de Biología Tropical ISSN Impreso: 0034-7744 ISSN electrónico: 2215-2075

Arbuscular mycorrhizal, Mucoromycotina and dark septate fungi in ferns and lycophytes with distribution in Mexico: a global review

Luis Alberto Lara-Pérez, Ramón Zulueta-Rodríguez, Antonio Andrade-Torres



Ferns and lycophytes are a group of vascular plants of interest to understand the evolution of mycorrhizal interactions; their preservation is of relevance for their multiple ecological relations. The record of different taxonomic groups of fungi associated with ferns and lycophytes is fragmentary, and the criteria for it identification is inconsistent, which hinders the understanding and determination of mycorrhizal status. The aim of this study was to determine the percentage of the species of ferns and lycophytes with distribution in Mexico, and with information of fungal interactions. A checklist of the presence of arbuscular mycorrhizal fungi (AMF), Mucoromycotina and dark septate fungi (DSF) associated with ferns and lycophytes was integrated through an exhaustive global literature search. In this study, mycorrhizal species was considered by the presence of arbuscules to differentiate with hyphal, vesicular and coils colonization. The study gathered a checklist of mycorrhizal occurrences of 27 families, 61 genus and 137 species of ferns and lycophytes, which covers 13.4 % of the species, 91 % of the genus and 77 % of the families distributed in Mexico. The 78.1 % of the species showed colonization, 56.2 % by AMF, 29.9 % by DSF and 0.72 % by Mucoromycotina fungi. From the total of the species, the higher presences of colonization were in terrestrial, epiphytic, saxicolous, and aquatic plants with 76.6 %, 33.3 %, 20 %, and 6.3 %, respectively. The families of ferns and lycophytes with the higher number of species colonized were Pteridaceae, Polypodiaceae, Aspleniaceae and Dryopteridaceae. The present study showed the widespread associations of AMF and DSF in ferns and lycophytes of Mexico. It is urgently needed to include ferns and lycophytes in studies focused on endomycorhizal interactions, since only 28 species (28 %) were studied in Mexican ecosystems. The majority of studies were focused on sporophytic face (80 %). Nonetheless, to understand the role that plays the mycorrhiza in the establishment of ferns and lycophytes, it is necessary to include the gametophytic face in ecological, molecular and physiological experimental studies. This information is important to implement conservation strategies, because a considerable number of ferns and lycophytes species, depend on these mycorrhizal associations for their growth and survival.



arbuscular mycorrhizal fungi, dark septate fungi, Mucoromycotina, Lycophyta, Monilophyta, ferns.


Alarcón, A., Hernández-Cuevas, L. V., Ferrera-Cerrato, R., & Franco-Ramírez, A. (2012). Diversity and agricultural applications of arbuscular mycorrhizal fungi in Mexico. Journal of Fertilizers Pesticides, 3, 115 doi:104172/2155-62021000115

Andrade, A. C. S., Queiroz, M. H., Hermes, R. A. L., & Oliveira, V. L. (2000). Mycorrhizal status of some plants of the araucaria forest and the Atlantic rainforest in Santa Catarina, Brazil. Mycorrhiza, 10, 131-136.

Bagyaraj, D. J., Manjunath, A., & Patil, R. B. (1979). Occurrence of vesicular-arbuscular mycorrhizas in some tropical aquatic plants. Transactions of the British Mycological Society, 72, 164-167.

Bajwa, R., Yaqoob, A., & Javaid, A. (2001). Seasonal variation in VAM in wetland plants Pakisan. Journal of Biological Sciences, 4, 464-470.

Bautista-Cruz, A. A., Montaño, N. M., Camargo-Ricalde, S. L, & Pacheco, L. (2014). Hongos micorrizógenos arbusculares y nutrimentos del suelo asociados a cuatro especies de helechos en dos ecosistemas de Oaxaca, México. Revista Chapingo. Serie ciencias forestales y del ambiente, 20, 199-212.

Berch, S. M., & Kendrick, B. (1982). Vesicular-arbuscular mycorrhizae of southern Ontario ferns and ferns allies. Mycologia, 74, 769-776.

Berliner, R., & Torrey, J. G. (1989). Studies on mycorrhizal associations in Harvard Forest, Massachusetts. Canadian Journal of Botany, 67, 2245-2251.

Bidartondo, M. I., Read, D. J., Trappe, J. M., Merckx, V., Ligrone, R., & Duckett, J. G. (2011). The dawn of symbiosis between plants and fungi. Biology Letters, 7, 574-577.

Bonfante, P., & Genre, A. (2008). Plants and arbuscular mycorrhizal fungi: an evolutionary-developmental perspective. Trends in Plant Science, 13, 492-498.

Bonfante, P., & Selosse, M. A. (2010). A glimpse into the past of land plants and of their mycorrhizal affairs: from fossils to evo-devo. New Phytologist, 186, 267-270.

Boullard, B. (1979). Considérations sur les symbioses fongiques chez les Ptéridophytes. Syllogeus, 19, 1-58.

Brundrett, M. C. (2002). Coevolution of roots and mycorrhizas of land plants. New Phytologist, 154, 275-304.

Brundrett, M. C. (2009). Mycorrhizal associations and other means of nutrition of vascular plants: understanding the global diversity of host plants by resolving conflicting information and developing reliable means of diagnosis. Plant and Soil, 320, 37-77.

Campbell, D. H. (1908). Symbiosis in fern protallia. The American Naturalist, 42, 154-165.

Carrillo, R., Godoy, R., & Peredo, H. (1992). Simbiosis micorrícica en comunidades boscosas del Valle Central en el sur de Chile. Bosque, 13, 57-67.

Castillo, C. G., Borie, F., Godoy, R., Rubio, R., & Sieverding, E. (2006). Diversity of mycorrhizal plant species and arbuscular mycorrhizal fungi in evergreen forest, deciduous forest and grassland ecosystems of Southern Chile. Journal of Applied Botany and Food Quality, 80, 40-47.

Christenhusz, M. J. M., Zhang, X. C., & Schneider, H. (2011). A linear sequence of extant families and genera of lycophytes and ferns. Phytotaxa, 19, 7-54.

Cooke, J. C., & Lefor, M. W. (1998). The mycorrhizal status of selected plant species from Connecticut wetlands and transition zones. Restoration Ecology, 6, 214-222.

Cooper, K. M. (1976). A field survey of mycorrhizas in New Zealand ferns. New Zealand Journal of Botany, 14, 169-181.

de la Rosa-Mera, C., Ferrera-Cerrato, R., Alarcón, A., Sánchez-Colín, M. J., & Franco-Ramírez, A. (2012). Aislamiento de consorcios de hongos micorrícicos arbusculares de plantas medicinales y su efecto en el crecimiento de vinca (Catharanthus roseus). Revista Chilena de Historia Natural, 85, 187-198.

de Marins, J. F., Carrenho, R., & Thomaz, S. M. (2009). Occurrence and coexistence of arbuscular mycorrhizal fungi and dark septate fungi in aquatic macrophytes in a tropical river-floodplain system. Aquatic Botany, 91, 13-19.

Dhillion, S. S. (1993). Vesicular-arbuscular mycorrhizas of Equisetumspecies in Norway and the USA -occurrence and mycotrophy. Mycological Research, 97, 656-660.

Fernández, N., Messuti, M. I., & Fontenla, S. (2008). Arbuscular mycorrhizas and dark septate fungi in Lycopodium paniculatum (Lycopodiaceae) and Equisetum bogotense (Equisetaceae) in a Valdivian temperate forest of Patagonia, Argentina. American Fern Journal, 98, 117-127.

Fernández, N. V., Messuti, M. I., & Fontenla, S. B. (2013). Occurrence of arbuscular mycorrhizas and dark septate endophytes in pteridophytes from a patagonian rainforest, Argentina. Journal of Basic Microbiology, 53, 498-508.

Field, K. J., Leake, Jr., Tille, S., Allinson, K. E., Rimington, W. R., Bidartondo, M. I., Beerling, D. J., & Cameron, D. D. (2015). From mycohetereotrophy to mutualism: mycorrhizal specificity and functioning in Ophioglossum vulgatum sporophytes. New Phytologist, 205, 1492-1502.

Gemma, J. N., Koske, R. E., & Flynn, T. (1992). Mycorrhizae in Hawaiian pteridophytes: occurrence and evolutionary significance. American Journal of Botany, 79, 843-852.

Harley, J. L., & Harley, E. L. (1987). A check-list of mycorrhiza in the British flora. New Phytologist Supplement, 105, 1-100.

Hepden, P. M. (1960). Studies in vesicular-arbuscular endophytes II Endophytes in the Pteridophyta, with special references to leptosporangiate ferns. Transactions of the British Mycological Society, 43, 559-570.

Iqbal, S. H, Yousaf, M., & Younus, M. (1981). A field survey of mycorrhizal associations in ferns of Pakistan. New Phytologist, 87, 69-79.

James, T. Y., Kauff, F., Schoch, C. L., Matheny, P. B., Hofstetter, V., Cox, … & Vilgalys, R. (2006). Reconstructing the early evolution of fungi using a six-gene phylogeny. Nature, 443, 818-822.

Janos, D. P., & Sahley, C. T. (1995). Rodent dispersal of vesicular-arbuscular mycorrhizal fungi in amazonian Peru. Ecology, 76, 1852-1858.

Jumpponen, A. (2001). Dark septate endophyte -are they mycorrhizal. Mycorrhiza, 11, 207-211.

Kessler, M., Güdel, R., Salazar, L., Homeier, J., & Kluge, J. (2014). Impact of mycorrhization on the abundance, growth and leaf nutrient status of ferns along a tropical elevational gradient. Oecologia, 175, 887-900.

Kessler, M., Jonas, R., Cicuzza, D., Kluge, J., Piatek, K., Naks, P., & Lehnert, M. (2009). A survey of the mycorrhization of Southeast Asian ferns and lycophytes. Plant Biology, 12, 788-793.

Kessler, M., Jonas, R., Strasberg, D., & Lehnert, M. (2010). Mycorrhizal colonizations of ferns and lycophytes on the island of La Réunion in relation to nutrient availability. Basic and Applied Ecology, 11, 329-336.

Khade, S. W., & Rodrigues, B. F. (2002). Arbuscular mycorrhizal fungi associated with some pteridophytes from western ghat region of Goa. Tropical Ecology, 43, 251-256.

Kottke, I., Haug, I., Setaro, S., Suárez, J. P., Weiß, M., Preußing, M., Nebel, M., & Oberwinkler, F. (2008). Guilds of mycorrhizal fungi and their relation to trees, ericads, orchids and liverworts in a neotropical mountain rain forest. Basic and Applied Ecology, 9, 13-23.

Kovács, G. M., Balazs, T., & Penzes, Z. (2007). Molecular study of arbuscular mycorrhizal fungi colonizing the sporophyte of the eusporangiate rattlesnake fern (Botrychium virginianum, Ophioglossaceae). Mycorrhiza, 17, 597-605.

Kovács, G. M., Kottke, I., & Oberwinkler, F. (2003). Light and electron microscopic study on the mycorrhizae of sporophytes of Botrychium virginianum-arbuscular structure resembling fossil forms. Plant Biology, 5, 574-580.

Laferrière, J. L., & Koske, R. E. (1981). Occurrence of VA-mycorrhizas in some Rhode Island Pteridophytes. Transactions of the British Mycological Society, 76, 331-332.

Lara-Pérez, L. A. (2007). Efecto de los hongos micorrizógenos arbusculares en la supervivencia de gametofitos micropropagados de Cyathea fulva Pteridophyta: Cyatheaceae (Tesis de Licenciatura). Universidad Veracruzana, México.

Lara-Pérez, L. A., Noa-Carrazana, J. C., López-Landa, J., Hernández-González, S., Oros-Ortega I., & Andrade-Torres, A. (2014). Colonization and structure of arbuscular mycorrhizal fungi community in Alsophila firma (Cyatheales: Cyatheaceae) from a tropical montane cloud forest in Veracruz, México. Revista de Biología Tropical, 62, 1609-1623.

Lara-Pérez, L. A., Valdés-Baizabal, M. D., Noa-Carrazana, J. C., Zulueta-Rodríguez, R., Lara-Capistrán, L., & Andrade-Torres, A. (2015). Mycorrhizal associations of ferns and lycopods of central Veracruz, Mexico. Symbiosis, 65, 185-192.

Laursen, G. A., Treu, R., Seppelt, R. D., & Stephenson, S. L. (1997). Mycorrhizal assessment of vascular plants from subantarctic Macquarie Island. Arctic and Alpine Research, 29, 483-491.

Leake, J. R. (1994). The biology of mycoheterotrophic ('saprophitic') plants. New Phytologist, 127, 171-216.

Lee, J. K., Eom, A. H., Lee, S. S, & Lee, C. H. (2001). Mycorrhizal symbioses found in roots of fern and its relatives in Korea. Journal of Plant Biology, 44, 81-86.

Lehnert, M., Kottke, I., Setaro, S., Pazmino, L. F., Suárez, J. P., & Kessler M. (2009). Mycorrhizal associations in ferns from Southern Ecuador. American Fern Journal, 99, 292-306.

Lehnert, M., Krug, M., & Kessler, M. (2016). A review of symbiotic fungal endophytes in lycophytes and ferns-a global phylogenetic and ecological perspective. Symbiosis, 1-13. doi:10.1007/s13199-016-0436-5

Lesica, P., & Antibus, R. K. (1990). The occurrence of mycorrhizae in vascular epiphytes of two Costa Rican rain forests. Biotropica, 33, 250-258.

Leung, H. M., Ye, Z. H., & Wong, M. H. (2007). Survival strategies of plants associated with arbuscular mycorrhizal fungi on toxic mine tailings. Chemosphere, 66, 905-915.

Liu, Y., Zhu, Y. G., Chen, B. D., Christie, P., & Li, X. L. (2005). Influence of the arbuscular mycorrhizal fungus Glomus mosseae on uptake of arsenate by the As hyperaccumulator fern Pteris vittata L. Mycorrhiza, 15, 187-192.

Lloyd, R. M. (1974). Mating systems and genetic load in pioneer and non‐pioneer Hawaiian Pteridophyta. Botanical Journal of the Linnean Society, 69, 23-35.

Martínez, A. E., Chiocchio, V., Em, L. T., Rodríguez, M. A., & Godeas, A. M. (2012). Mycorrhizal association in gametophytes and sporophytes of fern Pteris vittata (Peridaceae) with Glomus intraradices. Revista de Biología Tropical, 60, 857-865.

McIlveen, W. D., & Cole, Jr., H. (1976). Spore dispersal of Endogonaceae by worms, ants, wasps, and birds. Canadian Journal of Botany, 54, 1486-1489.

Mehltreter, K. (2010). Interactions of ferns with fungi and animals. In K. Mehltreter, L. R. Walker, & J. M. Sharpe (Eds.), Fern Ecology (pp. 220-254). Cambridge: Cambridge University Press.

Merckx, V., & Bidartondo, M. I. (2008). Breakdown and delayed cospeciation in the arbuscular mycorrhizal mutualism. Proceedings of the Royal Society B: Biological Sciences, 275, 1029-1035.

Mickel, J. T., & Smith, A. R. (2004). The pteridophytes of Mexico. Memoirs of the New York Botanical Garden, 88, 1-1054.

Mishra, R. R., Sharma, G. D., & Gatphoh, A. R. (1980). Mycorrhizas in the ferns of North Eastern India. Proceedings of the Indian Academy of Science, 46, 546-551.

Montaño, N. M., Alarcón, A., Camargo-Ricalde, S. L., Hernández-Cuevas, L. V., Álvarez-Sánchez, J., González-Chávez, M. D. C. A., … & Ferrera-Cerrato, R. (2012). Research on arbuscular mycorrhizae in Mexico: an historical synthesis and future prospects. Symbiosis, 57, 111-126.

Moran, R. C. (2008). Diversity, biogeography and floristics. In: T. A. Ranker, & C. H. Haufler, (Eds.), Biology and evolution of ferns and lycophytes (pp. 367-394). Cambridge, UK: Cambridge University Press.

Moteetee, A., Duckett. J. G., & Russell, A. J. (1996). Mycorrhizas in the ferns of Lesotho. In J. M. Camus, M. Gibby, & R. J. Johns (Eds.), Pteridology in perspective (pp. 621-632). UK: Royal Botanic Gardens, Kew.

Muthukumar, T., & Prabha, K. (2012). Fungal associations in gametophytes and young sporophytic roots of the fern Nephrolepis exaltata. Acta Botanica Croatica, 71, 139-146.

Muthukumar, T., & Prabha, K. (2013). Arbuscular mycorrhizal and septate endophyte fungal associations in lycophytes and ferns of South India. Symbiosis, 59, 15-33

Muthukumar, T., Sathiyaraj, G., Priyadharsini, P., Uma, E., & Sathiyadash, K. (2014). Arbuscular mycorrhizal and dark septate endophyte fungal associations in ferns and lycophytes of Palni Hills, Western Ghats, southern India. Brazilian Journal of Botany, 37, 561-581.

Muthukumar, T., & Udaiyan, K. (2000). Arbuscular mycorrhizas of plants growing in the Western Ghats region, southern India. Mycorrhiza, 9, 297-313.

Muthuraja, R., Muthukumar, T., Sathiyadash, K., Uma, E., & Priyadharsini, P. (2014). Arbuscular mycorrhizal (AM) and dark septate endophyte (DSE) fungal association in lycophytes and ferns of the Kolli Hills Eastern Ghats, Southern India. American Fern Journal, 104, 67-102.

Nadarajah, P., & Nawawi, A. (1993). Mycorrhizal status of epiphytes in Malaysian oil palm plantations. Mycorrhiza, 4, 21-25.

Newman, E. I., & Reddell, P. (1987). The distribution of mycorhizas among families of vascular plants. New Phytologist, 106, 745-751.

Newsham, K. K. (2011). A meta-analysis of plant responses to dark septate root endophytes. New Phytologist, 190, 783-793.

Ogura-Tsujita, Y., Hirayama, Y., Sakoda, A., Suzuki, A., Ebihara, A., Morita, N., & Imaichi, R. (2015). Arbuscular mycorrhizal colonization in field-collected terrestrial cordate gametophytes of pre-polypod leptosporangiate ferns (Osmundaceae, Gleicheniaceae, Plagiogyriaceae, Cyatheaceae). Mycorrhiza, 26, 87-97.

Palmieri, M., & Swatzell, L. J. (2004). Mycorrhizal fungi associated with the fern Cheilanthes lanosa in southeast Missouri and southern Illinois. Northeastern Naturalist, 11, 57-66.

Pirozynski, K. A., & Malloch, D. W. (1975). The origin of land plants: a matter of mycotrophism. Biosystems, 6, 153-164.

Pryer, K. M., Schuettpelz, E., Wolf, P. G., Schneider, H., Smith, A. R., & Cranfill, R. (2004). Phylogeny and evolution of ferns monilophytes with a focus on the early leptosporangiate divergences. American Journal of Botany, 91, 1582-1598.

Radhika, K. P. & Rodrigues, B. F. (2007). Arbuscular mycorrhizae in association with aquatic and marshy plant species in Goa, India. Aquatic Botany, 86, 291-294.

Raghavan, V. (1992). Germination of fern spores. American Scientist, 80, 176-185.

Redecker, D., Kodner, R., & Graham, L. E. (2000). Glomalean fungi from the Ordovician. Science, 289, 1920-1921.

Reyes-Jaramillo, I., Camargo-Ricalde, S. L., & Aquiahuatl-Ramos, M. A. (2008). Mycorrhizal-like interaction between gametophytes and young sporophytes of the fern Dryopteris muenchii (Filicales) and its fungal endophyte. Revista de Biología Tropical, 56, 1101-1107.

Rimington, W. R., Pressel, S., Duckett, J. G., & Bidartondo, M. I. (2015). Fungal associations of basal vascular plants: reopening a closed book? New Phytologist, 205, 1394-1398.

Rzedowski, J. (1978). La Vegetación de México. Ciudad de México: Limusa.

Santhoshkumar, S., & Nagarajan, N. (2014). AM fungal association in the rhizosphere soil of some pteridophytic plant species in Valparai Hills, Western Ghats of Tamilnadu, India. International Journal of Life Sciences, 2, 201-206.

Schmid, E., & Oberwinkler, F. (1993). Mycorrhiza-like interaction between the achlorophyllous gametophyte of Lycopodium clavatum L. and its fungal endophyte studied by light and electron microscopy. New Phytologist, 124, 69-81.

Schmid, E., & Oberwinkler, F. (1996). Light and electron microscopy of a distinctive VA mycorrhiza in mature sporophytes of Ophioglossum reticulatum. Mycological Research, 100, 843-849.

Schmid, E., Oberwinkler, F., & Gómez, L. D. (1995). Light and electron microscopy of a host-fungus interaction in the roots of some epiphytic ferns from Costa Rica. Canadian Journal of Botany, 73, 991-996.

Schneider, J., Oliveira, L. M., Guilherme, L. R. G., Stürmer, S. L., & Soares, C. R. F. S. (2012). Espécies tropicais de pteridófitas em associação com fungos micorrízicos arbusculares em solo contaminado com arsênio. Química Nova, 35, 709-714.

Schüßler, A., & Walker, C. (2010). The Glomeromycota: a species list with new families and new genera Gloucester. England: The Royal Botanic Garden Edinburgh/The Royal Botanic Garden Kew/Botanische Staatssammlung Munich/Oregon State University.

Seerangan, K., & Thangavelu, M. (2014). Arbuscular mycorrhizal and dark septate endophyte fungal associations in South Indian aquatic and wetland macrophytes. Journal of Botany, 1-14 doiorg/101155/2014/173125

Smith, A. R., Pryer, K. M., Schuettpelz, E., Korall, P., Schneider. H., & Wolf, P. G. (2006). A classification for extant ferns. Taxon, 55, 705-731.

Smith, S. E., & Read, D. J. (2008). Mycorrhizal Symbiosis New York: Academic Press.

Strullu-Derrien, C., Kenrick, P., Pressel, S., Duckett, J. G., Rioult, J. P., & Strullu, D. G. (2014). Fungal associations in Horneophyton ligneri from the Rhynie Chert c 407 million year old closely resemble those in extant lower land plants: novel insights into ancestral plant-fungus symbioses. New Phytologist, 203, 964-979.

Tejero-Díez, J. D., Torres-Díaz, A., Mickel, J., Mehltreter, K., & Krömer, T. (2009). Helechos y licófitas. In G. Ceballos, R. List, G. Garduño, R. López-Cano, M. J. Muñozcano-Quintanar, E. Collado, & J. E. San Román (Eds.), La diversidad biológica del Estado de México: Estudio de Estado México (pp. 167-176). México: Gobierno del Estado de México y Comisión Nacional para el Conocimiento y Uso de la Biodiversidad.

Treu, R., Laursen, G. A., Stephenson, S. L., Landolt, J. C., & Densmore, R. (1996). Mycorrhizae from Denali National Park and Preserve, Alaska. Mycorrhiza, 6, 21-29.

Trotta, A., Falaschi, P., Cornara, L., Minganti, V., Fusconi, A., Drava, G., & Berta, G. (2006). Arbuscular mycorrhizae increase the arsenic translocation factor in the As hyperaccumulating fern Pteris vittata L. Chemosphere, 65, 74-81.

Tryon, R. M., & Tryon, A. F. (1982). Ferns and allied plants with special reference to tropical America. New York: Springer-Verlag.

Turnau, K., Anielska, T., & Jurkiewicz, A. (2005). Mycothallic/mycorrhizal symbiosis of chlorophyllous gametophytes and sporophytes of a fern, Pellaea viridis (Forsk) Prantl. (Pellaeaceae, Pteridales). Mycorrhiza, 15, 121-128.

van der Heijden, M. G. A., Martin, F., Selosse, M. A., & Sanders, I. R. (2015). Mycorrhizal ecology and evolution: the past, the present, and the future. New Phytologist, 205, 1406-1423.

Wang, B., & Qiu, Y. L. (2006). Phylogenetic distribution and evolution of mycorrhizas in land plants. Mycorrhiza, 16, 299-363.

Wang, Y., Zhan, M., Zhu, H., Guo, S., Wang, W., & Xue, B. (2012). Distribution and accumulation of metals in soils and plant from a lead-zinc mineland in Guangxi, South China. Bulletin of Environmental Contamination and Toxicology, 88, 198-203.

West, B., Brandt, J., Holstien, K., Hill, A., & Hill, M. (2009). Fern-associated arbuscular mycorrhizal fungi are represented by multiple Glomus spp: do environmental factors influence partner identity? Mycorrhiza, 19, 295-304.

Winther, J. L., & Friedman, W. E. (2008). Arbuscular mycorrhizal associations in Lycopodiaceae. New Phytologist, 177, 790-801.

Winther, J. L., & Friedman, W. E. (2009). Phylogenetic affinity of arbuscular mycorrhizal symbionts in Psilotum nudum. Journal of Plant Research, 122, 485-496.

Yi, W., Fu, S., Cao, H., & Ding, M. (1994). Vesicular-arbuscular mycorrhizal associations in three forest types in the Dinghushan Biosphere Reserve, subtropical China. Proceedings of Australian Center for international Agricultural Research, 62, 41-46.

Zhang, Y., Guo, L. D, & Liu, R. J. (2004). Arbuscular mycorrhizal fungi associated with common pteridophytes in Dujiangyan, southwest China. Mycorrhiza, 14, 25-30.

Zhao, Z. W. (2000). The arbuscular mycorrhizas of pteridophytes in Yunnan, southwestern China: evolutionary interpretations. Mycorrhiza, 10, 145-149.

Zubek, S., Piatek, K., Naks, P., Heise, W., Wayda, M., & Mleczko, P. (2010). Fungal root endophyte colonization of fern and lycophyte species from the Celaque National Park. American Fern Journal, 100, 126-136.


  • There are currently no refbacks.

© 2017 Universidad de Costa Rica. Para ver más detalles sobre la distribución de los artículos en este sitio visite el aviso legal. Este sitio es desarrollado por UCRIndex y Open Journal Systems.