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Drought Effects on Fine-Root and Ectomycorrhizal-Root Biomass in Managed Pinus Oaxacana Mirov Stands in Oaxaca, Mexico

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Drought Effects on Fine-Root and Ectomycorrhizal-Root Biomass in Managed Pinus Oaxacana Mirov Stands in Oaxaca, Mexico Mycorrhiza (2006) 16: 117–124 DOI 10.1007/s00572-005-0022-9 ORIGINAL PAPER María Valdés . Heidi Asbjornsen . Martín Gómez-Cárdenas . Margarita Juárez . Kristiina A. Vogt Drought effects on fine-root and ectomycorrhizal-root biomass in managed Pinus oaxacana Mirov stands in Oaxaca, Mexico Received: 10 January 2005 / Accepted: 7 September 2005 / Published online: 2 December 2005 # Springer-Verlag 2005 Abstract The effects of a severe drought on fine-root and teractions between multiple disturbances that may lead to ectomycorrhizal biomass were investigated in a forest eco- more severe stress in the host plant and subsequent re- system dominated by Pinus oaxacana located in Oaxaca, ductions in ectomycorrhizal colonization. Mexico. Root cores were collected during both the wet and dry seasons of 1998 and 1999 from three sites subjected to Keywords Ectomycorrhizae . Pinus oaxacana . different forest management treatments in 1990 and as- Prescribed burning . Harvesting . Drought sessed for total fine-root biomass and ectomycorrhizal-root biomass. Additionally, a bioassay experiment with P. oaxacana seedlings was conducted to assess the ectomy- Introduction corrhizal inoculum potential of the soil for each of the three stands. Results indicated that biomasses of both fine roots Mycorrhizal fungi function as an interface between plants and ectomycorrhizal roots were reduced by almost 60% in and the soil (Bethlenfalvay 1992), and most tree seedlings the drought year compared to the nondrought year. There without these associations grow poorly because of a re- were no significant differences in ectomycorrhizal and duced capacity to acquire the soil resources needed for fine-root biomass between the wet and dry seasons. growth (Perry et al. 1987). Research has shown that many Further, the proportion of total root biomass consisting of factors can reduce the presence of mycorrhizal fungi needed ectomycorrhizal roots did not vary between years or seasons. to form these symbiotic associations (Vogt et al. 1993), These results suggest that both total fine-root biomass and including timber harvesting (Jones et al. 2002), fire (Dahlberg ectomycorrhizal-root biomass are strongly affected by 2002; Smith et al. 2004), and drought (Swaty et al. 1998). severe drought in these high-elevation tropical pine forests, Further, ecosystems dominated by plants obligately depen- and that these responses outweigh seasonal effects. Forest dent on ectomycorrhizal associations tend to be high-stress management practices in these tropical pine forests should environments—with stresses ranging from drought, fire, consider the effects of drought on the capacity of P. low temperatures, and flooding (Harley and Smith 1983). oaxacana to maintain sufficient levels of ectomycorrhizae As a genus, Pinus is considered to be obligately dependent especially when there is a potential for synergistic in- on ectomycorrhizal fungal associations for their growth and survival (Trappe 1962; Read 1998). Pine forest ecosystems are typically impacted by periodic drought and fire, and the high stress tolerance of Pinus species is a critical physio- M. Valdés . M. Juárez logical trait enabling pine trees to maintain competitiveness Escuela Nacional de Ciencias Biológicas, in stressful environments (Smith and Hinckley 1995). Instituto Politécnico Nacional, México, D.F., Mexico Although ectomycorrhizal fungal associations have been implicated in maintaining positive water balance in plants H. Asbjornsen (*) . M. Gómez-Cárdenas in general (e.g., Bowen 1973) and in the genus Pinus in Department of Natural Resource Ecology and Management, particular (Parke et al. 1983), few studies have explicitly 234 Science II, Iowa State University, Ames, IA 50011, USA and simultaneously considered the effects of moisture stress e-mail: [email protected] on the dynamics of fine-root biomass production and Tel.: +1-515-2947703 ectomycorrhizal-root biomass and colonization rates. Most Fax: +1-515-2942995 studies that have examined interactions between mycor- K. A. Vogt rhizae and moisture availability have focused on assessing College of Forest Resources, University of Washington, the positive and negative changes in the percent of Seattle, WA, USA ectomycorrhizal colonization of root tips. For example, 118 Meier et al. (1990) and Davies et al. (1996) reported drought, harvesting, and burning) would have greater re- increases in the proportion of root tips colonized by my- ductions in ectomycorrhizal-root biomass compared to the corrhizal fungi in response to moisture stress, while Becker control stand that was only exposed to drought. et al. (1987) and Lansac and Martin (1995)reportedde- creases in colonization rates. Variable patterns of mycor- rhizal colonization have been suggested to result from Materials and methods changes in plant–fungal associations in response to in- herent differences in site conditions related to soil type Description of the study site and moisture availability (Swaty et al. 1998, 2004). However, we are aware of no studies that have quantified The study site is a secondary pine–oak forest located in the changes in total fine-root and ectomycorrhizal-root bio- Sierra Norte region of Oaxaca, Mexico, on lands com- masses in response to moisture stress caused by severe munally owned by Ixtlán de Juárez. The study plots were drought. located between 17°18′16″ latitude and 17°34′00″N, and Both harvesting and prescribed burning are important 96°31′38″ and 96°20′00″W longitude. Elevation ranged forest management practices integral to the management of between 2,190 and 2,400 m above sea level (masl), and pine forests (Smith et al. 1996). Although a relatively slopes averaged 40% (Gómez-Cárdenas et al. 1994). Parent extensive body of literature exists on the effects of these material consists of igneous, sedimentary, and metamorphic management activities on ectomycorrhizal-root coloniza- rocks, and the predominant soils in the region are Vertisols, tion rates and soil inoculum potentials (see Jones et al. 2002 Regosols, and Litosols (INEGI 1984). Both subhumid for a review), the study results have been highly variable, temperate and humid temperate climatic zones are rep- with some reporting negative and others reporting stimu- resented in the region (García 1981). Mean annual average latory effects on mycorrhizal-root colonization. Further, few temperature ranges 12–17°C, and mean annual precipita- studies have quantified changes in ectomycorrhizal-root tion ranges 1,000–1,300 mm, mostly falling in the wet biomass in forest ecosystems under field conditions (Vogt summer season. However, during the El Niño Southern et al. 1982a,b; Dahlberg et al. 1997; Stendell et al. 1999). Oscillation event of 1997–1998, a severe drought occurred Pine forests dominated by Pinus oaxacana Mirov are an throughout southern Mexico (Román-Cuesta et al. 2004). important component of the Sierra Norte mountains of Three forest stands, each approximately 100 ha in size Oaxaca, Mexico, encompassing a total area of approxi- and originally supporting 76–85 pine trees per hectare and mately 12,000 ha (Perry 1991). These forests occur primarily 14–19 oak trees per hectare, were used in this study. Each on communal lands that are managed by local people for stand had been subjected to different silvicultural treatments timber production. Clear-cutting was the traditional silvi- in 1990: (1) seed tree (ST) harvesting, in which 17–27 trees cultural practice for these forests, although prescribed fire is per hectare were left on the site following harvesting to being increasingly used following harvesting operations to provide a seed source for regeneration; (2) ST harvesting facilitate pine regeneration by exposing the mineral soil and (also reducing stand density to 17–27 trees per hectare) reducing understory competition (Keeley and Zedler 1998). with prescribed burning (STB) applied following the Previous studies in these forests have focused on assessing harvesting treatment; and (3) a control site (C) that did the effectiveness of different silvicultural treatments (Becerra not receive any treatment. Prior to the treatments, the ST et al. 1993); however, little is known about the response of and STB stands supported 76 pine trees per hectare and 19 mycorrhizal fungal associations to these anthropogenic oak trees per hectare; the control stand had 85 pine trees per disturbances or to the periodic droughts that commonly hectare and 14 oak trees per hectare. These forests re- occur during the growing season in this region. generated on agricultural lands after abandonment by local Understanding how disturbances caused by both anthro- farmers approximately 50 years earlier. The three stands pogenic activities and naturally occurring extreme climatic had comparable land-use histories and biophysical condi- events influence mycorrhizal dynamics and soil inoculum tions, as indicated by local written and oral accounts and potential in these tropical pine ecosystems is needed for developing sustainable management practices for timber production that ensure the maintenance of symbiotic as- Table 1 Biomasses of total roots, fine roots, and ectomycorrhizal sociations required for healthy ecosystem functioning. This roots (g m−2) and proportion of total root biomass composed of study examined changes in nonmycorrhizal fine-root ectomycorrhizal-root biomass for a drought year (YR1, 1998) and a biomass and ectomycorrhizal-root biomass in P. oaxacana nondrought year (YR2, 1999) forests during a drought and nondrought
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