., Satoh N. The spawning and early development of the Hawaiian acorn worm (Hemichordate), Ptychodera flava. Zool. Sci. 1998;15(1):85–91. DOI: 10.2108/zsj.15.85Tagawa K., Nishino A., Humphreys T., Satoh N. The spawning and early development of the Hawaiian acorn worm (Hemichordate), Ptychodera flava. Zool. Sci. 1998;15(1):85–91. DOI: 10.2108/zsj.15.85Miyamoto N., Saito Y. Morphology and develop­ment of a new species of Balanoglossus (Hemichordata: En­teropneusta: Ptychoderidae) from Shimoda. Zool. Sci. 2007;24(12):1278–1285. DOI: 10.2108/zsj.24.1278Miyamoto N., Saito Y. Morphology and develop­ment of a new species of Balanoglossus (Hemichordata: En­teropneusta: Ptychoderidae) from Shimoda. Zool. Sci. 2007;24(12):1278–1285. DOI: 10.2108/zsj.24.1278Akesson B. The embryology of the polychaete Eu­nice kobiensis. Acta Zoologica. 1967;48(1–2):141–192. DOI: 10.1111/j.1463-6395.1967.tb00136.xAkesson B. The embryology of the polychaete Eu­nice kobiensis. Acta Zoologica. 1967;48(1–2):141–192. DOI: 10.1111/j.1463-6395.1967.tb00136.xDautert E. Die Bildung der Kaimblatter bei Paludi­na. Zool. Jb. Abt. Anat. 1926;50(4):433–496.Dautert E. Die Bildung der Kaimblatter bei Paludi­na. Zool. Jb. Abt. Anat. 1926;50(4):433–496.Zilch R. Embryologische Untersuchungen an der holoblastischen Ontogenese von Penaeus trisulcatus Leach (Crustacea, Decapoda). Zoomorphologie. 1978;90(1):67–100.Zilch R. Embryologische Untersuchungen an der holoblastischen Ontogenese von Penaeus trisulcatus Leach (Crustacea, Decapoda). Zoomorphologie. 1978;90(1):67–100.Martín-Durán J.M., Janssen R., Wennberg S., Budd G.E., Hejnol A. Deuterostomic development in the pro­tostome Priapulus caudatus. Curr. Biol. 2012;22(22):2161–2166. DOI: 10.1016/j.cub.2012.09.037Martín-Durán J.M., Janssen R., Wennberg S., Budd G.E., Hejnol A. Deuterostomic development in the pro­tostome Priapulus caudatus. Curr. Biol. 2012;22(22):2161–2166. DOI: 10.1016/j.cub.2012.09.037Малахов В.В., Спиридонов С.Э. Эмбриональ­ное развитие Gordius sp. из Туркмении с обсуждением положения волосатиков в системе животного царства. Зоол. журн. 1984;63(9):1285–1296.Малахов В.В., Спиридонов С.Э. Эмбриональ­ное развитие Gordius sp. из Туркмении с обсуждением положения волосатиков в системе животного царства. Зоол. журн. 1984;63(9):1285–1296.Balfour F.M. Anatomy and development of Peri­patopsis capensis. Quart. J. Micr. Soc. 1883;23(2):213–259.Balfour F.M. Anatomy and development of Peri­patopsis capensis. Quart. J. Micr. Soc. 1883;23(2):213–259.Sedgwick A. The development of Peripatus capensis. Part I. Quart. J. Micr. Soc. 1885;25(4):449–466.Sedgwick A. The development of Peripatus capensis. Part I. Quart. J. Micr. Soc. 1885;25(4):449–466.Malakhov V.V. Nematodes. Structure, Development, Classification, Phylogeny. Washington and London: Smithso­nian Institution Press; 1994. 286 pp.Malakhov V.V. Nematodes. Structure, Development, Classification, Phylogeny. Washington and London: Smithso­nian Institution Press; 1994. 286 pp.Malakhov V.V. Embryological and histological pe­culiarities of the order Enoplida, a primitive group of nema­todes. Russ. J. Nematol. 1998;6(1):41–46.Malakhov V.V. Embryological and histological pe­culiarities of the order Enoplida, a primitive group of nema­todes. Russ. J. Nematol. 1998;6(1):41–46.de Beauchamp P.M. La development des Gastro­triches. Bull. Soc. Zool. France. 1929;54(5):549–558.de Beauchamp P.M. La development des Gastro­triches. Bull. Soc. Zool. France. 1929;54(5):549–558.Teuchert G. Zur Fortpflanzung und Entwicklung der Macrodasyoidea (Gastrotricha). Z. Morphol. Tiere. 1968;63(2):343–418.Teuchert G. Zur Fortpflanzung und Entwicklung der Macrodasyoidea (Gastrotricha). Z. Morphol. Tiere. 1968;63(2):343–418.Treadwell A.L. The cytogeny of Podarke obscura. J. Morphol. 1901;17(2):399–476.Treadwell A.L. The cytogeny of Podarke obscura. J. Morphol. 1901;17(2):399–476.Woltereck R. Trochophorastudien. 1. Ueber die En­twicklung des Annelids bei den Polygordiusarten der Nord­see. Bibliotheca Zoologica (Stuttgart). 1902;34:1–71.Woltereck R. Trochophorastudien. 1. Ueber die En­twicklung des Annelids bei den Polygordiusarten der Nord­see. Bibliotheca Zoologica (Stuttgart). 1902;34:1–71.Shearer C. The development of the trochophore of Hydroides (Eupomatus) uncinatus. Quart. J. Microsc. Sci. 1911;56(2):543–590.Shearer C. The development of the trochophore of Hydroides (Eupomatus) uncinatus. Quart. J. Microsc. Sci. 1911;56(2):543–590.Akesson В. The embryology of Tomopteris helgo­landicus (Polychaeta). Acta Zoologica. 1962;48(1):135–199.Akesson В. The embryology of Tomopteris helgo­landicus (Polychaeta). Acta Zoologica. 1962;48(1):135–199.Adrianov A.V., Mayorova A.S., Malakhov V.V. Embry­onic development of the peanut worm Themiste pyroides (Sipun-cula: Sipunculoidea) from the Sea of Japan. Invertebr. Reprod. Dev. 2008;52(1):143–151. DOI: 10.1080/07924259.2008.9652282.Adrianov A.V., Mayorova A.S., Malakhov V.V. Embry­onic development of the peanut worm Themiste pyroides (Sipun-cula: Sipunculoidea) from the Sea of Japan. Invertebr. Reprod. Dev. 2008;52(1):143–151. DOI: 10.1080/07924259.2008.9652282.Patten W. The embryology of Patella. Arb. Zool. Inst. Univ. Wien und Zool. Stat. Triest. 1886;6(2):149–174.Patten W. The embryology of Patella. Arb. Zool. Inst. Univ. Wien und Zool. Stat. Triest. 1886;6(2):149–174.Conklin E.G. The embryology of Crepidula. J. Mor­phol. 1897;13(1):1–226.Conklin E.G. The embryology of Crepidula. J. Mor­phol. 1897;13(1):1–226.Heath H. The development of Ischnochiton. Zool. Jb. Abt. Anat. 1898;12(2):567–565.Heath H. The development of Ischnochiton. Zool. Jb. Abt. Anat. 1898;12(2):567–565.Малахов В.В., Медведева Л.А. Эмбриональное развитие двустворчатого моллюска Mytilus edulis (Mytili­da, Mytilidae). Зоол. журн. 1985;64(12):1808–1815.Малахов В.В., Медведева Л.А. Эмбриональное развитие двустворчатого моллюска Mytilus edulis (Mytili­da, Mytilidae). Зоол. журн. 1985;64(12):1808–1815.Lartillot N., Lespinet O., Vervoort M., Adoutte A. Expression pattern of Brachyury in the mollusk Patella vul­gate suggests a conserved role in the establishment of the AP axis in Bilateria. Development. 2002;129(6):1411–1421. DOI: 10.1242/dev.129.6.1411Lartillot N., Lespinet O., Vervoort M., Adoutte A. Expression pattern of Brachyury in the mollusk Patella vul­gate suggests a conserved role in the establishment of the AP axis in Bilateria. Development. 2002;129(6):1411–1421. DOI: 10.1242/dev.129.6.1411Lyons D.C., Perry K.J., Henry J.Q. Spiralian gas­trulation: germ layer formation, morphogenesis, and fate of the blastopore in the slipper snail Crepidula fornicate. EvoDe­vo. 2015;6:24. DOI:10.1186/s13227-015-0019-1Lyons D.C., Perry K.J., Henry J.Q. Spiralian gas­trulation: germ layer formation, morphogenesis, and fate of the blastopore in the slipper snail Crepidula fornicate. EvoDe­vo. 2015;6:24. DOI:10.1186/s13227-015-0019-1Rattenbury J. C. The embryology of Phoronopsis viridis. J. Morph. 1954;95(2):289–349.Rattenbury J. C. The embryology of Phoronopsis viridis. J. Morph. 1954;95(2):289–349.Temereva E.N., Malakhov V.V. Embryogenesis and larval development of Phoronopsis harmeri Pixell, 1912 (Phoro­nida): dual origin of the coelomic mesoderm. Invertebr. Reprod. Dev. 2007;50(2):57–66. DOI: 10.1080/07924259.2007.9652228.Temereva E.N., Malakhov V.V. Embryogenesis and larval development of Phoronopsis harmeri Pixell, 1912 (Phoro­nida): dual origin of the coelomic mesoderm. Invertebr. Reprod. Dev. 2007;50(2):57–66. DOI: 10.1080/07924259.2007.9652228.Marcus E. Briozoários marinhos brasileiros. 3. Bo­letins da Faculdade de Filosofia, Ciências e Letras. Univ. San Paulo. Zool. 1939;3(1):111–299.Marcus E. Briozoários marinhos brasileiros. 3. Bo­letins da Faculdade de Filosofia, Ciências e Letras. Univ. San Paulo. Zool. 1939;3(1):111–299.Малахов В.В. Описание развития Ascopodaria discreta (Coloniales, Barentsiidae) и обсуждение положе­ния Kamptozoa в системе животного царства. Зоол. журн. 1990;69(10):20–30.Малахов В.В. Описание развития Ascopodaria discreta (Coloniales, Barentsiidae) и обсуждение положе­ния Kamptozoa в системе животного царства. Зоол. журн. 1990;69(10):20–30.Conklin E.G. The embryology of a brachiopod, Terebratulina septentrionalis Couthouy. Proc. Am. Philos. Soc. 1902;41(1):41–76.Conklin E.G. The embryology of a brachiopod, Terebratulina septentrionalis Couthouy. Proc. Am. Philos. Soc. 1902;41(1):41–76.Martín-Durán J., Passamaneck Y., Martindale M., Hejnol A. The developmental basis for the recurrent evolu­tion of deuterostomy and protostomy. Nat. Ecol. Evol. 2016;1(1):5. DOI: 10.1038/s41559-016-0005Martín-Durán J., Passamaneck Y., Martindale M., Hejnol A. The developmental basis for the recurrent evolu­tion of deuterostomy and protostomy. Nat. Ecol. Evol. 2016;1(1):5. DOI: 10.1038/s41559-016-0005Ковалевский А.О. Наблюдения над развитием Вгасhiороdа. Изв. Имп. О-ва любит. естествозн., антро­пол. этногр. 1874;14(1):1–40.Ковалевский А.О. Наблюдения над развитием Вгасhiороdа. Изв. Имп. О-ва любит. естествозн., антро­пол. этногр. 1874;14(1):1–40.Nielsen C. The development of the brachiopod Cra­nia (Neocrania) anomala (O.F. Müller) and its phylogenetic significance. Acta Zool. 1991;72(1):7–28. DOI: 10.1111/j.1463-6395.1991.tb00312.xNielsen C. The development of the brachiopod Cra­nia (Neocrania) anomala (O.F. Müller) and its phylogenetic significance. Acta Zool. 1991;72(1):7–28. DOI: 10.1111/j.1463-6395.1991.tb00312.xVellutini B.C., Hejnol A. Expression of segment po­larity genes in brachiopods supports a non-segmental ances­tral role of engrailed for bilaterians. Sci. Rep. 2016;6:32387. DOI: 10.1038/srep32387Vellutini B.C., Hejnol A. Expression of segment po­larity genes in brachiopods supports a non-segmental ances­tral role of engrailed for bilaterians. Sci. Rep. 2016;6:32387. DOI: 10.1038/srep32387Heymons R. Die Entwicklungsgeschichte der Scol­opender. Bibliotheca Zoologica (Stuttgart). 1901;13:1–224.Heymons R. Die Entwicklungsgeschichte der Scol­opender. Bibliotheca Zoologica (Stuttgart). 1901;13:1–224.Иванов П.П. Эмбриональное развитие сколо­пендры в связи с эмбриологией и морфологией Trachea­ta. Изв. АН СССР. Отдел. биолог. наук. 1940;6:831–861.Иванов П.П. Эмбриональное развитие сколо­пендры в связи с эмбриологией и морфологией Trachea­ta. Изв. АН СССР. Отдел. биолог. наук. 1940;6:831–861.Wheeler W.M. A contribution to insect embryology. J. Morphol. 1893;8(1):1–161.Wheeler W.M. A contribution to insect embryology. J. Morphol. 1893;8(1):1–161.Jura C. Embryogenesis of the alimentary system of the weevil, Phyllobius glaucus. Zool. Poloniae. 1956;7(1):155–176.Jura C. Embryogenesis of the alimentary system of the weevil, Phyllobius glaucus. Zool. Poloniae. 1956;7(1):155–176.Brauer A. Entwicklungsgeschichte des Scorpions. Z. wiss. Zool. 1894;57(2):409–432.Brauer A. Entwicklungsgeschichte des Scorpions. Z. wiss. Zool. 1894;57(2):409–432.Weygoldt P. Embryologische Untersuchungen an Ostracoden: Die Entwicklung von Cyprides litoralis. Zool. Jb. Abt. Anat. 1960;78(2):370–426.Weygoldt P. Embryologische Untersuchungen an Ostracoden: Die Entwicklung von Cyprides litoralis. Zool. Jb. Abt. Anat. 1960;78(2):370–426.Nielsen C., Hay-Schmidt A. Development of the enteropneust Ptychodera flava: ciliary bands and nervous system. J. Morphol. 2007;268(7):551–570. DOI: 10.1002/jmor.10533Nielsen C., Hay-Schmidt A. Development of the enteropneust Ptychodera flava: ciliary bands and nervous system. J. Morphol. 2007;268(7):551–570. DOI: 10.1002/jmor.10533Малахов В.В. Проблема основного плана строе­ния в различных группах вторичноротых животных. Журн. общ. биологии. 1977;38(4):485–499.Малахов В.В. Проблема основного плана строе­ния в различных группах вторичноротых животных. Журн. общ. биологии. 1977;38(4):485–499.Arendt D., Nübler-Jung K. Inversion of dorsoven­tral axis. Nature. 1994;371(6492):26. DOI: 10.1038/371026a0Arendt D., Nübler-Jung K. Inversion of dorsoven­tral axis. Nature. 1994;371(6492):26. DOI: 10.1038/371026a0Arendt D., Nübler-Jung K. Dorsal or ventral: simi­larities in fate maps and gastrulation patterns in annelids, ar­thropods and chordates. Mech. Dev. 1997;61(1–2):7–21. DOI: 10.1016/s0925-4773(96)00620-xArendt D., Nübler-Jung K. Dorsal or ventral: simi­larities in fate maps and gastrulation patterns in annelids, ar­thropods and chordates. Mech. Dev. 1997;61(1–2):7–21. DOI: 10.1016/s0925-4773(96)00620-xMacBride E. W. The formation of the layers in am­phioxus and its bearing on the interpretation of the early on­togenetic processes in other vertebrates. Quart. J. Microsc. Sci. 1909;54(3):279–345.MacBride E. W. The formation of the layers in am­phioxus and its bearing on the interpretation of the early on­togenetic processes in other vertebrates. Quart. J. Microsc. Sci. 1909;54(3):279–345.Bergström J. Origin of high-rank groups of organ­isms. Paleontol. Res. 1997;1(1):1–14.Bergström J. Origin of high-rank groups of organ­isms. Paleontol. Res. 1997;1(1):1–14.Nübler-Jung K.N., Arendt D. Dorsoventral axis in­version. Enteropneust anatomy links invertebrates to chor­dates turned upside down. J. Zool. Syst. Evol. Res. 1999;37(2):93–100.Nübler-Jung K.N., Arendt D. Dorsoventral axis in­version. Enteropneust anatomy links invertebrates to chor­dates turned upside down. J. Zool. Syst. Evol. Res. 1999;37(2):93–100.Lacalli T.C. The emergence of the chordate body plan: some puzzles and problems. Acta Zoologica (Stockholm). 2010;91(1):4–10. DOI: 10.1111/j.1463-6395.2008.00384.xLacalli T.C. The emergence of the chordate body plan: some puzzles and problems. Acta Zoologica (Stockholm). 2010;91(1):4–10. DOI: 10.1111/j.1463-6395.2008.00384.xVeeman M.T., Newman-Smith E., El-Nachef D., Smith W.C. The ascidian mouth opening is derived from the anterior neuropore: reassessing the mouth/neural tube relation­ship in chordate evolution. Dev. Biol. 2010;344(1):138–149. DOI: 10.1016/j.ydbio.2010.04.028Veeman M.T., Newman-Smith E., El-Nachef D., Smith W.C. The ascidian mouth opening is derived from the anterior neuropore: reassessing the mouth/neural tube relation­ship in chordate evolution. Dev. Biol. 2010;344(1):138–149. DOI: 10.1016/j.ydbio.2010.04.028Максимов А.А. Основы гистологии. Петроград: К.Л. Риккер; 1914. 382 с.Максимов А.А. Основы гистологии. Петроград: К.Л. Риккер; 1914. 382 с.Jiménez A.J., Domínguez-Pinos M.D., Guer­ra M.M., Fernández-Llebrez P., Pérez-Fígares J.M. Struc­ture and function of the ependymal barrier and diseases as­sociated with ependyma disruption. Tissue Barriers. 2014;2:e28426. DOI: 10.4161/tisb.28426Jiménez A.J., Domínguez-Pinos M.D., Guer­ra M.M., Fernández-Llebrez P., Pérez-Fígares J.M. Struc­ture and function of the ependymal barrier and diseases as­sociated with ependyma disruption. Tissue Barriers. 2014;2:e28426. DOI: 10.4161/tisb.28426Denes A.S., Jékely G., Steinmetz P.R., Raible F., Sny­man H., Prud’homme B., Ferrier D.E., Balavoine G., Arendt D. Molecular architecture of annelid nerve cord supports com­mon origin of nervous system centralization in bilateria. Cell. 2007;129(2):277–288. DOI: 10.1016/j.cell.2007.02.040Denes A.S., Jékely G., Steinmetz P.R., Raible F., Sny­man H., Prud’homme B., Ferrier D.E., Balavoine G., Arendt D. Molecular architecture of annelid nerve cord supports com­mon origin of nervous system centralization in bilateria. Cell. 2007;129(2):277–288. DOI: 10.1016/j.cell.2007.02.040Arenas-Mena C. Brachyury, Tbx2/3 and sall expres­sion during embryogenesis of the indirectly developing poly­chaete Hydroides elegans. Int. J. Dev. Biol. 2013;57(1):73–83. DOI: 10.1387/ijdb.120056caArenas-Mena C. Brachyury, Tbx2/3 and sall expres­sion during embryogenesis of the indirectly developing poly­chaete Hydroides elegans. Int. J. Dev. Biol. 2013;57(1):73–83. DOI: 10.1387/ijdb.120056caLauri A., Brunet T., Handberg-Thorsager M., Fis­cher A.H., Simakov O., Steinmetz P.R., Tomer R., Keller P.J., Arendt D. Development of the annelid axochord: insights into notochord evolution. Science. 2014;345(6202):1365–1368. DOI: 10.1126/science.1253396Lauri A., Brunet T., Handberg-Thorsager M., Fis­cher A.H., Simakov O., Steinmetz P.R., Tomer R., Keller P.J., Arendt D. Development of the annelid axochord: insights into notochord evolution. Science. 2014;345(6202):1365–1368. DOI: 10.1126/science.1253396Boyle M.J., Yamaguchi E., Seaver E.C. Molecular conservation of metazoan gut formation: evidence from ex­pression of endomesoderm genes in Capitella teleta (Anneli­da). EvoDevo. 2014;5:39. DOI: 10.1186/2041-9139-5-39Boyle M.J., Yamaguchi E., Seaver E.C. Molecular conservation of metazoan gut formation: evidence from ex­pression of endomesoderm genes in Capitella teleta (Anneli­da). EvoDevo. 2014;5:39. DOI: 10.1186/2041-9139-5-39Martín-Durán J. M., Hejnol A. The study of Pria­pulus caudatus reveals conserved molecular patterning un­derlying different gut morphogenesis in the Ecdysozoa. BMC Biol. 2015;13:29. DOI: 10.1186/s12915-015-0139-zMartín-Durán J. M., Hejnol A. The study of Pria­pulus caudatus reveals conserved molecular patterning un­derlying different gut morphogenesis in the Ecdysozoa. BMC Biol. 2015;13:29. DOI: 10.1186/s12915-015-0139-zArendt D., Technau U., Wittbrodt J. Evolution of the bilaterian larval foregut. Nature. 2001;409(6816):81–85. DOI: 10.1038/35051075Arendt D., Technau U., Wittbrodt J. Evolution of the bilaterian larval foregut. Nature. 2001;409(6816):81–85. DOI: 10.1038/35051075Tagawa K., Humphreys T., Satoh N. Novel pattern of Brachyury gene expression in hemichordate embryos. Mech. Dev. 1998;75(1–2):139–143. DOI: 10.1016/s0925-4773(98)00078-1Tagawa K., Humphreys T., Satoh N. Novel pattern of Brachyury gene expression in hemichordate embryos. Mech. Dev. 1998;75(1–2):139–143. DOI: 10.1016/s0925-4773(98)00078-1Shoguchi E., Satoh N., Maruyama Y.K. Pattern of Brachyury gene expression in starfish embryos resembles that of hemichordate embryos but not of sea urchin embryos. Mech. Dev. 1999;82(1–2):185–189. DOI: 10.1016/s0925-4773(99)00008-8Shoguchi E., Satoh N., Maruyama Y.K. Pattern of Brachyury gene expression in starfish embryos resembles that of hemichordate embryos but not of sea urchin embryos. Mech. Dev. 1999;82(1–2):185–189. DOI: 10.1016/s0925-4773(99)00008-8Davidson A.J., Zon L.I. The caudal-related homeo­box genes cdx1a and cdx4 act redundantly to regulate hox gene expression and the formation of putative hematopoietic stem cells during zebrafish embryogenesis. Dev. Biol. 2006;292(2):506–518. DOI: 10.1016/j.ydbio.2006.01.003Davidson A.J., Zon L.I. The caudal-related homeo­box genes cdx1a and cdx4 act redundantly to regulate hox gene expression and the formation of putative hematopoietic stem cells during zebrafish embryogenesis. Dev. Biol. 2006;292(2):506–518. DOI: 10.1016/j.ydbio.2006.01.003Mudumana S.P., Hentschel D., Liu Y., Vasilyev A., Drummond I.A. odd skipped related1 reveals a novel role for endoderm in regulating kidney versus vascular cell fate. Deve-lopment. 2008;135(20):3355–3367. DOI: 10.1242/dev.022830Mudumana S.P., Hentschel D., Liu Y., Vasilyev A., Drummond I.A. odd skipped related1 reveals a novel role for endoderm in regulating kidney versus vascular cell fate. Deve-lopment. 2008;135(20):3355–3367. DOI: 10.1242/dev.022830Hejnol A., Martindale M. Acoel development indi­cates the independent evolution of the bilaterian mouth and anus. Nature. 2008;456(7220):382–386. DOI: 10.1038/nature07309Hejnol A., Martindale M. Acoel development indi­cates the independent evolution of the bilaterian mouth and anus. Nature. 2008;456(7220):382–386. DOI: 10.1038/nature07309Andrikou C., Pang K., Børve A., Lu T-M., He­jnol A. Molecular evidence from xenacoelomorph gonopore formation supports homology with the bilaterian anus. Nat. Ecol. Evol. 2025;9:2116–2126. DOI: 10.1038/s41559-025-02866-6Andrikou C., Pang K., Børve A., Lu T-M., He­jnol A. Molecular evidence from xenacoelomorph gonopore formation supports homology with the bilaterian anus. Nat. Ecol. Evol. 2025;9:2116–2126. DOI: 10.1038/s41559-025-02866-6Marlétaz F., Peijnenburg K.T.C.A., Goto T., Sa­toh N., Rokhsar D.S. A new spiralian phylogeny places the enigmatic arrow worms among gnathiferans. Curr. Biol. 2019;29(2):312–318. DOI: 10.1016/j.cub.2018.11.042Marlétaz F., Peijnenburg K.T.C.A., Goto T., Sa­toh N., Rokhsar D.S. A new spiralian phylogeny places the enigmatic arrow worms among gnathiferans. Curr. Biol. 2019;29(2):312–318. DOI: 10.1016/j.cub.2018.11.042Piovani L., Gavriouchkina D., Parey E., Sarre L.A., Peijnenburg K.T.C.A., Martín-Durán J.M., Rokhsar D.S., Satoh N., de Mendoza A., Goto T., Marlétaz F. The geno-mic origin of the unique chaetognath body plan. Nature. 2025;645(8082):964–973. DOI: 10.1038/s41586-025-09403- 2. DOI: 10.1038/s41586-025-09403-2Piovani L., Gavriouchkina D., Parey E., Sarre L.A., Peijnenburg K.T.C.A., Martín-Durán J.M., Rokhsar D.S., Satoh N., de Mendoza A., Goto T., Marlétaz F. The geno-mic origin of the unique chaetognath body plan. Nature. 2025;645(8082):964–973. DOI: 10.1038/s41586-025-09403- 2. DOI: 10.1038/s41586-025-09403-2Lankaster E.R. On the use of the term homology in modern zoology, and the distinction between homogenetic and homoplastic agreements. Ann. Mag. Nat. Hist. 1870;6(31):34–43.Lankaster E.R. On the use of the term homology in modern zoology, and the distinction between homogenetic and homoplastic agreements. Ann. Mag. Nat. Hist. 1870;6(31):34–43.Любищев А.А. Понятие сравнительной анатомии. В кн.: Вопросы общей зоологии и медицинской паразитологии. М.: Биомедгиз; 1962:189–214.Любищев А.А. Понятие сравнительной анатомии. В кн.: Вопросы общей зоологии и медицинской паразитологии. М.: Биомедгиз; 1962:189–214.Beklemishev V.N. Principles of comparative anatomy of invertebrates: V. 1 Promorphology, vol. 2 Organology. Univ. Chicago Press; 1969. 1058 pp.Beklemishev V.N. Principles of comparative anatomy of invertebrates: V. 1 Promorphology, vol. 2 Organology. Univ. Chicago Press; 1969. 1058 pp.Иванов Д.Л., Цетлин А.Б. Возниковение и эво­люция кутикулярного глоточного вооружения у трохо­форных животных с вентральной глоткой. Зоол. журн. 1981;60(11):1445–1454.Иванов Д.Л., Цетлин А.Б. Возниковение и эво­люция кутикулярного глоточного вооружения у трохо­форных животных с вентральной глоткой. Зоол. журн. 1981;60(11):1445–1454.Tzetlin A., Purschke G. Pharynx and intestine. Hy­drobiologia. 2005;535/536(1):199–225. DOI: 10.1007/1-4020-3240-4_12Tzetlin A., Purschke G. Pharynx and intestine. Hy­drobiologia. 2005;535/536(1):199–225. DOI: 10.1007/1-4020-3240-4_12Remane A. Die Grundlagen des natürlichen Systems der vergleichenden Anatomie und der Phylogenetik. Leipzig; 1952. 400 S.Remane A. Die Grundlagen des natürlichen Systems der vergleichenden Anatomie und der Phylogenetik. Leipzig; 1952. 400 S.Spengel J.W. Die Enteropneusten des Golfes von Ne­apel und der angrenzenden Meeres-Abschnitte. Fauna und Flo­ra des Golfes von Neapel und der angrenzenden Meeres-Ab­schnitte. Herausgegeben von der Zoologischen Station zu Neapel. Berlin: Friedländer. 1893;18:1–757.Spengel J.W. Die Enteropneusten des Golfes von Ne­apel und der angrenzenden Meeres-Abschnitte. Fauna und Flo­ra des Golfes von Neapel und der angrenzenden Meeres-Ab­schnitte. Herausgegeben von der Zoologischen Station zu Neapel. Berlin: Friedländer. 1893;18:1–757.van der Horst C.J. Hemichordata. Klassen und Ord­nungen des Tierreichs wissenschaftlich dargestellt in Wort und Bild. Ed. Bronn H.G. Leipzig: Akademische Verlags­gesellschaft; 1939: 1–737.van der Horst C.J. Hemichordata. Klassen und Ord­nungen des Tierreichs wissenschaftlich dargestellt in Wort und Bild. Ed. Bronn H.G. Leipzig: Akademische Verlags­gesellschaft; 1939: 1–737.Barrington E.J.W. Observations on feeding and di­gestion in Glossobalanus minutus. Quart. J. Micr. Sci. 1940;82(2):227–260.Barrington E.J.W. Observations on feeding and di­gestion in Glossobalanus minutus. Quart. J. Micr. Sci. 1940;82(2):227–260.Burdon-Jones C. The feeding mechanism of Bal­anoglossus gigas. Bol. Fac. Filos. Cienc. Letr. Univ. Sao Paulo. Zoologia.1962;24(261):255–280.Burdon-Jones C. The feeding mechanism of Bal­anoglossus gigas. Bol. Fac. Filos. Cienc. Letr. Univ. Sao Paulo. Zoologia.1962;24(261):255–280.Cameron C.B. Particle retention and flow in the phar­ynx of the enteropneust worm Harrimania planktophilus: the fil­ter-feeding pharynx may have evolved before the chordates. Biol. Bull. 2002;202(2):192–200. DOI: 10.2307/1543655Cameron C.B. Particle retention and flow in the phar­ynx of the enteropneust worm Harrimania planktophilus: the fil­ter-feeding pharynx may have evolved before the chordates. Biol. Bull. 2002;202(2):192–200. DOI: 10.2307/1543655Slack J.M.W., Holland P.W.H., Graham C.F. The zootype and the phylotypic stage. Nature. 1993;361(6412):490–492. DOI: 10.1038/361490a0Slack J.M.W., Holland P.W.H., Graham C.F. The zootype and the phylotypic stage. Nature. 1993;361(6412):490–492. DOI: 10.1038/361490a0Ruppert E.E., Cameron C.B., Frick J.E. Endo­style-like features of the dorsal epibranchial ridge of an en­teropneust and the hypothesis of dorso-ventral inversion in chordates. Invert. Biol. 1999;118(2):202–212. DOI: 10.2307/3227061Ruppert E.E., Cameron C.B., Frick J.E. Endo­style-like features of the dorsal epibranchial ridge of an en­teropneust and the hypothesis of dorso-ventral inversion in chordates. Invert. Biol. 1999;118(2):202–212. DOI: 10.2307/3227061Sato T. Vorlaufige mitteilung fiber Atubaria heterolo­pha gen. nov., sp. nov., einen in freiem zustand aufgefunde­nen Pterobranchier aus dem stillen ozean. Zoologische An­zeiger. 1936;15(7):97–106.Sato T. Vorlaufige mitteilung fiber Atubaria heterolo­pha gen. nov., sp. nov., einen in freiem zustand aufgefunde­nen Pterobranchier aus dem stillen ozean. Zoologische An­zeiger. 1936;15(7):97–106.Komai T. Internal structure of the pterobranch Atubaria heterolopha with an appendix on the homology of the ‘notochord’. Proc. Japan Acad. 1949;25(7):19–24.Komai T. Internal structure of the pterobranch Atubaria heterolopha with an appendix on the homology of the ‘notochord’. Proc. Japan Acad. 1949;25(7):19–24.Halanych K. Convergence in the feeding appara­tuses of lophophorates and pterobranch hemichordates re­vealed by 18S rDNA: An interpretation. Biol. Bull. 1996;190(1):1–5. DOI: 10.2307/1542669Halanych K. Convergence in the feeding appara­tuses of lophophorates and pterobranch hemichordates re­vealed by 18S rDNA: An interpretation. Biol. Bull. 1996;190(1):1–5. DOI: 10.2307/1542669Щепотьев А. Pterobranchia. Санкт-Петербург: Типография Стасюлевича; 1907. 248 с.Щепотьев А. Pterobranchia. Санкт-Петербург: Типография Стасюлевича; 1907. 248 с.Lester S.M. Cephalodiscus sp. (Hemichordata: Pterobranchia): observations of functional morphology, be­havior and occurrence in shallow water around Bermuda. Mar. Biol. 1985;85(3):263–268. DOI: 10.1007/BF00393246Lester S.M. Cephalodiscus sp. (Hemichordata: Pterobranchia): observations of functional morphology, be­havior and occurrence in shallow water around Bermuda. Mar. Biol. 1985;85(3):263–268. DOI: 10.1007/BF00393246Maletz J., Cameron C.B. Treatise on invertebrate paleontology, part V, chapter 3: introduction to class Ptero­branchia Lankester, 1877. Treatise Online. 2016;82:1–15.Maletz J., Cameron C.B. Treatise on invertebrate paleontology, part V, chapter 3: introduction to class Ptero­branchia Lankester, 1877. Treatise Online. 2016;82:1–15.Bather-London F.A. Vortrage und diskussionen auf der Wiener Tagung der Palaeontologischen Gesellschaft im September 1923: Cothurnocystis: a study in adaptation. Paläontologische Zeitschrift. 1926;7(1):1–15.Bather-London F.A. Vortrage und diskussionen auf der Wiener Tagung der Palaeontologischen Gesellschaft im September 1923: Cothurnocystis: a study in adaptation. Paläontologische Zeitschrift. 1926;7(1):1–15.Jefferies R.P.S. Some fossil chordates with echino­derm affinities. Symp. Zool. Soc. Lond. 1967;20:163–208.Jefferies R.P.S. Some fossil chordates with echino­derm affinities. Symp. Zool. Soc. Lond. 1967;20:163–208.Jefferies R.P.S. The ancestry of the vertebrates. Bri-tish Museum (Natural History) & Cambridge University Press; 1986. 376 pp.Jefferies R.P.S. The ancestry of the vertebrates. Bri-tish Museum (Natural History) & Cambridge University Press; 1986. 376 pp.Ubaghs G. Stylophora. Treatise on Invertebrate Paleontology. Part S. Echinodermata, vol. 1. Ed. R.C. Moore. Lawrence: Geological Society of America and University of Kansas Press; 1968:495–565.Ubaghs G. Stylophora. Treatise on Invertebrate Paleontology. Part S. Echinodermata, vol. 1. Ed. R.C. Moore. Lawrence: Geological Society of America and University of Kansas Press; 1968:495–565.Ubaghs G. Early Paleozoic echinoderms. Ann. Rev. Earth Planet. Sci. 1975;3(1):79–98.Ubaghs G. Early Paleozoic echinoderms. Ann. Rev. Earth Planet. Sci. 1975;3(1):79–98.Ruta M. Brief review of the stylophoran debate. Evol. Dev. 1999;1(2):123–135. DOI: 10.1046/j.1525-142x.1999.99008.xRuta M. Brief review of the stylophoran debate. Evol. Dev. 1999;1(2):123–135. DOI: 10.1046/j.1525-142x.1999.99008.xKapli P., Natsidis P., Leite D.J., Fursman M., Jef­frie N., Rahman I.A., Philippe H., Copley R.R., Telford M.J. Lack of support for Deuterostomia prompts reinterpretation of the first Bilateria. Sci. Adv. 2021;7(12):eabe2741. DOI: 10.1126/sciadv.abe2741Kapli P., Natsidis P., Leite D.J., Fursman M., Jef­frie N., Rahman I.A., Philippe H., Copley R.R., Telford M.J. Lack of support for Deuterostomia prompts reinterpretation of the first Bilateria. Sci. Adv. 2021;7(12):eabe2741. DOI: 10.1126/sciadv.abe2741Silva A.S., Natsidis P., Piovani L., Kapli P., Tel­ford M.J. Is the deuterostome clade an artifact? Curr. Biol. 2025;35(15):3611–3623. DOI: 10.1016/j.cub.2025.06.045Silva A.S., Natsidis P., Piovani L., Kapli P., Tel­ford M.J. Is the deuterostome clade an artifact? Curr. Biol. 2025;35(15):3611–3623. DOI: 10.1016/j.cub.2025.06.045Cannon J.T., Vellutini B.C., Smith J. 3rd, Ron­quist F., Jondelius U., Hejnol A. Xenacoelomorpha is the sister group to Nephrozoa. Nature. 2016;530(7588):89–93. DOI: 10.1038/nature16520Cannon J.T., Vellutini B.C., Smith J. 3rd, Ron­quist F., Jondelius U., Hejnol A. Xenacoelomorpha is the sister group to Nephrozoa. Nature. 2016;530(7588):89–93. DOI: 10.1038/nature16520Philippe H., Brinkmann H., Copley R.R., Moroz L.L., Nakano H., Poustka A.J., Wallberg A., Peterson K.J., Telford M.J. Acoelomorph flatworms are deuterostomes related to Xenoturbella. Nature. 2011;470(7333):255–258. DOI: 10.1038/nature09676Philippe H., Brinkmann H., Copley R.R., Moroz L.L., Nakano H., Poustka A.J., Wallberg A., Peterson K.J., Telford M.J. Acoelomorph flatworms are deuterostomes related to Xenoturbella. Nature. 2011;470(7333):255–258. DOI: 10.1038/nature09676Philippe H., Poustka A.J. Chiodin M. et al. Miti­gating anticipated effects of systematic errors supports sis­ter-group relationship between Xenacoelomorpha and Am­bulacraria. Curr. Biol. 2019;29(11):1818–1826. DOI: 10.1016/j.cub.2019.04.009Philippe H., Poustka A.J. Chiodin M. et al. Miti­gating anticipated effects of systematic errors supports sis­ter-group relationship between Xenacoelomorpha and Am­bulacraria. Curr. Biol. 2019;29(11):1818–1826. DOI: 10.1016/j.cub.2019.04.009Martynov A., Lundin K., Picton B., Fletcher K., Malmberg K., Korshunova T. Multiple paedomorphic line­ages of soft-substrate burrowing invertebrates: parallels in the origin of Xenocratena and Xenoturbella. PLoS One. 2020;15(1):e0227173. DOI: 10.1371/journal.pone.0227173Martynov A., Lundin K., Picton B., Fletcher K., Malmberg K., Korshunova T. Multiple paedomorphic line­ages of soft-substrate burrowing invertebrates: parallels in the origin of Xenocratena and Xenoturbella. PLoS One. 2020;15(1):e0227173. DOI: 10.1371/journal.pone.0227173Международный кодекс зоологической номенкла­туры. Изд. 4-e. М.: Т-во науч. изд. КМК; 2004. 223 с.Международный кодекс зоологической номенкла­туры. Изд. 4-e. М.: Т-во науч. изд. КМК; 2004. 223 с.Kuhn T.S. The Structure of Scientific Revolutions. University of Chicago Press; 1962. 172 pp.Kuhn T.S. The Structure of Scientific Revolutions. University of Chicago Press; 1962. 172 pp.Гинзбург В.Л. О физике и астрофизике: Ста­тьи и выступления. М.: Наука; 1992. 528 с.Гинзбург В.Л. О физике и астрофизике: Ста­тьи и выступления. М.: Наука; 1992. 528 с.de Beaumarchais P.-A.C. La Folle Journee, ou le Mariage de Figaro. Paris: De l’Imprimerie de la Societe Lit­teraire-Typographique; 1785. 201 pp.de Beaumarchais P.-A.C. La Folle Journee, ou le Mariage de Figaro. Paris: De l’Imprimerie de la Societe Lit­teraire-Typographique; 1785. 201 pp.

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