Morphological and molecular characterization of native ectomycorrhizae from forest soil inoculated in Pinus montezumae under greenhouse conditions
DOI:
https://doi.org/10.22201/ib.20078706e.2026.97.5787Keywords:
Ectomycorrhizal fungi, ITS, Morphotypes, Piedra Canteada, TlaxcalaAbstract
Ectomycorrhizal fungi represent an important component of the fungal diversity in Mexico’s temperate forests. However, studies characterizing ectomycorrhizae without the use of fruiting bodies are scarce. Therefore, in this study, 3 ectomycorrhizal morphotypes present in the roots of Pinus montezumae, inoculated with forest soil containing native ectomycorrhizal mycelium, were characterized from the Piedra Canteada Protected Natural Area, Tlaxcala. Detailed descriptions of the morphotypes, designated as yellow, white, and black, were provided. The ITS2 region of rDNA was subsequently amplified by PCR using the ITS5 and ITS4 primers. Through a combined morphological and molecular approach, the native ectomycorrhizal taxa colonizing the root system of P. montezumae were identified. The yellow and white morphotypes were identified as Tylospora sp. and Aureoboletus sp., respectively. Both are new records of ectomycorrhizal fungi in Mexico. Morphological characterization was sufficient for the identification of the black morphotype as Cenococcum geophilum. These results contribute to the knowledge of the native fungal diversity of the forest soil associated with P. montezumae, whose presence and ecological function favor the sustainability of this protected area.
References
Aceves-Rangel, L. D., Méndez-González, J., García-Aranda, M. A. y Nájera-Luna, J. A. (2018). Distribución potencial de 20 especies de pinos en México. Agrociencias, 52, 1043–1057.
Agerer, R., Hartmann, A., Pristch, K., Raidl, S., Schloter, M. y Verma, R. (2012). Plants and their ectomycorrhizosphere: cost and benefit of symbiotic soil organisms. En R. Matyssek, H. Schnyder, W. Oßwald, D. Ernst, J. C. Munch y H. Pretzsch (Eds.), Growth and defence in plants (pp. 213–242). Heidelberg: Springer-Verlag. https://doi.org/10.1007/978-3-642-30645-7_10
Agerer, R. y Raidl, S. (2004). Distance-related semi-quantitative estimation of the extramatrical ectomycorrhizal mycelia of Cortinarius obtusus and Tylospora asterophora. Mycological Progress, 3, 57–64. https://doi.org/10.1007/s11557-006-0077-9
Agerer, R. y Rambold, G. (s.f.). DEEMY. An information system for characterization and determination of ectomycorrhizae. Recuperado el 03 de septiembre del 2025 de: http://www.deemy.de/
Almaraz-Llamas, A., Pérez-Moreno, J., Torres-Aquino, M., Carcaño-Montiel, M. G. y Hernández Ríos, I. (2019). Cenococcum geophilum y su asociación con Pinus maximartinezii árbol nativo de México en peligro de extinción. Scientia Fungorum, 49, 1–15. https://doi.org/10.33885/sf.2019.49.1222
Altschul, S. F., Madden, T. L., Schäffer, A. A., Zhang, J., Zhang, Z., Miller, W. et al. (1997). Gapped BLAST and PSI-BLAST: a new generation of protein database search programs. Nucleic Acids Research, 25, 3389–3402. https://doi.org/10.1093/nar/25.17.3389
Arteaga-León, C., Pérez-Moreno, J., Espinosa-Victoria, D., Almaraz-Suárez, J. J., Silva-Rojas, H. y Delgado-Alvarado, A. (2018). La inoculación ectomicorrízica con hongos comestibles incrementa el crecimiento vegetal y el contenido nutrimental de Pinus ayacahuite. Revista Mexicana de Biodiversidad, 89, 1089–1099. https://doi.org/10.22201/ib.20078706e.2018.4.2235
Ayala-Vásquez, O., Martínez-Reyes, M., Pérez-Moreno, J., Martínez-González, C. R., Pinzón, J. P., de la Fuente, J. I. et al. (2023). Five new species of Aureoboletus and Chalciporus (Boletaceae, Boletales) and their ethnomycological aspects. Journal of Fungi, 9, 1041. https://doi.org/10.3390/jof9101041
Baeza-Guzman, Y., Camargo-Ricalde, S. L., Trejo Aguilar, D. y Montano, N. M. (2023). Fungal and bacterial communities in a forest relict of Pinus pseudostrobus var. coatepecensis. iForest Biogeosciences and Forestry, 16, 299–306. https://doi.org/10.3832/ifor4284-016
Baeza-Guzmán, Y., Medel-Ortiz, R. y Garibay-Orijel, R. (2017). Caracterización morfológica y genética de los hongos ectomicorrízicos asociados a bosques de Pinus hartwegii en el Parque Nacional Cofre de Perote, Veracruz. Revista Mexicana de Biodiversidad, 88, 41–48. https://doi.org/10.1016/j.rmb.2017.01.027
Barragán-Soriano, J. L., Pérez-Moreno, J., Almaraz-Suárez, J. J., Carcaño-Montiel, M. G. y Medrano-Ortiz, K. I. (2018). Inoculation with an edible ectomycorrhizal fungus and bacteria increases growth and improves the physiological quality of Pinus montezumae Lamb. Revista Chapingo serie Ciencias Forestales y del Ambiente, 24, 3–16. https://doi.org/10.5154/r.rchscfa.2017.01.010
Bickford, D., Lohman, D. J., Sodhi, N. S., Ng, P. K., Meier, R., Winker, K. et al. (2007). Cryptic species as a window on diversity and conservation. Trends in Ecology and Evolution, 22, 148–155. https://doi.org/10.1016/j.tree.2006.11.004
Binder, M. y Hibbett, D. S. (2006). Molecular systematics and biological diversification of Boletales. Mycologia, 98, 971–983. https://doi.org/10.1080/15572536.2006.11832626
Bzdyk, R. M., Sikora, K., Studnicki, M. y Aleksandrowicz-Trzcińska, M. (2022). Communities of mycorrhizal fungi among seedlings of scots pine (Pinus sylvestris L.) growing on a clearcut in microsites generated by different site-preparation methods. Forests, 13, 353. https://doi.org/10.3390/f13020353
Cairney, J. W. G. y Burke, R. M. (1994). Fungal enzymes degrading plant cell walls: their possible significance in the ectomycorrhizal symbiosis. Mycological Research, 98, 1345–1356. https://doi.org/10.1016/S0953-7562(09)81062-9
Chávez-Aguilar, G., Burrola-Aguilar, C., González-Morales, A. y Pérez-Suárez, M. (2020). Almacén de carbono orgánico del suelo y abundancia ectomicorrízica bajo dos especies de coníferas en el Nevado de Toluca, México. Agro Productividad, 13, 75–82. https://doi.org/10.32854/agrop.vi0.1621
Clasen, B. E., Silveira, A. D. O., Baldoni, D. B., Montagner, D. F., Jacques, R. J. S. y Antoniolli, Z. I. (2018). Characterization of Ectomycorrhizal species through molecular biology tools and morphotyping. Scientia Agricola, 75, 246–254. https://doi.org/10.1590/1678-992X-2016-0419
Dentinger, B. T. M., Ammirati, J. F., Both, E. E., Desjardin, D. E., Halling, R. E., Henkel, T. W. et al. (2010). Molecular phylogenetics of porcini mushrooms (Boletus section Boletus). Molecular Phylogenetics and Evolution, 57, 1276–1292. https://doi.org/10.1016/j.ympev.2010.10.004
Douhan, G. W., Huryn, K. L. y Douhan, L. I. (2007). Significant diversity and potential problems associated with inferring population structure within the Cenococcum geophilum species complex. Mycologia, 99, 812–819. https://doi.org/10.1080/15572536.2007.11832513
Eckhart, L., Bach, J., Ban, J. y Tschachler, E. (2000). Melanin binds reversibly to thermostable DNA polymerase and inhibits its activity. Biochemical and Biophysical Research Communications, 271, 726–730. https://doi.org/10.1006/bbrc.2000.2716
Farid, A., Bessette, A. E., Bessette, A. R., Bolin, J. A., Kudzma, L. V., Franck, A. R. et al. (2021). Investigations in the boletes (Boletaceae) of southeastern USA: four novel species and three novel combinations. Mycosphere, 12, 1038–1076. https://doi.org/10.5943/mycosphere/12/1/12
Fernández, C. W. y Koide, R. T. (2013). The function of melanin in the ectomycorrhizal fungus Cenoccocum geophilum under water stress. Fungal Ecology, 6, 479–486. https://doi.org/10.1016/j.funeco.2013.08.004
Flores, R. (2020). Luciérnagas iluminan la conservación de un bosque en México. Mongabay. Recuperado el 03 de septiembre del 2025 de https://es.mongabay.com/2020/08/luciernagas-iluminan-la-conservacion-de-un-bosque-en-mexico/
Flores-García, A., Pineda-Ojeda, T. y Flores-Ayala, E. (2019). Potencial de reforestación de seis especies de pino para la restauración de zonas degradadas. Revista Mexicana de Ciencias Forestales, 10, 171–179. https://doi.org/10.29298/rmcf.v10i55.604
Garay-Serrano, E., Ortega–Larrocea, M. P., Reverchon, F. y Suárez-Quijada, I. (2018). Persistence of ecto- and ectendomycorrhizal fungi associated with Pinus montezumae in experimental microcosms. Symbiosis, 74, 67–78. https://doi.org/10.1007/s13199-017-0496-1
Gardes, M. y Bruns, T. D. (1993). ITS primers with enhanced specificity for basidiomycetes‐application to the identification of mycorrhizae and rusts. Molecular Ecology, 2, 113–118. https://doi.org/10.1111/j.1365-294X.1993.tb00005.x
Garibay-Orijel, R., Morales-Marañon, E., Domínguez-Gutiérrez, M. y Flores-García, A. (2013). Caracterización morfológica y genética de las ectomicorrizas formadas entre Pinus montezumae y los hongos presentes en los bancos de esporas en la Faja Volcánica Transmexicana. Revista Mexicana de Biodiversidad, 84, 153–169. https://doi.org/10.7550/rmb.29839
González-Ocampo, H. A., Cortés-Calva, P., Íñiguez-Dávalos, L. I. y Ortega-Rubio, A. (2014). Las áreas naturales protegidas de México. Investigación y Ciencia de la Universidad Autónoma de Aguascalientes, 60, 7–15.
Goodman, D. M., Trofymow, J. A. y Thomson, A. J. (2000). Developing an online database of descriptions of ectomycorrhizae. Journal of Ecosystems and Management, 1, 1–8. https://doi.org/10.22230/jem.2001v1n1a211
Guerra-De la Cruz, V., Carrillo-Anzures, F. y Acosta-Mireles, M. (2008). El aprovechamiento de los recursos forestales maderables. En V. Guerra-de la Cruz y C. Mallén-Rivera (Comp.), Tlaxcala, sus recursos forestales: conservación, aprovechamiento y bases para su manejo sustentable. Libro Técnico Núm. 4 (pp. 30–43). Ciudad de México: INIFAP-Cenid-Comef.
Guerra-De la Cruz, V., Islas-Gutiérrez, F., Flores-Ayala, E., Acosta-Mireles, M., Buendía-Rodríguez, E., Carrillo-Anzures, F. et al. (2019). Modelos locales altura-diámetro para Pinus montezumae Lamb. y Pinus teocote Schiede ex Schltdl. en Nanacamilpa, Tlaxcala. Revista Mexicana de Ciencias Forestales, 10, 133–156. https://doi.org/10.29298/rmcf.v10i51.407
Guevara-Rojas, M. (2019). Inoculación con dosis de tierra micorrizada y Suillus luteus comercial en la producción de plantones de Pinus radiata D. Don, en Paquecc (2510 m snm), Huanta – Ayacucho (Tesis -Ingeniería). Facultad de Ciencias Agrarias, Universidad Nacional De San Cristóbal de Huamanga. Ayacucho, Perú.
Gutiérrez-Arancibia, G. G. (2019). Tipos y niveles de inóculos de micorrizas en la producción de dos especies de pinos en tubetes, Vivero Forestal Huamanga 2761 msnm, Ayacucho (Tesis -Ingeniería). Facultad de Ciencias Agrarias, Universidad Nacional De San Cristóbal de Huamanga. Ayacucho, Perú.
Guzmán, G. (2009). La diversidad de hongos en México. Ciencias, 39, 52–57.
He, X. L., Li, Q., Peng, W. H., Zhou, J., Cao, X. L, Wang, D. et al. (2017). Intra- and inter-isolate variation of ribosomal and protein-coding genes in Pleurotus: implications for molecular identification and phylogeny on fungal groups. Bmc Microbiology, 17, 139. https://doi.org/10.1186/s12866-017-1046-y
Herrera, M., Yu, F. Q., Ramos-Rendón, D., Martínez-Reyes, M., Hernández-Santiago, F., Chater, C. C. et al. (2022). Morphoanatomical and phylogenetic characterization of the ectomycorrhiza between Laccaria squarrosa with Pinus pseudostrobus and its relevance for reforestation programs. Botanical Sciences, 100, 397–411. https://doi.org/10.17129/botsci.2830
Herrera-Hernández, R., López-Upton, J., Muñoz-Gutiérrez, L. y Ramírez-Herrera, C. (2024). Potencial de producción de semilla de Pinus montezumae Lambert en el centro de México. Ciência Florestal, 34, e71911. https://doi.org/10.5902/1980509871911
Horton, T. R. (2002). Molecular approaches to ectomycorrhizal diversity studies: variation in ITS at a local scale. Plant and Soil, 244, 29–39. https://doi.org/10.1023/A:1020268020563
Horton, T. R. y Bruns, T. D. (2001). The molecular revolution in ectomycorrhizal ecology: peeking into the black‐box. Molecular Ecology, 10, 1855–1871. https://doi.org/10.1046/j.0962-1083.2001.01333.x
Hu, Q., Liu, Y., Yi, S. y Huang, D. (2015). A comparison of four methods for PCR inhibitor removal. Forensic Science International: Genetics, 16, 94–97. https://doi.org/10.1016/j.fsigen.2014.12.001
Huang, J., Han, Q. y Li, J. (2018). Soil propagule bank of ectomycorrhizal fungi associated with Masson pine (Pinus massoniana) grown in a manganese mine wasteland. Plos One, 13, e0198628. https://doi.org/10.1371/journal.pone.0198628
INEGI. (2021a). Aspectos geográficos Tlaxcala. Instituto Nacional de Estadística y Geografía. México. Recuperado el 25 de agosto del 2025 de https://www.inegi.org.mx/contenidos/app/areasgeograficas/resumen/resumen_29.pdf
INEGI (Instituto Nacional de Estadística y Geografía). (2021b). Diccionario de datos de climas. Instituto Nacional de Estadística y Geografía. México. Recuperado el 25 de agosto del 2025 de https://www.inegi.org.mx/contenidos/productos/prod_serv/contenidos/espanol/bvinegi/productos/nueva_estruc/702825199258.pdf
Janowski, D. y Leski, T. (2023). Methods for identifying and measuring the diversity of ectomycorrhizal fungi. Forestry: An International Journal of Forest Research, 96, 639–652. https://doi.org/10.1093/forestry/cpad017
Karlsen-Ayala, E., Smith, M. E., Askey, B. C. y Gazis, R. (2022). Native ectomycorrhizal fungi from the endangered pine rocklands are superior symbionts to commercial inoculum for slash pine seedlings. Mycorrhiza, 32, 465–480. https://doi.org/10.1007/s00572-022-01092-3
Kyaschenko, J., Clemmensen, K. E., Hagenbo, A., Karltun, E. y Lindahl, B. D. (2017). Shift in fungal communities and associated enzyme activities along an age gradient of managed Pinus sylvestris stands. International Society for Microbial Ecology Journal, 11, 863–874. https://doi.org/10.1038/ismej.2016.184
Lücking, R., Aime, M. C., Robbertse, B., Miller, A. N., Ariyawansa, H. A., Aoki, T. et al. (2020). Unambiguous identification of fungi: Where do we stand and how accurate and precise is fungal DNA barcoding?. Ima Fungus, 11, 14. https://doi.org/10.1186/s43008-020-00033-z
Manzanilla-Quiñones, U., Delgado-Valerio, P., Hernández-Ramos, J., Molina-Sánchez, A., García-Magaña J. J. y Rocha-Granados, M. C. (2019). Similaridad del nicho ecológico de Pinus montezumae y P. pseudostrobus (Pinaceae) en México: implicaciones para la selección de áreas productoras de semillas y de conservación. Acta Botanica Mexicana, 126, e1398. https://doi.org/10.21829/abm126.2019.1398
Martínez-González, C. R., Ramírez-Mendoza, R., Jiménez-Ramírez, J., Gallegos-Vázquez, C. y Luna-Vega, I. (2017). Improved method for genomic DNA extraction for Opuntia Mill. (Cactaceae). Plant Methods, 13, 82. https://doi.org/10.1186/s13007-017-0234-y
Massicotte, H. B., Trappe, J. M., Peterson, R. L. y Melville, L. H. (1992) Studies on Cenococcum geophilum. II. Sclerotium morphology, germination, and formation in pure culture and growth pouches. Canadian Journal of Botany, 70, 125–132. https://doi.org/10.1139/b92-017
Netherway, T. y Bahram, M. (2024). Melanized root-associated fungi: key players in plant–soil systems. Trends in Microbiology, 12, 1190–1199. https://doi.org/10.1016/j.tim.2024.06.006
Obase, K., Douhan, G. W., Matsuda, Y. y Smith, M. E. (2016). Revisiting phylogenetic diversity and cryptic species of Cenococcum geophilum sensu lato. Mycorrhiza, 26, 529–540. https://doi.org/10.1007/s00572-016-0690-7
Obase, K., Douhan, G. W., Matsuda, Y. y Smith, M. E. (2017). Progress and challenges in understanding the biology, diversity, and biogeography of Cenococcum geophilum. En L. Tedersoo (Ed.), Biogeography of mycorrhizal symbiosis, Vol 230 (pp. 299–317). Cham: Springer. https://doi.org/10.1007/978-3-319-56363-3_14
Obase K., Lee, J. K., Lee, S. Y. y Chun, K. W. (2011). Diversity and community structure of ectomycorrhizal fungi in Pinus thunbergii coastal forests in the eastern region of Korea. Mycoscience, 52, 383–391. https://doi.org/10.1007/S10267-011-0123-6
Palfner, G., Casanova-Katny, M. A. y Read, D. J. (2005). The mycorrhizal community in a forest chronosequence of Sitka spruce [Picea sitchensis (Bong.) Carr.] in Northern England. Mycorrhiza, 15, 571–579. https://doi.org/10.1007/s00572-005-0364-3
Pérez-Alavez, Y., Rodríguez-Ortiz, G., Santiago-García, W., Martin, M. P., Enríquez-del Valle, J. R. y Aguilar-Pinacho, I. J. (2025). Efectos de aclareos secuenciales en rodal de Pinus pseudostrobus Lindl. Ecosistemas y Recursos Agropecuarios, 12, e4006.
Pérez-Moreno, J., Fuentes-García, O., Martínez-Reyes, M., Martínez-González, C. R., Lagunes-Reyes, M., Díaz-Aguilar, I. et al. (2025). The genus Xerocomellus (Boletales, Boletaceae) in Mexico: a new species, a new record, and notes on its biocultural importance. Phytotaxa, 689, 1–14. https://doi.org/10.11646/phytotaxa.689.1.1
Pérez-Moreno, J. y Martínez-Reyes, M. (2014). Edible ectomycorrhizal mushrooms: biofactories for sustainable development. En R. Guevara-Gonzalez, I. Torres-Pacheco (Eds.), Biosystems engineering: biofactories for food production in the Century XXI (pp. 151–233). Cham: Springer. https://doi.org/10.1007/978-3-319-03880-3_6
Pérez-Moreno, J., Martínez-Reyes, M., Hernández-Santiago, F. y Ortiz-Lopez, I. (2020). Climate change, biotechnology, and mexican neotropical edible ectomycorrhizal mushrooms. En J. Pérez-Moreno, A. Guerin-Laguette, R. Flores-Arzú, F. Q. Yu (Eds.), Mushrooms, humans and nature in a changing world (pp. 61–99). Cham: Springer. https://doi.org/10.1007/978-3-030-37378-8_3
Pérez-Moreno, J. y Read, D. J. (2000). Mobilization and transfer of nutrients from litter to tree seedlings via the vegetative mycelium of ectomycorrhizal plants. New Phytologist, 145, 301–309. https://doi.org/10.1046/j.1469-8137.2000.00569.x
Ramírez-Miguel, A. A., Hernández-Díaz, A. F., Valenzuela-Encinas, C., Garibay-Orijel, R. y Truong, C. (2021). Hongos ectomicorrízicos asociados a plantas jóvenes de Pinus patula y Quercus crassifolia en plantaciones del sistema matarrasa de la Sierra Juárez de Oaxaca, México. Scientia Fungorum, 51, e1289. https://doi.org/10.33885/sf.2021.51.1289
Rentería-Chávez, M. C., Pérez-Moreno, J., Cetina-Alcalá, V. M., Ferrera-Cerrato, R. y Xoconostle-Cázares, B. (2017). Transferencia de nutrientes y crecimiento de Pinus greggii Engelm. inoculado con hongos comestibles ectomicorrícicos en dos sustratos. Revista Argentina de Microbiología, 49, 93–104. https://doi.org/10.1016/j.ram.2016.06.004
Reverchon, F., Ortega-Larrocea, M. P., Bonilla-Rosso, G. y Pérez-Moreno, J. (2012). Structure and species composition of ectomycorrhizal fungal communities colonizing seedlings and adult trees of Pinus montezumae in Mexican neotropical forests. Fems Microbiology Ecology, 80, 479–487. https://doi.org/10.1111/j.1574-6941.2012.01314.x
Reverchon, F., Ortega-Larrocea, M. P. y Pérez-Moreno, J. (2014). Structure and diversity of ectomycorrhizal resistant propagules in Pinus montezumae neotropical forests and implications for seedling establishment. Mycoscience, 56, 214–223. https://doi.org/10.1016/j.myc.2014.06.005
Robles-Villanueva, F. A., Rodríguez-Trejo, D. A. y Villanueva-Morales, A. (2017). Calidad de planta y supervivencia en reforestación de Pinus montezumae Lamb. Revista Mexicana de Ciencias Forestales, 8, 55–76. https://doi.org/10.29298/rmcf.v8i42.19
Rodríguez-Gómez Tagle, G., Vargas-Hernández, J. J., López-Upton, J. y Pérez-Moreno, J. (2024). Diversidad de morfotipos de hongos ectomicorrizógenos y adaptación al hospedero en poblaciones contrastantes de Pinus greggii var. australis (Pinaceae). Acta Botanica Mexicana, 24, e2151. https://doi.org/10.21829/abm131.2024.2151
Rodríguez-Gutiérrez, I., Garibay-Orijel, R., Santiago-Morales, B. y Lindig-Cisneros, R. (2020). Comparación entre las abundancias de esporomas y ectomicorrizas del género Laccaria en Ixtlán de Juárez, Oaxaca. Revista Mexicana de Biodiversidad, 91, e913340. https://doi.org/10.22201/ib.20078706e.2020.91.3340
Rodríguez-Tovar, A., Xoconostle-Cásarez, B. y Valdés, M. (2004). Ecología molecular de los hongos ectomicorrízicos. Revista Fitotecnia Mexicana, 27, 267–278. https://doi.org/10.35196/rfm.2004.3.267
Rosenthal, L. M., Larsson, K. H., Branco, S., Chung, J. A., Glassman, S. I., Liao, H. L. et al. (2017). Survey of corticioid fungi in North American pinaceous forests reveals hyperdiversity, underpopulated sequence databases, and species that are potentially ectomycorrhizal. Mycologia, 109, 115–127. https://doi.org/10.1080/00275514.2017.1281677
Rudawska, M., Leski, T. y Stasińska, M. (2011). Species and functional diversity of ectomycorrhizal fungal communities on Scots pine (Pinus sylvestris L.) trees on three different sites. Annals of Forest Science, 68, 5–15. https://doi.org/10.1007/s13595-010-0002-x
Sakakibara, S. M., Jones, M. D., Gillespie, M., Hagerman, S. M., Forrest, M. E., Simard, S. W. et al. (2002). A comparison of ectomycorrhiza identification based on morphotyping and PCR-RFLP analysis. Mycological Research, 106, 868–878. https://doi.org/10.1017/S0953756202006263
Semarnat (Secretaría de Medio Ambiente y Recursos Naturales). (2009). Restauración de ecosistemas forestales: guía básica para comunicadores. Ciudad de México: Secretaría de Medio Ambiente y Recursos Naturales-Comisión Nacional Forestal.
Simard, S. W., Jones, M. D. y Durall, D. M. (2003). Carbon and nutrient fluxes within and between mycorrhizal plants. En M. G. A. van der Heijden y I. R. Sanders (Eds.), Mycorrhizal ecology, Vol. 157 (pp. 33–74). Heidelberg: Springer. https://doi.org/10.1007/978-3-540-38364-2_2
Smith, M. E., Douhan, G. W. y Rizzo, D. M. (2007). Intra-specific and intra-sporocarp ITS variation of ectomycorrhizal fungi as assessed by rDNA sequencing of sporocarps and pooled ectomycorrhizal roots from a Quercus woodland. Mycorrhiza, 18, 15–22. https://doi.org/10.1007/s00572-007-0148-z
Smith, S. E. y Read, D. J. (2008). Mycorrhizal symbiosis. Nueva York: Academic Press.
Soto-Gil, A. L., Velázquez-Martínez, A., Pérez-Moreno, J., Fierros-González, A. M. y Martínez-Reyes, M. (2022). Morfotipos ectomicorrícicos en retención estructural variable de Pinus patula Schltdl et Cham. Madera y Bosques, 28, e2822388. https://doi.org/10.21829/myb.2022.2822388
Spatafora, J. W., Owensby, C. A., Douhan, G. W., Boehm, E. W. A. y Schoch, C. L. (2012). Phylogenetic placement of the ectomycorrhizal genus Cenococcum in Gloniaceae (Dothideomycetes). Mycologia, 104, 758–765. https://doi.org/10.3852/11-233
Taylor, A. F. S. y Alexander, I. J. (1991). Ectomycorrhizal synthesis with Tylospora fibrillosa, a member of the Corticiaceae. Mycological Research, 95, 381–384. https://doi.org/10.1016/S0953-7562(09)81256-2
Tedersoo, L., May, T. W. y Smith, M. E. (2010) Ectomycorrhizal lifestyle in fungi: global diversity, distribution, and evolution of phylogenetic lineages. Mycorrhiza, 20, 217–263. https://doi.org/10.1007/s00572-009-0274-x
Tedersoo, L. y Smith, M. E. (2013). Lineages of ectomycorrhizal fungi revisited: foraging strategies and novel lineages revealed by sequences from belowground. Fungal Biology Reviews, 27, 83–99. http://dx.doi.org/10.1016/j.fbr.2013.09.001
Velasco-Bautista, E., Zamora-Martínez, M. C., Nieto de Pascual-Pola, C., Martínez-Valdez, J. I. y Montoya, A. (2010). Modelos predictivos de la producción de hongos silvestres comestibles en bosques de coníferas, Tlaxcala, México. Revista Mexicana de Ciencias Forestales, 1, 95–104. https://doi.org/10.29298/rmcf.v1i1.657
Viveros-Viveros, H., Sáenz-Romero, C., López-Upton, J. y Vargas-Hernández, J. J. (2007). Growth and frost damage variation among Pinus pseudostrobus, P. montezumae and P. hartwegii tested in Michoacán, México. Forest Ecology and Management, 253, 81–88. https://doi.org/10.1016/j.foreco.2007.07.005
Wang, H., Kohler, A. y Martin, F. M. (2025a). Biology, genetics, and ecology of the cosmopolitan ectomycorrhizal ascomycete Cenococcum geophilum. Frontiers in Microbiology, 16, 1502977. https://doi.org/10.3389/fmicb.2025.1502977
Wang, Y., Zhang, W., Cao, Q., Yang, R., Qin, Y. y Zhang, G. (2025b). Description, identification, and growth of ectomycorrhizae in tuber sinense-mycorrhized Castanea mollissima seedlings. Agriculture, 15, 868. https://doi.org/10.3390/agriculture15080868
Wenny, D. L. y Dumroese, R. K. (1987). Germination of conifer seeds surface-sterilized with bleach. Tree Planters’ Notes, 38, 18–21.
White, T. J., Bruns, T. D., Lee, S. B. y Taylor, J. W. (1990) Amplification and direct sequencing of fungal ribosomal RNA genes for phylogenetics. En M. A. Innis, D. H. Gelfand, J. J. Sninsky y T. J. White (Eds.). Protocols and applications. A laboratory manual (pp. 315−322). Nueva York: Academic Press. https://doi.org/10.1016/B978-0-12-372180-8.50042-1
Zhang, M., Li, T. H., Wang, C. Q., Song, B. y Xu, J. (2015). Aureoboletus formosus, a new bolete species from Hunan Province of China. Mycological Progress, 14, 118. https://doi.org/10.1007/s11557-015-1142-z
Zhang, M., Li, T. H., Wang, C. Q., Zeng, N. K. y Deng, W. Q. (2019). Phylogenetic overview of Aureoboletus (Boletaceae, Boletales), with descriptions of six new species from China. Mycokeys, 61, 111−145. https://doi.org/10.3897/mycokeys.61.47520
Zhang, X., Zhang, J., He, J., Li, M., Matsushita, N., Geng, Q. et al. (2024). Physiological and transcriptome responses of Pinus massoniana seedlings inoculated by various ecotypes of the ectomycorrhizal fungus Cenococcum geophilum during the early stage of drought stress. Journal of Fungi, 10, 71. https://doi.org/10.3390/jof10010071
Zhang, P., Zhang, Y., Pang, W., Alonazi, M. A., Alwathnani, H., Rensing, C. et al. (2024). Cenococcum geophilum impedes cadmium toxicity in Pinus massoniana by modulating nitrogen metabolism. Science of the Total Environment, 946, 174296. https://doi.org/10.1016/j.scitotenv.2024.174296
