Núm. 127 (2020)
Artículo de investigación

Gametophytic phase of three taxa of Anemia (Anemiaceae)

Cristina Janet Chambi
Universidad Nacional de Salta
Olga Gladys Martínez
Universidad Nacional de Salta, IBIGEO (Instituto de Bio y Geociencias del Noroeste Argentino, CONICET-UNSa), Facultad de Ciencias Naturales

Publicado 2020-07-09

Palabras clave

  • ferns,
  • gametophytes,
  • germination,
  • spores,
  • trichomes



Background and Aims: Anemiaceae has a single genus, Anemia, with 115 species distributed in the Neotropics, Africa, India, and islands in the Indian Ocean. The aim of this work is characterize living spores and gametophytic phase of three Anemia taxa from South America, little studied or with fragmentary studies, A. herzogii, A. tomentosa var. anthriscifolia and A. tomentosa var. tomentosa.
Methods: Gametophytes of the three taxa have been obtained through in vitro cultures of spores. The spores and gametophytes were studied with light and scanning electron microscopes. The germination pattern of the spores, the evolution of the gametophytic development and the sexual expression were described.
Key results: Filamentous gametophytes were formed by 4-20 cells, the adults were cordate, symmetrical or asymmetrical, bisexual and protandric, on their surfaces and margins clavate trichomes were observed. Gametangia corresponded to the leptosporangiate type. The dehiscence of antheridia was produced by a perforation in the opercular cell. Archegonia had necks with four to seven cells and four triangular cells at the distal end of the neck. Anemia herzogii showed branched trichomes on the gametophytes and later developed sporophytes.
Conclusions: Germination in all taxa corresponds to the Anemia-type and the gametophytic development to the Ceratopteris-type. Anemia herzogii is distinguished from the two varieties by the presence of branched trichomes exclusive of the gametophytic phase. The studied taxa are characterized by the morphology of gametophyte, type of trichomes and sexuality.


  1. Atkinson, L. R. 1962. The Schizaeaceae: The gametophyte of Anemia. Phytomorphology 12: 264-288.
  2. De la Sota, E. R. and J. T. Mickel. 1968. Sinopsis de las especies argentinas del género Anemia Swartz (Schizaeaceae). Revista del Museo de La Plata, Nueva serie Sección Botánica 11(61): 133-152.
  3. De la Sota, E. R. and M. A. Morbelli. 1987. Schizaeales. Phytomorphology 37(4): 365-393.
  4. Dyer, A. F. 1979. The culture of fern gametophytes for experimental investigation. In: Dyer, A. F. (ed.). The experimental biology of ferns. Academic Press. London, England. Pp. 254-305.
  5. Edwards, M. E. and J. H. Miller. 1972. Growth regulation by ethylene in fern gametophytes III. Inhibition of spore germination. American Journal of Botany 59(5): 458-465. DOI: https://doi.org/10.1002/j.1537-2197.1972.tb10117.x
  6. Escamilla-Aquino, A. M., M. L. Arreguín-Sánchez and R. Fernández-Nava. 2008. Ciclos Biológicos de Anemia munchi Christ (Schizaeaceae-Pteridophyta) y Pityrogramma calomelanos (L.) Link (Adiantaceae-Pteridophyta). Polibotánica 26: 101-111.
  7. Grill, R. 1988. Photocontrol of Gibberellin-induced precocious antheridium formation in the fern Anemia phyllitidis L. Sw. Journal of Plant Physiology 133(3): 381-384. DOI: https://doi.org/10.1016/S0176-1617(88)80222-0
  8. Hernández-Solano, J. 2004. Descripción de las fases del ciclo de vida de Adiantum concinnum Humb. et Blonpl. ex Willd. (Adiantaceae-Pteridophyta y Anemia adiantifolia (Schizaceae-Pteridophyta). Tesis de licenciatura. Escuela Nacional de Ciencias Biológicas, Instituto Politécnico Nacional. Cd. Mx., México. 52 pp.
  9. Labiak, P. H., J. T. Mickel and J. G. Hanks. 2015. Molecular phylogeny and character evolution of Anemiaceae (Schizaeales). Taxon 64(6): 1141-1158. DOI: https://doi.org/10.12705/646.3
  10. Luján, M. C., R. Morero and G. E. Barboza. 2011. Estudios epidérmicos en helechos y licófitas medicinales de la Provincia de Córdoba, Argentina. Hoehnea 38(4): 609-659. DOI: https://doi.org/10.1590/S2236-89062011000400007
  11. Martínez, O. G. 2010. Gametófitos y esporófitos jóvenes de cuatro especies naturalizadas de Pteris (Pteridaceae) en América. Revista de Biología Tropical 58(1): 81-92. DOI: https://doi.org/10.15517/rbt.v58i1.5196
  12. Martínez, O. G., M. A. Hernández and M. M. Ponce. 2017. Reproductive expression of cheilanthoid ferns (Pteridaceae) from South America. Flora 236-237: 126-131. DOI: https://doi.org/10.1016/j.flora.2017.10.003
  13. Mickel, J. T. 1962. A monographic study of the fern genus Anemia, subgenus Coptophyllum. Iowa State College Journal of Science 36: 349-482.
  14. Mickel, J. T. 2016. Anemia (Anemiaceae). Flora Neotropica Monograph 118: 1-182.
  15. Miller, J. H. 1968. Fern gametophytes as experimental material. The Botanical Review 34: 361-440. DOI: https://doi.org/10.1007/BF02859133
  16. Näf, U. 1969. On the control of antheridium formation in ferns. In: Gunckel, J. E. (ed.). Current Topics in Plant Science. Academic Press. New York, USA. Pp. 97-116.
  17. Näf, U., K. Nakanishi and M. Endo. 1975. On the physiology and chemistry of fern antheridiogens. The Botanical Review 41(3): 315-359. DOI: https://doi.org/10.1007/BF02860840
  18. Nayar, B. K. and S. Kaur. 1971. Gametophytes of homosporous ferns. The Botanical Review 37: 330-331. DOI: https://doi.org/10.1007/BF02859157
  19. Nester, J. E. 1985. Scanning Electron Microscopy of antheridia and archegonia of Anemia mexicana Klotzsch. American Journal of Botany 72(5): 777-780. DOI: https://doi.org/10.2307/2443692
  20. Nester, J. E. and R. C. Coolbaugh. 1986. Factors influencing spore germination and early gametophyte development in Anemia mexicana and Anemia phyllitidis. Plant Physiology 82(1): 230-235. DOI: https://doi.org/10.1104/pp.82.1.230
  21. Nester, J. E. and M. D. Schedlbauer. 1981. Gametophyte Development in Anemia mexicana Klotzsch. Botanical Gazette 142(2): 242-250. DOI: https://doi.org/10.1086/337219
  22. Nester, J. E., S. Veysey and R. C. Coolbaugh. 1987. Partial characterization of an antheridiogen of Anemia mexicana: Comparison with the antheridiogen of A. phyllitidis. Planta 170: 26-33. DOI: https://doi.org/10.1007/BF00392377
  23. Pinto, S. C., G. G. Leitão, A. Castellar, D. S. D’Elia, C. L. Lage, A. B. Henriques, J. Fernandes, G. S. Motta, H. R. Bizzo and S. G. Leitão. 2013. Chemical composition of the volatile fractions from wild and in vitro plants of Anemia tomentosa var. anthriscifolia (Pteridophyta). Journal of Essential Oil Research 25(3): 198-202. DOI: https://doi.org/10.1080/10412905.2012.755477
  24. PPG I. 2016. A community-based classification for extant ferns and lycophytes. Journal of Systematics and Evolution 54(6): 563-603. DOI: https://doi.org/10.1111/jse.12229
  25. Pray, T. R. 1971. The gametophyte of Anemia colimensis. American Journal of Botany 58(4): 323-328. DOI: https://doi.org/10.1002/j.1537-2197.1971.tb09980.x
  26. Raghavan, V. and C. S. Huckaby. 1980. A comparative study of cell division patterns during germination of spores of Anemia, Lygodium and Mohria (Schizaeaceae). American Journal of Botany 67(5): 653-663. DOI: https://doi.org/10.1002/j.1537-2197.1980.tb07696.x
  27. Raghavan, V. 1989. Developmental biology of fern gametophytes. Cambridge University Press. Cambridge, UK. Pp. 362.
  28. Ramos Giacosa, J. P. 2014. Abnormal spore morphology and wall ultrastructure in Anemia tomentosa var. anthriscifolia and A. tomentosa var. tomentosa (Anemiaceae). Plant Systematics and Evolution 300: 1571-1578. DOI: https://doi.org/10.1007/s00606-014-0983-2
  29. Ramos Giacosa, J. P., M. A. Morbelli and G. E. Giudice. 2012. Spore morphology and wall ultrastructure of Anemia Swartz species (Anemiaceae) from Argentina. Review of Palaeobotany and Palynology 174: 27-38. DOI: https://doi.org/10.1016/j.revpalbo.2012.02.004
  30. Reynolds, T. L. 1979. Apical dominance in Anemia phyllitidis gametophytes. American Fern Journal 69(3): 92-94. DOI: https://doi.org/10.2307/1546388
  31. Smith, A. R. and M. Kessler. 2017. Prodromus of a fern flora for Bolivia. XIII. Anemiaceae. Phytotaxa 329(1): 080-086. DOI: https://doi.org/10.11646/phytotaxa.329.1.5
  32. Smith, A. R., K. M. Pryer, E. Schuettpelz, P. Korall, H. Schneider and P. G. Wolf. 2006. A classification for extant ferns. Taxon 55(3): 705-731. DOI: https://doi.org/10.2307/25065646
  33. Takahashi, N., C. Kami, N. Morita and R. Imaichi. 2012. Comparative development of heavily asymmetric-cordate gametophytes of Anemia phyllitidis (Anemiaceae) focusing on meristem behavior. Journal of Plant Research 125: 371-380. DOI: https://doi.org/10.1007/s10265-011-0453-7
  34. Thiers, B. 2018 continuously updated. Index Herbariorum: A global directory of public herbaria and associated staff. New York Botanical Garden’s Virtual Herbarium. New York, USA. http://sweetgum.nybg.org/ih/ (consulted April, 2018).
  35. Tryon, A. F. and B. Lugardon. 1991. Spores of pteridophytes. Springer. New York, USA. Pp. 107-113.
  36. Von Aderkas, P. and E. G. Cutrer. 1983. Gametophyte plasticity and its bearing on sex expression in Todea barbara (L.) Moore. Botanical Gazette 144(4): 525-532. DOI: https://doi.org/10.1086/337406
  37. Yamane, H. 1998. Fern Antheridiogens. International Review of Cytology 184: 1-32. DOI: https://doi.org/10.1016/S0074-7696(08)62177-4