<?xml version="1.0" encoding="UTF-8"?>
<!DOCTYPE article PUBLIC "-//NLM//DTD JATS (Z39.96) Journal Publishing DTD v1.3 20210610//EN" "JATS-journalpublishing1-3.dtd">
<article article-type="research-article" dtd-version="1.3" xmlns:mml="http://www.w3.org/1998/Math/MathML" xmlns:xlink="http://www.w3.org/1999/xlink" xmlns:xsi="http://www.w3.org/2001/XMLSchema-instance" xml:lang="ru"><front><journal-meta><journal-id journal-id-type="publisher-id">boolt</journal-id><journal-title-group><journal-title xml:lang="ru">Бюллетень Государственного Никитского ботанического сада</journal-title><trans-title-group xml:lang="en"><trans-title>Bulletin of the State Nikitsky Botanical Gardens</trans-title></trans-title-group></journal-title-group><issn pub-type="ppub">0513-1634</issn><publisher><publisher-name>Federal State Funded Institution of Science “The Labour Red Banner Order Nikitsky Botanical Gardens – National scientific Center of the RAS”</publisher-name></publisher></journal-meta><article-meta><article-id custom-type="elpub" pub-id-type="custom">boolt-1105</article-id><article-categories><subj-group subj-group-type="heading"><subject>Research Article</subject></subj-group><subj-group subj-group-type="section-heading" xml:lang="ru"><subject>БОТАНИКА И ЭКОЛОГИЯ</subject></subj-group></article-categories><title-group><article-title>К вопросу о классификации жизненных форм тропических и субтропических видов папоротникообразных растений</article-title><trans-title-group xml:lang="en"><trans-title>The Approaches to the Classification of Life Forms of Fern-Like Plants (Polypodiophyta, Including Lycopodiophyta)</trans-title></trans-title-group></title-group><contrib-group><contrib contrib-type="author" corresp="yes"><name-alternatives><name name-style="eastern" xml:lang="ru"><surname>Калюжный</surname><given-names>С. С.</given-names></name><name name-style="western" xml:lang="en"><surname>Kalyuzhny</surname><given-names>S. S.</given-names></name></name-alternatives><bio xml:lang="en"><p>Sergey Sergeevich Kalyuzhny,</p><p>197022, St. Petersburg, internal territory of the city, municipal district Aptekarsky Island, Professor Popov Street, Bld. 2, litera V.</p><p> </p></bio><email xlink:type="simple">fernbaikal@yandex.ru</email><xref ref-type="aff" rid="aff-1"/></contrib><contrib contrib-type="author" corresp="yes"><name-alternatives><name name-style="eastern" xml:lang="ru"><surname>Шевчук</surname><given-names>О. М.</given-names></name><name name-style="western" xml:lang="en"><surname>Shevchuk</surname><given-names>O. M.</given-names></name></name-alternatives><bio xml:lang="en"><p>Oksana Mikhailovna Shevchuk,</p><p>298648, Yalta, NBG-NSC’s territory, bld. 1, facility 1.</p></bio><email xlink:type="simple">oksana_shevchuk1970@mail.ru</email><xref ref-type="aff" rid="aff-2"/></contrib></contrib-group><aff xml:lang="en" id="aff-1"><institution>Federal State Funded Institution of Science Botanical Institute named after V.L. Komarov of the Russian Academy of Sciences, Peter the Great Botanical Garden</institution><country>Russian Federation</country></aff><aff xml:lang="en" id="aff-2"><institution>Federal State Funded Institution of Science “The Labour Red Banner Order Nikitsky Botanical Gardens – National Scientific Center of the RAS”</institution><country>Russian Federation</country></aff><pub-date pub-type="collection"><year>2026</year></pub-date><pub-date pub-type="epub"><day>23</day><month>05</month><year>2026</year></pub-date><volume>0</volume><issue>158</issue><fpage>44</fpage><lpage>58</lpage><permissions><copyright-statement>Copyright &amp;#x00A9; Калюжный С.С., Шевчук О.М., 2026</copyright-statement><copyright-year>2026</copyright-year><copyright-holder xml:lang="ru">Калюжный С.С., Шевчук О.М.</copyright-holder><copyright-holder xml:lang="en">Kalyuzhny S.S., Shevchuk O.M.</copyright-holder><license xml:lang="ru" license-type="creative-commons-attribution" xlink:href="https://creativecommons.org/licenses/by/4.0/" xlink:type="simple"><license-p>Данная работа распространяется под лицензией Creative Commons Attribution 4.0.</license-p></license><license xml:lang="en" license-type="creative-commons-attribution" xlink:href="https://creativecommons.org/licenses/by/4.0/" xlink:type="simple"><license-p>This work is licensed under a Creative Commons Attribution 4.0 License.</license-p></license></permissions><self-uri xlink:href="https://boolt.elpub.ru/jour/article/view/1105">https://boolt.elpub.ru/jour/article/view/1105</self-uri><abstract><p>Жизненная форма (ЖФ) папоротникообразных растений (Polypodiophyta, включая Lycopodiophyta) является отражением приспособленности к прошлым и существующим почвенноклиматическим и ценотическим условиям местообитаний. Анализ созданных к настоящему времени классификаций папоротникообразных показывает вовлеченность как спорофитов, так и гаметофитов. В частности, исследование морфологических структур гаметофитов папоротников, начатое в XIX веке еще немецкими ботаниками, выявило важность использования сравнительной морфологии при подходах к классификации папоротникообразных растений (Orth, 1936; Goebel, 1928). Различные способы прорастания спор были проанализированы S. Momose и отражены им в многочисленных публикациях (1937 a, b; 1938 a, b, c; 1939; 1941; 1942). Nayar B.K., Kaur S при изучении папоротникообразных растений Индии предложили новую классификацию. Варминг Й.Э. (1884) при выделении жизненных форм папоротников выделял две группы: I главная (3 подгруппы) самая малая и II главная (2 подгруппы, 3 типа) со множеством вариантов и подвариантов. Первые подходы в нашей стране по выделению ЖФ успешно были развиты И.Г. Серебряковым для высших цветковых растений (Серебряков, 1964), где не рассматривались споровые растения. Первая классификация споровых растений (папоротников, хвощей и плаунов) нашей страны нашла отражение в работах А.П. Хохрякова (1976) и Шориной Н.И. (1994). При Изучение папоротников Юга Российского Дальнего Востока О.В. Храпко была разработана достаточно подробная классификация: выделено 14 морфологических типов вай, 6 типов ветвления и 15 типов корневищ, что в итоге выделило 15 жизненных форм по надземным и подземным частям папоротников (Храпко, 1996). Рядом авторов (Halle, Oldeman, 1970 И.И. Гуреевой (2001) был предложен термин «архитектурная модель», которая базируется на основе дихотомического ключа в рассмотрении подземных корневищ и содержит большое количество моделей, типов, групп, секций, вариантов, но не отражает биоморфологических особенностей строения многолетних структур папоротникообразных растений. По мнению некоторых исследователей, (Серебряков, 1964; Raunkiaer, 1934) концепция модульности при классификации жизненных форм папоротникообразных, которая развита на не экологических принципах «архитектурная модель» не может признаваться структурной основой жизненной формы. Раункиером разработана (1934) классификация, в которой выделены 131 биоморфа из основных 13 и 185 вариантов. Нами предложена объединенная классификация жизненных форм растений, базирующаяся на классификациях Ch. Raunkiaer, с дополнением по T. Hosokawa (1949). Разработанная система жизненных форм Ch. Raunkiaer не теряет своей актуальности и может быть использована большинством ботаников в наше время, и прекрасно подходит для выделения жф папоротникообразных растений, а дополнения T. Hosokawa по эпифитным растениям только усиливают данную систему.</p></abstract><trans-abstract xml:lang="en"><p>The life form (LF) of plants reflects their adaptation to past and current soil, climatic, and cenotic conditions of their habitats. A distinctive feature of fern-like plant classifications (Polypodiophyta, including Lycopodiophyta) is the need to consider both the structure of sporophytes and gametophytes. The study of the morphological structures of fern gametophytes, begun in the 19th century by German botanists, revealed the importance of using comparative morphology in approaches to the classification of ferns (Orth, 1936; Goebel,</p><p>1928). The methods of spore germination were analyzed by S. Momose (1937 a,b; 1938 a,b,c; 1939; 1941; 1942). The author distinguished three types of germination: centrifugal, tangential, and centripetal. Nayar B.K. and Kaur S. when studying pteridophytes of India proposed a new classification, which reflected all existing types of germination of spores and formation of prothallia. J.E. Warming (1884), when distinguishing the life forms of ferns, distinguished two groups: I main (3 subgroups) the smallest and II main (2 subgroups, 3 types) with many variants and subvariants. The first approaches in our country to the allocation of LF of plants were successfully developed by I.G. Serebryakov for higher angiosperms (Serebryakov, 1964), where spore plants were not considered. The first classification of spore plants (ferns, horsetails and club mosses) was reflected in the works of A.P. Khokhryakov (1976) and Shorina N.I. (1994). When studying ferns of the south of the Russian Far East, O.V. Khrapko (1996) developed a fairly detailed classification that identified 14 morphological types of fronds, 6 types of branching and 15 types of rhizomes, which ultimately identified 15 LF of the above-ground and underground parts of ferns. A number of authors (Halle, Oldeman, 1970; I.I. Gureeva (2001)) proposed the term "architectural model" which is based on a dichotomous key in the consideration of underground rhizomes and contains a large number of models, types, groups, sections, variants, but does not reflect the biomorphological features of the perennial structure for fern’s-like plants. According to some authors (Serebryakov, 1964; Raunkiaer, 1934) the concept of modularity in the classification of LF of ferns, which is developed on non-ecological principles, the "architectural model" cannot be recognized as the structural basis of a life form. Raunkiaer developed (1934) a classification in which 131 biomorphs from the main 13 and 185 variants are distinguished. For fern-like plants of the boreal, subtropical and tropical zones, distinguished by the presence of evergreen and tree-like species, we are segregated classification of plant life forms, based on the classification of Ch. Raunkiaer (1934), with additions according to T. Hosokawa (1949) for epiphytic plants, which includes 12 main life forms and 31 variants. </p></trans-abstract><kwd-group xml:lang="ru"><kwd>папоротникообразные растения</kwd><kwd>жизненная форма</kwd><kwd>биоморфа</kwd><kwd>классификация</kwd><kwd>экологический принцип</kwd></kwd-group><kwd-group xml:lang="en"><kwd>pteridophytes</kwd><kwd>life form</kwd><kwd>biomorph</kwd><kwd>classification</kwd><kwd>ecological principle</kwd></kwd-group><funding-group><funding-statement xml:lang="en">The study carried out within the framework of the state assignment «History of creation, state, and development potential for living plant collections of the Peter the Great Botanical Garden of the Botanical Institute of the Russian Academy of Sciences» (124020100075-2)</funding-statement></funding-group></article-meta></front><back><ref-list><title>References</title><ref id="cit1"><label>1</label><citation-alternatives><mixed-citation xml:lang="ru">Antonova I.S., Lagunova N.G. O modul'noy organizatsii nekotorykh grupp vysshikh rasteniy // Zhurnal obshchey biologii. – 1999. – Vol. 60, №1. – P. 49-59.</mixed-citation><mixed-citation xml:lang="en">Antonova I.S., Lagunova N.G. O modul'noy organizatsii nekotorykh grupp vysshikh rasteniy // Zhurnal obshchey biologii. – 1999. – Vol. 60, №1. – P. 49-59.</mixed-citation></citation-alternatives></ref><ref id="cit2"><label>2</label><citation-alternatives><mixed-citation xml:lang="ru">Atkinson L.R., Stokey A.G. Comparative morphology of homosporous ferns. Phytomorphology. – 1964. – Vol. 14, No. 1. – p. 51-70.</mixed-citation><mixed-citation xml:lang="en">Atkinson L.R., Stokey A.G. Comparative morphology of homosporous ferns. Phytomorphology. – 1964. – Vol. 14, No. 1. – p. 51-70.</mixed-citation></citation-alternatives></ref><ref id="cit3"><label>3</label><citation-alternatives><mixed-citation xml:lang="ru">Blasi C., Mazzoleni S., Spada F. &amp; Stanisci A. Life forms variability of mediterranean sclerophyllous forests. Vegetatio, 1990. – Vol. 88 (2). – p. 93-102.</mixed-citation><mixed-citation xml:lang="en">Blasi C., Mazzoleni S., Spada F. &amp; Stanisci A. Life forms variability of mediterranean sclerophyllous forests. Vegetatio, 1990. – Vol. 88 (2). – p. 93-102.</mixed-citation></citation-alternatives></ref><ref id="cit4"><label>4</label><citation-alternatives><mixed-citation xml:lang="ru">Brown J.G. Prothallia of Lycopodium in America. Wiley on behalf of the Ecological Society of America. The Plant World, 1917. –Vol. 20, No. 7. – p. 225-226</mixed-citation><mixed-citation xml:lang="en">Brown J.G. Prothallia of Lycopodium in America. Wiley on behalf of the Ecological Society of America. The Plant World, 1917. –Vol. 20, No. 7. – p. 225-226</mixed-citation></citation-alternatives></ref><ref id="cit5"><label>5</label><citation-alternatives><mixed-citation xml:lang="ru">Bruchmann H. Uber die prothallien und die keimpflanzen mehrerer europaischer Lycopodien und zwar uber die von Lycopodium clavatum, L. annotinum, L. complanatum und L. selago. Gotha, Friedrich Adreas Perthes, 1898. – P. 1-119</mixed-citation><mixed-citation xml:lang="en">Bruchmann H. Uber die prothallien und die keimpflanzen mehrerer europaischer Lycopodien und zwar uber die von Lycopodium clavatum, L. annotinum, L. complanatum und L. selago. Gotha, Friedrich Adreas Perthes, 1898. – P. 1-119</mixed-citation></citation-alternatives></ref><ref id="cit6"><label>6</label><citation-alternatives><mixed-citation xml:lang="ru">Burk W. Vorlaugige mittheilung uber die entwickelungsgeschichte des prothalliums von Anemia. Bot. Zeit. Bd., 1875. – p. 499-501.</mixed-citation><mixed-citation xml:lang="en">Burk W.  Vorlaugige mittheilung uber die entwickelungsgeschichte des prothalliums von Anemia. Bot. Zeit. Bd., 1875. – p. 499-501.</mixed-citation></citation-alternatives></ref><ref id="cit7"><label>7</label><citation-alternatives><mixed-citation xml:lang="ru">Cain S.A. Life-forms and phytoclimate. The Botanical Review, 1950. – Vol. 16(1). – p. 1-32.</mixed-citation><mixed-citation xml:lang="en">Cain S.A. Life-forms and phytoclimate. The Botanical Review, 1950. – Vol. 16(1). – p. 1-32.</mixed-citation></citation-alternatives></ref><ref id="cit8"><label>8</label><citation-alternatives><mixed-citation xml:lang="ru">Chamberlain C.J. Prothallia and sporelings of three New Zealand species of Lycopodium. Bot. Gaz. – 1917. – p. 51-65.</mixed-citation><mixed-citation xml:lang="en">Chamberlain C.J. Prothallia and sporelings of three New Zealand species of Lycopodium. Bot. Gaz. – 1917. – p. 51-65.</mixed-citation></citation-alternatives></ref><ref id="cit9"><label>9</label><citation-alternatives><mixed-citation xml:lang="ru">Conti L., Mendez-Castro F.E., Chytry M., Götzenberger L., Hajek M., Horsak M. et al. Insularity promotes plant persistence strategies in edaphic Island systems. Global Ecology and Biogeography, 2022. – No. 31(4). – p. 753-764.</mixed-citation><mixed-citation xml:lang="en">Conti L., Mendez-Castro F.E., Chytry M., Götzenberger L., Hajek M., Horsak M. et al. Insularity promotes plant persistence strategies in edaphic Island systems. Global Ecology and Biogeography, 2022. – No. 31(4). – p. 753-764.</mixed-citation></citation-alternatives></ref><ref id="cit10"><label>10</label><citation-alternatives><mixed-citation xml:lang="ru">Danin A. &amp; Orshan G. The distribution of Raunkiaer life forms in Israel in relation to the environment. Journal of Vegetation Science, 1990. – No. 1(1). – p. 41-48.</mixed-citation><mixed-citation xml:lang="en">Danin A. &amp; Orshan G. The distribution of Raunkiaer life forms in Israel in relation to the environment. Journal of Vegetation Science, 1990. – No. 1(1). – p. 41-48.</mixed-citation></citation-alternatives></ref><ref id="cit11"><label>11</label><citation-alternatives><mixed-citation xml:lang="ru">Ellenberg I. &amp; Mueller-Doubois D. A key to Raunkier plant life forms with revised subdivisions. Berichte des Geobotanischen Institutes der Eidg. Technischen Hochschule, Stiftung Rübel, 1967. – No. 378. – p. 56-73.</mixed-citation><mixed-citation xml:lang="en">Ellenberg I. &amp; Mueller-Doubois D. A key to Raunkier plant life forms with revised subdivisions. Berichte des Geobotanischen Institutes der Eidg. Technischen Hochschule, Stiftung Rübel, 1967. – No. 378. – p. 56-73.</mixed-citation></citation-alternatives></ref><ref id="cit12"><label>12</label><citation-alternatives><mixed-citation xml:lang="ru">Goebel K. Verzweigung der Farne. Organographie der Pflancen insbesondere der Archegoniaten und Samenpflancen bel. Jena, Verlag von Gustav Fischer, 1928. – p. 95-100.</mixed-citation><mixed-citation xml:lang="en">Goebel K.  Verzweigung der Farne. Organographie der Pflancen insbesondere der Archegoniaten und Samenpflancen bel. Jena, Verlag von Gustav Fischer, 1928. – p. 95-100.</mixed-citation></citation-alternatives></ref><ref id="cit13"><label>13</label><citation-alternatives><mixed-citation xml:lang="ru">Goebel K. Von Entwicklungsgeschichte des prothalliums von Gymnogramme leptophylla Desv. Bor. Zeit., Bd. – p. 671-711.</mixed-citation><mixed-citation xml:lang="en">Goebel K.  Von Entwicklungsgeschichte des prothalliums von Gymnogramme leptophylla Desv. Bor. Zeit., Bd. – p. 671-711.</mixed-citation></citation-alternatives></ref><ref id="cit14"><label>14</label><citation-alternatives><mixed-citation xml:lang="ru">Golubev V.N. Ob analitiko-sinteticheskom modelirovanii zhiznennykh form antofitov // Uspekhi sovremen. biologii. – 1977. – P. 281-293.</mixed-citation><mixed-citation xml:lang="en">Golubev V.N. Ob analitiko-sinteticheskom modelirovanii zhiznennykh form antofitov // Uspekhi sovremen. biologii. – 1977. – P. 281-293.</mixed-citation></citation-alternatives></ref><ref id="cit15"><label>15</label><citation-alternatives><mixed-citation xml:lang="ru">Gureyeva I.I. Ravnosporovyye paporotniki Yuzhnoy Sibiri: sistematika, proiskhozhdeniye, biomorfologiya, populyatsionnaya biologiya. – Tomsk: Izd-vo Tom. gos. un-ta, 2001. – 158 p.</mixed-citation><mixed-citation xml:lang="en">Gureyeva I.I. Ravnosporovyye paporotniki Yuzhnoy Sibiri: sistematika, proiskhozhdeniye, biomorfologiya, populyatsionnaya biologiya. – Tomsk: Izd-vo Tom. gos. un-ta, 2001. – 158 p.</mixed-citation></citation-alternatives></ref><ref id="cit16"><label>16</label><citation-alternatives><mixed-citation xml:lang="ru">Halle F. and Oldeman R.A.A. Essai sur l’architecture et la dynamique de croissance des arbres tropicaux. Masson, Paris, 1970. – 178 p.</mixed-citation><mixed-citation xml:lang="en">Halle F. and Oldeman R.A.A. Essai sur l’architecture et la dynamique de croissance des arbres tropicaux. Masson, Paris, 1970. – 178 p.</mixed-citation></citation-alternatives></ref><ref id="cit17"><label>17</label><citation-alternatives><mixed-citation xml:lang="ru">Halle F., Oldeman R.A.A., Tomlinson R.B. Tropical trees and forests. An Architectural analysis. N.-Y., 1978. – 312 p.</mixed-citation><mixed-citation xml:lang="en">Halle F., Oldeman R.A.A., Tomlinson R.B. Tropical trees and forests. An Architectural analysis. N.-Y., 1978. – 312 p.</mixed-citation></citation-alternatives></ref><ref id="cit18"><label>18</label><citation-alternatives><mixed-citation xml:lang="ru">Harrison S.P., Prentice I.C., Barboni D., Kohfeld K.E., Ni J. &amp; Sutra, J.-P. Ecophysiological and bioclimatic foundations for a global plant functional classification. Journal of Vegetation Science, 2010. – No. 21 (2). – p. 300-317.</mixed-citation><mixed-citation xml:lang="en">Harrison S.P., Prentice I.C., Barboni D., Kohfeld K.E., Ni J. &amp; Sutra, J.-P. Ecophysiological and bioclimatic foundations for a global plant functional classification. Journal of Vegetation Science, 2010. – No. 21 (2). – p. 300-317.</mixed-citation></citation-alternatives></ref><ref id="cit19"><label>19</label><citation-alternatives><mixed-citation xml:lang="ru">Hosokawa Takahide. Studies of the life-form of vascular epiphytes and the spectrum of their life-forms. Journ. Jap. Bot., Vol. XXIV, Nos. 1-12. – 1949. – p. 41-45.</mixed-citation><mixed-citation xml:lang="en">Hosokawa Takahide. Studies of the life-form of vascular epiphytes and the spectrum of their life-forms. Journ. Jap. Bot., Vol. XXIV, Nos. 1-12. – 1949. – p. 41-45.</mixed-citation></citation-alternatives></ref><ref id="cit20"><label>20</label><citation-alternatives><mixed-citation xml:lang="ru">Irl S.D.H., Obermeier A., Beierkuhnlein C. &amp; Steinbauer M.J. Climate controls plant life-form patterns on a high-elevation oceanic island. Journal of Biogeography, 2020. – No. 47 (10). – p. 2261-2273.</mixed-citation><mixed-citation xml:lang="en">Irl S.D.H., Obermeier A., Beierkuhnlein C. &amp; Steinbauer M.J. Climate controls plant life-form patterns on a high-elevation oceanic island. Journal of Biogeography, 2020. – No. 47 (10). – p. 2261-2273.</mixed-citation></citation-alternatives></ref><ref id="cit21"><label>21</label><citation-alternatives><mixed-citation xml:lang="ru">Kamelin R.V. Florgeneticheskiy analiz yestestvennoy flory gornoy Sredney Azii – L.: Nauka, 1973. – 355 p.</mixed-citation><mixed-citation xml:lang="en">Kamelin R.V. Florgeneticheskiy analiz yestestvennoy flory gornoy Sredney Azii – L.: Nauka, 1973. – 355 p.</mixed-citation></citation-alternatives></ref><ref id="cit22"><label>22</label><citation-alternatives><mixed-citation xml:lang="ru">Khokhryakov A.P. Evolyutsiya biomorf rasteniy. M.: Nauka, 1981. – 167 p.</mixed-citation><mixed-citation xml:lang="en">Khokhryakov A.P. Evolyutsiya biomorf rasteniy. M.: Nauka, 1981. – 167 p.</mixed-citation></citation-alternatives></ref><ref id="cit23"><label>23</label><citation-alternatives><mixed-citation xml:lang="ru">Khrapko O.V. Paporotniki yuga Dal'nego Vostoka Rossii (biologiya, ekologiya, voprosy okhrany genofonda). – Vladivostok: Dal'nauka, 1996. – 200 p.</mixed-citation><mixed-citation xml:lang="en">Khrapko O.V. Paporotniki yuga Dal'nego Vostoka Rossii (biologiya, ekologiya, voprosy okhrany genofonda). – Vladivostok: Dal'nauka, 1996. – 200 p.</mixed-citation></citation-alternatives></ref><ref id="cit24"><label>24</label><citation-alternatives><mixed-citation xml:lang="ru">Khokhryakov A.P. Zhiznennyye formy paporotnikoobraznykh // Izv. AN SSSR. Seriya biol. №2. M.; L.: Nauka, 1979. – p. 251-264</mixed-citation><mixed-citation xml:lang="en">Khokhryakov A.P. Zhiznennyye formy paporotnikoobraznykh // Izv. AN SSSR. Seriya biol. №2. M.; L.: Nauka, 1979. – p. 251-264</mixed-citation></citation-alternatives></ref><ref id="cit25"><label>25</label><citation-alternatives><mixed-citation xml:lang="ru">Khokhryakov A.P. Zhiznennyye formy paporotnikoobraznykh i vozmozhnyye puti ikh evolyutsii // Materialy 5-go Moskovskogo soveshchaniya po filogenii rasteniy. M.: Nauka, 1976. – p. 184-187.</mixed-citation><mixed-citation xml:lang="en">Khokhryakov A.P. Zhiznennyye formy paporotnikoobraznykh i vozmozhnyye puti ikh evolyutsii // Materialy 5-go Moskovskogo soveshchaniya po filogenii rasteniy. M.: Nauka, 1976. – p. 184-187.</mixed-citation></citation-alternatives></ref><ref id="cit26"><label>26</label><citation-alternatives><mixed-citation xml:lang="ru">Loidi J., Chytry M., Jimenez-Alfaro B., Alessi N., Biurrun I., Campos J.A. et al. Life-form diversity across temperate deciduous forests of Western Eurasia: a different story in the understory. Journal of Biogeography, 2021. – p. 2932-2945.</mixed-citation><mixed-citation xml:lang="en">Loidi J., Chytry M., Jimenez-Alfaro B., Alessi N., Biurrun I., Campos J.A. et al. Life-form diversity across temperate deciduous forests of Western Eurasia: a different story in the understory. Journal of Biogeography, 2021. – p. 2932-2945.</mixed-citation></citation-alternatives></ref><ref id="cit27"><label>27</label><citation-alternatives><mixed-citation xml:lang="ru">Marini L., Battisti A., Bona E., Federici G., Martini F., Pautasso M. et al. Alien and native plant life-forms respond differently to human and climate pressures. Global Ecology and Biogeography, 2011. – p. 534-544.</mixed-citation><mixed-citation xml:lang="en">Marini L., Battisti A., Bona E., Federici G., Martini F., Pautasso M. et al. Alien and native plant life-forms respond differently to human and climate pressures. Global Ecology and Biogeography, 2011. – p. 534-544.</mixed-citation></citation-alternatives></ref><ref id="cit28"><label>28</label><citation-alternatives><mixed-citation xml:lang="ru">Matteodo M., Wipf S., Stöckli V., Rixen C. &amp; Vittoz P. Elevation gradient of successful plant traits for colonizing alpine summits under climate change. Environmental Research Letters, 2013. – No.8 (2). – p. 024-043.</mixed-citation><mixed-citation xml:lang="en">Matteodo M., Wipf S., Stöckli V., Rixen C. &amp; Vittoz P. Elevation gradient of successful plant traits for colonizing alpine summits under climate change. Environmental Research Letters, 2013. – No.8 (2). – p. 024-043.</mixed-citation></citation-alternatives></ref><ref id="cit29"><label>29</label><citation-alternatives><mixed-citation xml:lang="ru">Mochalov A.S. Paporotniki Urala: avtoref. dis. ... kand-ta biol. nauk / Tomsk: Izdvo Tom. gos. un-ta, 2010. – 23 p.</mixed-citation><mixed-citation xml:lang="en">Mochalov A.S. Paporotniki Urala: avtoref. dis. ... kand-ta biol. nauk / Tomsk: Izdvo Tom. gos. un-ta, 2010. – 23 p.</mixed-citation></citation-alternatives></ref><ref id="cit30"><label>30</label><citation-alternatives><mixed-citation xml:lang="ru">Moles A.T., Warton D.I., Warman L., Swenson N.G., Laffan S.W., Zanne A.E. et al. Global patterns in plant height. Journal of Ecology, 2009. – No. 97 (5). – p. 923-932.</mixed-citation><mixed-citation xml:lang="en">Moles A.T., Warton D.I., Warman L., Swenson N.G., Laffan S.W., Zanne A.E. et al. Global patterns in plant height. Journal of Ecology, 2009. – No. 97 (5). – p. 923-932.</mixed-citation></citation-alternatives></ref><ref id="cit31"><label>31</label><citation-alternatives><mixed-citation xml:lang="ru">Momose S. Studies on the gametophyte of ferns. V. On the prothallium of eudryopteroid ferns. Journ. Jap. Bot, 1938. – Vol. 14, No 7. – p. 445-453.</mixed-citation><mixed-citation xml:lang="en">Momose S.  Studies on the gametophyte of ferns. V. On the prothallium of eudryopteroid ferns. Journ. Jap. Bot, 1938. – Vol. 14, No 7. – p. 445-453.</mixed-citation></citation-alternatives></ref><ref id="cit32"><label>32</label><citation-alternatives><mixed-citation xml:lang="ru">Momose S. Studies on the gametophyte of ferns. I. On the prothallial development and prothallium of Dryopteris varia O. Kunze. Journ. Jap. Bot, 1937. – Vol.12, No 3. – p. 113-120.</mixed-citation><mixed-citation xml:lang="en">Momose S. Studies on the gametophyte of ferns. I. On the prothallial development and prothallium of Dryopteris varia O. Kunze. Journ. Jap. Bot, 1937. – Vol.12, No 3. – p. 113-120.</mixed-citation></citation-alternatives></ref><ref id="cit33"><label>33</label><citation-alternatives><mixed-citation xml:lang="ru">Momose S. Studies on the gametophyte of ferns. II. On the prothallial development and prothallium of Cyrtomium falcatum J. Smith and Rumohra standischii. Journ. Jap. Bot, 1937. – Vol. 13, No 2. – p. 113-120.</mixed-citation><mixed-citation xml:lang="en">Momose S. Studies on the gametophyte of ferns. II. On the prothallial development and prothallium of Cyrtomium falcatum J. Smith and Rumohra standischii. Journ. Jap. Bot, 1937. – Vol. 13, No 2. – p. 113-120.</mixed-citation></citation-alternatives></ref><ref id="cit34"><label>34</label><citation-alternatives><mixed-citation xml:lang="ru">Momose S. Studies on the gametophyte of ferns. III. On the prothallium of Polystichum aculeatum Roth var. taiwanense Nakai et Momose and Leptogramma totta J. Smith. Journ. Jap. Bot, 1938. – Vol. 14, No 1. – p. 56-62.</mixed-citation><mixed-citation xml:lang="en">Momose S. Studies on the gametophyte of ferns. III. On the prothallium of Polystichum aculeatum Roth var. taiwanense Nakai et Momose and Leptogramma totta J. Smith. Journ. Jap. Bot, 1938. – Vol. 14, No 1. –  p. 56-62.</mixed-citation></citation-alternatives></ref><ref id="cit35"><label>35</label><citation-alternatives><mixed-citation xml:lang="ru">Momose S. Studies on the gametophyte of ferns. IV. On the prothallium of Diplazium thunbergii Nakai and D. latifolium Moore. Journ. Jap. Bot. – p. 265-273.</mixed-citation><mixed-citation xml:lang="en">Momose S. Studies on the gametophyte of ferns. IV. On the prothallium of Diplazium thunbergii Nakai and D. latifolium Moore. Journ. Jap. Bot. – p. 265-273.</mixed-citation></citation-alternatives></ref><ref id="cit36"><label>36</label><citation-alternatives><mixed-citation xml:lang="ru">Momose S. Studies on the gametophyte of ferns. XII. Prothallium of Athyrioid ferns. Journ. Jap. Bot, 1939. – p. 712-726.</mixed-citation><mixed-citation xml:lang="en">Momose S. Studies on the gametophyte of ferns. XII. Prothallium of Athyrioid ferns. Journ. Jap. Bot, 1939. – p. 712-726.</mixed-citation></citation-alternatives></ref><ref id="cit37"><label>37</label><citation-alternatives><mixed-citation xml:lang="ru">Momose S. Studies on the gametophyte of ferns. XVI. Prothallium of Thelypterioid ferns. Journ. Jap. Bot, 1941. – p. 35-51.</mixed-citation><mixed-citation xml:lang="en">Momose S. Studies on the gametophyte of ferns. XVI. Prothallium of Thelypterioid ferns. Journ. Jap. Bot, 1941. – p. 35-51.</mixed-citation></citation-alternatives></ref><ref id="cit38"><label>38</label><citation-alternatives><mixed-citation xml:lang="ru">Momose S. Studies on the gametophyte of ferns. XXVII.. On the systematic criteria of the prothallia polypodiaceous ferns. Journ. Jap. Bot, 1942. – p. 391-403.</mixed-citation><mixed-citation xml:lang="en">Momose S. Studies on the gametophyte of ferns. XXVII.. On the systematic criteria of the prothallia polypodiaceous ferns. Journ. Jap. Bot, 1942. – p. 391-403.</mixed-citation></citation-alternatives></ref><ref id="cit39"><label>39</label><citation-alternatives><mixed-citation xml:lang="ru">Mueller R.J. Shoot morphology of the climbing fern Lygodium (Schizaeaceae): general organography, leaf initiation, and branching. Bot. Gaz., 1982. – p. 319-330.</mixed-citation><mixed-citation xml:lang="en">Mueller R.J. Shoot morphology of the climbing fern Lygodium (Schizaeaceae): general organography, leaf initiation, and branching. Bot. Gaz., 1982. – p. 319-330.</mixed-citation></citation-alternatives></ref><ref id="cit40"><label>40</label><citation-alternatives><mixed-citation xml:lang="ru">Nayar B.K., Kaur S. Gametophytes of homosporous ferns. The Botanical Review, 1971. – Vol. 37, No. 3. – p. 295-395.</mixed-citation><mixed-citation xml:lang="en">Nayar B.K., Kaur S. Gametophytes of homosporous ferns. The Botanical Review, 1971. – Vol. 37, No. 3. – p. 295-395.</mixed-citation></citation-alternatives></ref><ref id="cit41"><label>41</label><citation-alternatives><mixed-citation xml:lang="ru">Nishida M. Types of spore germination in the ferns. Jour. Jap. Bot., 1965. – Vol. 40, No. 6. – p. 161-171.</mixed-citation><mixed-citation xml:lang="en">Nishida M. Types of spore germination in the ferns. Jour. Jap. Bot., 1965. – Vol. 40, No. 6. – p. 161-171.</mixed-citation></citation-alternatives></ref><ref id="cit42"><label>42</label><citation-alternatives><mixed-citation xml:lang="ru">Olson M.E., Soriano D., Rosell J.A., Anfodillo T., Donoghue M.J., Edwards E.J. et al. Plant height and hydraulic vulnerability to drought and cold. Proceedings of the National Academy of Sciences, 2018. – No. 115 (29). – p. 7551-7556.</mixed-citation><mixed-citation xml:lang="en">Olson M.E., Soriano D., Rosell J.A., Anfodillo T., Donoghue M.J., Edwards E.J. et al. Plant height and hydraulic vulnerability to drought and cold. Proceedings of the National Academy of Sciences, 2018. – No. 115 (29). – p. 7551-7556.</mixed-citation></citation-alternatives></ref><ref id="cit43"><label>43</label><citation-alternatives><mixed-citation xml:lang="ru">Orth R. Morphologische und physiologische untersuchungen an farnprothallien. Planta, Bd. – 1936, – p. 104-150.</mixed-citation><mixed-citation xml:lang="en">Orth R. Morphologische und physiologische untersuchungen an farnprothallien. Planta, Bd. – 1936, – p. 104-150.</mixed-citation></citation-alternatives></ref><ref id="cit44"><label>44</label><citation-alternatives><mixed-citation xml:lang="ru">Ottaviani G., Keppel G., Gotzenberger L., Harrison S., Opedal O.H., Conti L. et al. Linking plant functional ecology to island biogeography. Trends in Plant Science, 2020. – No. 25 (4). – p. 329-339.</mixed-citation><mixed-citation xml:lang="en">Ottaviani G., Keppel G., Gotzenberger L., Harrison S., Opedal O.H., Conti L. et al. Linking plant functional ecology to island biogeography. Trends in Plant Science, 2020. – No. 25 (4). – p. 329-339.</mixed-citation></citation-alternatives></ref><ref id="cit45"><label>45</label><citation-alternatives><mixed-citation xml:lang="ru">Pavon N.P., Hernandez-Trejo H. &amp; Rico-Gray V. Distribution of plant life forms along an altitudinal gradient in the semi-arid valley of Zapotitlan, Mexico. Journal of Vegetation Science, 2000. – No.11 (1). – p. 39-42.</mixed-citation><mixed-citation xml:lang="en">Pavon N.P., Hernandez-Trejo H. &amp; Rico-Gray V. Distribution of plant life forms along an altitudinal gradient in the semi-arid valley of Zapotitlan, Mexico. Journal of Vegetation Science, 2000. – No.11 (1).  – p. 39-42.</mixed-citation></citation-alternatives></ref><ref id="cit46"><label>46</label><citation-alternatives><mixed-citation xml:lang="ru">Raunkiaer Ch. The life forms of plants and statistical plant geography. Oxford, 1934. – p. 2-104.</mixed-citation><mixed-citation xml:lang="en">Raunkiaer Ch. The life forms of plants and statistical plant geography. Oxford, 1934. – p. 2-104.</mixed-citation></citation-alternatives></ref><ref id="cit47"><label>47</label><citation-alternatives><mixed-citation xml:lang="ru">Serebryakov I.G. Ekologicheskaya morfologiya rasteniy: zhiznennyye formy pokrytosemennykh i khvoynykh. – M.: Vyssh. shk., 1962. – 378 p.</mixed-citation><mixed-citation xml:lang="en">Serebryakov I.G. Ekologicheskaya morfologiya rasteniy: zhiznennyye formy pokrytosemennykh i khvoynykh. – M.: Vyssh. shk., 1962. – 378 p.</mixed-citation></citation-alternatives></ref><ref id="cit48"><label>48</label><citation-alternatives><mixed-citation xml:lang="ru">Serebryakova T.I. Morfogenez pobegov i evolyutsiya zhiznennykh form zlakov. Moskva: Nauka, 1971. – 359 p.</mixed-citation><mixed-citation xml:lang="en">Serebryakova T.I. Morfogenez pobegov i evolyutsiya zhiznennykh form zlakov. Moskva: Nauka, 1971. – 359 p.</mixed-citation></citation-alternatives></ref><ref id="cit49"><label>49</label><citation-alternatives><mixed-citation xml:lang="ru">Serebryakova T.I. Ob osnovnykh «arkhitekturnykh modelyakh» travyanistykh mnogoletnikov i modusakh ikh preobrazovaniya // Byul. MOIP. Otd. biol. – 1977. – P. 84169.</mixed-citation><mixed-citation xml:lang="en">Serebryakova T.I. Ob osnovnykh «arkhitekturnykh modelyakh» travyanistykh mnogoletnikov i modusakh ikh preobrazovaniya // Byul. MOIP. Otd. biol. – 1977. – P. 84169.</mixed-citation></citation-alternatives></ref><ref id="cit50"><label>50</label><citation-alternatives><mixed-citation xml:lang="ru">Serebryakova T.I. Zhiznennyye formy i modeli pobegoobrazovaniya nazemnopolzuchikh mnogoletnikh trav // Zhiznennyye formy: struktura, spektry i evolyutsiya. – M., 1981. – P. 161-179.</mixed-citation><mixed-citation xml:lang="en">Serebryakova T.I. Zhiznennyye formy i modeli pobegoobrazovaniya nazemnopolzuchikh mnogoletnikh trav // Zhiznennyye formy: struktura, spektry i evolyutsiya. – M., 1981. – P. 161-179.</mixed-citation></citation-alternatives></ref><ref id="cit51"><label>51</label><citation-alternatives><mixed-citation xml:lang="ru">Shorina N.I. Ekologicheskaya morfologiya i populyatsionnaya biologiya predstaviteley podklassa Polypodiidea: avtoref. dis.... d-ra biol. Nauk. – M., 1994. – 34 p.</mixed-citation><mixed-citation xml:lang="en">Shorina N.I. Ekologicheskaya morfologiya i populyatsionnaya biologiya predstaviteley podklassa Polypodiidea: avtoref. dis.... d-ra biol. Nauk. – M., 1994. – 34 p.</mixed-citation></citation-alternatives></ref><ref id="cit52"><label>52</label><citation-alternatives><mixed-citation xml:lang="ru">Shorina N.I. Metameriya sporofitov paporotnikov v svyazi s ikh vetvleniyem // Uspekhi sovremennoy botaniki: tezisy dokladov delegatskogo s"yezda VBO. Alma-Ata, 1988. – 397 p.</mixed-citation><mixed-citation xml:lang="en">Shorina N.I. Metameriya sporofitov paporotnikov v svyazi s ikh vetvleniyem // Uspekhi sovremennoy botaniki: tezisy dokladov delegatskogo s"yezda VBO. Alma-Ata, 1988. – 397 p.</mixed-citation></citation-alternatives></ref><ref id="cit53"><label>53</label><citation-alternatives><mixed-citation xml:lang="ru">Shorina N.I., Derzhavina N.M. Modeli rosta, osobennosti metamerii nekotorykh paporotnikov i vozmozhnyye modusy ikh morfologicheskoy evolyutsii. Morfologicheskaya evolyutsiya vysshikh rasteniy // Materialy VI Moskovskogo soveshchaniya po filogenii rasteniy. – M., 1981. – P. 14-23.</mixed-citation><mixed-citation xml:lang="en">Shorina N.I., Derzhavina N.M. Modeli rosta, osobennosti metamerii nekotorykh paporotnikov i vozmozhnyye modusy ikh morfologicheskoy evolyutsii. Morfologicheskaya evolyutsiya vysshikh rasteniy // Materialy VI Moskovskogo soveshchaniya po filogenii rasteniy. – M., 1981. – P. 14-23.</mixed-citation></citation-alternatives></ref><ref id="cit54"><label>54</label><citation-alternatives><mixed-citation xml:lang="ru">Smith W.G. Raunkiaer’s life forms and statistical methods. Journal of Ecology, 1913. – No. 1. – p. 16-26.</mixed-citation><mixed-citation xml:lang="en">Smith W.G. Raunkiaer’s life forms and statistical methods. Journal of Ecology, 1913. – No. 1. – p. 16-26.</mixed-citation></citation-alternatives></ref><ref id="cit55"><label>55</label><citation-alternatives><mixed-citation xml:lang="ru">Stanley A. Cain. Life-forms and phytoclamate. The botanical review, 1950. – Vol. XVI (1). – p. 1-32.</mixed-citation><mixed-citation xml:lang="en">Stanley A. Cain. Life-forms and phytoclamate. The botanical review, 1950. – Vol. XVI (1). – p. 1-32.</mixed-citation></citation-alternatives></ref><ref id="cit56"><label>56</label><citation-alternatives><mixed-citation xml:lang="ru">Troll W. Vergleichende Morphologie der hoheren Pflancen. Verlag von Gebruder Borntrager. Berlin, Bd. 1., Teil. 1. – 1937.</mixed-citation><mixed-citation xml:lang="en">Troll W. Vergleichende Morphologie der hoheren Pflancen. Verlag von Gebruder Borntrager. Berlin, Bd. 1., Teil. 1. – 1937.</mixed-citation></citation-alternatives></ref><ref id="cit57"><label>57</label><citation-alternatives><mixed-citation xml:lang="ru">Vin'kovskaya O.P. Flora Irkutskoy gorodskoy aglomeratsii i yeye dinamika za posledniye 125 let: avtoref. dis. … kand. biol. nauk / Irkutsk, 2005. – 241 p.</mixed-citation><mixed-citation xml:lang="en">Vin'kovskaya O.P. Flora Irkutskoy gorodskoy aglomeratsii i yeye dinamika za posledniye 125 let: avtoref. dis. … kand. biol. nauk / Irkutsk, 2005. – 241 p.</mixed-citation></citation-alternatives></ref><ref id="cit58"><label>58</label><citation-alternatives><mixed-citation xml:lang="ru">Violle C., Reich P.B., Pacala S.W., Enquist B.J. &amp; Kattge J. The emergence and promise of functional biogeography. Proceedings of the National Academy of Sciences. – 2014. – p. 13690-13696.</mixed-citation><mixed-citation xml:lang="en">Violle C., Reich P.B., Pacala S.W., Enquist B.J. &amp; Kattge J. The emergence and promise of functional biogeography. Proceedings of the National Academy of Sciences. – 2014. – p. 13690-13696.</mixed-citation></citation-alternatives></ref><ref id="cit59"><label>59</label><citation-alternatives><mixed-citation xml:lang="ru">Warming E. Om Skudbygning, Overvintring og Foryngelse [título traducido: Arquitectura de rodaje, perennización y rejuvenecimiento]. Naturhistorisk Forenings Festskrift, 1884. – p. 1-105.</mixed-citation><mixed-citation xml:lang="en">Warming E. Om Skudbygning, Overvintring og Foryngelse [título traducido: Arquitectura de rodaje, perennización y rejuvenecimiento]. Naturhistorisk Forenings Festskrift, 1884. – p. 1-105.</mixed-citation></citation-alternatives></ref></ref-list><fn-group><fn fn-type="conflict"><p>The authors declare that there are no conflicts of interest present.</p></fn></fn-group></back></article>
