The Approaches to the Classification of Life Forms of Fern-Like Plants (Polypodiophyta, Including Lycopodiophyta)

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,

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. 

1. Antonova I.S., Lagunova N.G. O modul'noy organizatsii nekotorykh grupp vysshikh rasteniy // Zhurnal obshchey biologii. – 1999. – Vol. 60, №1. – P. 49-59.

2. Atkinson L.R., Stokey A.G. Comparative morphology of homosporous ferns. Phytomorphology. – 1964. – Vol. 14, No. 1. – p. 51-70.

3. Blasi C., Mazzoleni S., Spada F. & Stanisci A. Life forms variability of mediterranean sclerophyllous forests. Vegetatio, 1990. – Vol. 88 (2). – p. 93-102.

4. 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

5. 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

6. Burk W. Vorlaugige mittheilung uber die entwickelungsgeschichte des prothalliums von Anemia. Bot. Zeit. Bd., 1875. – p. 499-501.

7. Cain S.A. Life-forms and phytoclimate. The Botanical Review, 1950. – Vol. 16(1). – p. 1-32.

8. Chamberlain C.J. Prothallia and sporelings of three New Zealand species of Lycopodium. Bot. Gaz. – 1917. – p. 51-65.

9. 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.

10. Danin A. & 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.

11. Ellenberg I. & 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.

12. Goebel K. Verzweigung der Farne. Organographie der Pflancen insbesondere der Archegoniaten und Samenpflancen bel. Jena, Verlag von Gustav Fischer, 1928. – p. 95-100.

13. Goebel K. Von Entwicklungsgeschichte des prothalliums von Gymnogramme leptophylla Desv. Bor. Zeit., Bd. – p. 671-711.

14. Golubev V.N. Ob analitiko-sinteticheskom modelirovanii zhiznennykh form antofitov // Uspekhi sovremen. biologii. – 1977. – P. 281-293.

15. Gureyeva I.I. Ravnosporovyye paporotniki Yuzhnoy Sibiri: sistematika, proiskhozhdeniye, biomorfologiya, populyatsionnaya biologiya. – Tomsk: Izd-vo Tom. gos. un-ta, 2001. – 158 p.

16. Halle F. and Oldeman R.A.A. Essai sur l’architecture et la dynamique de croissance des arbres tropicaux. Masson, Paris, 1970. – 178 p.

17. Halle F., Oldeman R.A.A., Tomlinson R.B. Tropical trees and forests. An Architectural analysis. N.-Y., 1978. – 312 p.

18. Harrison S.P., Prentice I.C., Barboni D., Kohfeld K.E., Ni J. & Sutra, J.-P. Ecophysiological and bioclimatic foundations for a global plant functional classification. Journal of Vegetation Science, 2010. – No. 21 (2). – p. 300-317.

19. 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.

20. Irl S.D.H., Obermeier A., Beierkuhnlein C. & 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.

21. Kamelin R.V. Florgeneticheskiy analiz yestestvennoy flory gornoy Sredney Azii – L.: Nauka, 1973. – 355 p.

22. Khokhryakov A.P. Evolyutsiya biomorf rasteniy. M.: Nauka, 1981. – 167 p.

23. Khrapko O.V. Paporotniki yuga Dal'nego Vostoka Rossii (biologiya, ekologiya, voprosy okhrany genofonda). – Vladivostok: Dal'nauka, 1996. – 200 p.

24. Khokhryakov A.P. Zhiznennyye formy paporotnikoobraznykh // Izv. AN SSSR. Seriya biol. №2. M.; L.: Nauka, 1979. – p. 251-264

25. 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.

26. 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.

27. 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.

28. Matteodo M., Wipf S., Stöckli V., Rixen C. & 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.

29. Mochalov A.S. Paporotniki Urala: avtoref. dis. ... kand-ta biol. nauk / Tomsk: Izdvo Tom. gos. un-ta, 2010. – 23 p.

30. 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.

31. 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.

32. 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.

33. 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.

34. 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.

35. 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.

36. Momose S. Studies on the gametophyte of ferns. XII. Prothallium of Athyrioid ferns. Journ. Jap. Bot, 1939. – p. 712-726.

37. Momose S. Studies on the gametophyte of ferns. XVI. Prothallium of Thelypterioid ferns. Journ. Jap. Bot, 1941. – p. 35-51.

38. 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.

39. Mueller R.J. Shoot morphology of the climbing fern Lygodium (Schizaeaceae): general organography, leaf initiation, and branching. Bot. Gaz., 1982. – p. 319-330.

40. Nayar B.K., Kaur S. Gametophytes of homosporous ferns. The Botanical Review, 1971. – Vol. 37, No. 3. – p. 295-395.

41. Nishida M. Types of spore germination in the ferns. Jour. Jap. Bot., 1965. – Vol. 40, No. 6. – p. 161-171.

42. 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.

43. Orth R. Morphologische und physiologische untersuchungen an farnprothallien. Planta, Bd. – 1936, – p. 104-150.

44. 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.

45. Pavon N.P., Hernandez-Trejo H. & 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.

46. Raunkiaer Ch. The life forms of plants and statistical plant geography. Oxford, 1934. – p. 2-104.

47. Serebryakov I.G. Ekologicheskaya morfologiya rasteniy: zhiznennyye formy pokrytosemennykh i khvoynykh. – M.: Vyssh. shk., 1962. – 378 p.

48. Serebryakova T.I. Morfogenez pobegov i evolyutsiya zhiznennykh form zlakov. Moskva: Nauka, 1971. – 359 p.

49. Serebryakova T.I. Ob osnovnykh «arkhitekturnykh modelyakh» travyanistykh mnogoletnikov i modusakh ikh preobrazovaniya // Byul. MOIP. Otd. biol. – 1977. – P. 84169.

50. Serebryakova T.I. Zhiznennyye formy i modeli pobegoobrazovaniya nazemnopolzuchikh mnogoletnikh trav // Zhiznennyye formy: struktura, spektry i evolyutsiya. – M., 1981. – P. 161-179.

51. Shorina N.I. Ekologicheskaya morfologiya i populyatsionnaya biologiya predstaviteley podklassa Polypodiidea: avtoref. dis.... d-ra biol. Nauk. – M., 1994. – 34 p.

52. 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.

53. 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.

54. Smith W.G. Raunkiaer’s life forms and statistical methods. Journal of Ecology, 1913. – No. 1. – p. 16-26.

55. Stanley A. Cain. Life-forms and phytoclamate. The botanical review, 1950. – Vol. XVI (1). – p. 1-32.

56. Troll W. Vergleichende Morphologie der hoheren Pflancen. Verlag von Gebruder Borntrager. Berlin, Bd. 1., Teil. 1. – 1937.

57. Vin'kovskaya O.P. Flora Irkutskoy gorodskoy aglomeratsii i yeye dinamika za posledniye 125 let: avtoref. dis. … kand. biol. nauk / Irkutsk, 2005. – 241 p.

58. Violle C., Reich P.B., Pacala S.W., Enquist B.J. & Kattge J. The emergence and promise of functional biogeography. Proceedings of the National Academy of Sciences. – 2014. – p. 13690-13696.

59. Warming E. Om Skudbygning, Overvintring og Foryngelse [título traducido: Arquitectura de rodaje, perennización y rejuvenecimiento]. Naturhistorisk Forenings Festskrift, 1884. – p. 1-105.