894 American Journal of Botany [Vol. 97 Fig. 1. Location of the two Mexican beech forests sampled in the state of Hidalgo, Mexico. Distributions of Fagus grandifolia and F. grandifolia var.mexicana were drawn from biodiversity occurrence data obtained from 42 databases accessed through GBIF Data Portal (http://www.gbif.net, 2009-11-14). The current distribution of the cloud forest in Hidalgo was drawn from Velázquez et al. (2002).In isolation, the two beech forests may have independently ac- in mean slope (t = −3.88, df =38, P ≤ 0.001), median arboreal cover (U = 285,cumulated different bolete species, leading to differences in N = 20, P = 0.02) and the dbh of the nearest tree (U = 125, N = 20, P = 0.04). Slope is steeper and there is more cover in La Mojonera than in Medio Monte,alpha diversity and high beta diversity between forests. Alter- and dbh is greater in Medio Monte than in La Mojonera. Median litter depth (U =natively, the bolete communities may have uniform alpha 207.5, N = 20, P = 0.850) and mean distance to the nearest tree (t = −0.242,diversity and low beta diversity, reﬂecting the shared environ- df = 38, P = 0.810) are not statistically different between the two forests.mental conditions or common historical origins of the relictbeech forests. Understanding the patterns of diversity among Field sampling—Bolete fruit bodies were collected once every 15 d overectomycorrhizal fungi will permit us to make conservation rec- 5 mo in the rainy season of 2007 (from July to November), for a total of 22ommendations that take into account not only the trees, but also samples, 11 in each forest. No fruit bodies were found during four of those 11 sampling forays (July and late November in La Mojonera; July, late October,the microbial environment upon which the trees depend. and early November in Medio Monte), resulting in seven successful samplings at each forest. We followed two sampling procedures: sampling in permanent plots and MATERIALS AND METHODS opportunistic sampling (Mueller et al., 2004). Permanent plots of 100 × 100 m2 were set up in representative areas of each forest, as far as possible from human Study areas—The study was carried out in two well-conserved, isolated settlements and roads. In each plot, we set 10 transects, 100 m in length andforests where Fagus grandifolia var. mexicana is monodominant in the state of 10 m apart. We marked 20 sampling sites in each transect, each separated byHidalgo, Mexico (Fig. 1). One forest is La Mojonera, in the Zacualtipan mu- 5 m, for a total 200 sites in the plot. At each site, all bolete fruit bodies werenicipality, located ca. 20°37′40″N and 98°37′15″W, at 1958 to 1991 m a.s.l. collected in a circular, 5-m2 subplot around the point, for a total sampling areaThe other forest is Medio Monte in the municipality of San Bartolo Tutotepec, of 0.1 ha in each forest, resampled 11 times. During each sampling, opportunis-located ca. 20°24′50″N and 98°14′24″W, at 1800 to 1944 m a.s.l. In both for- tic sampling was conducted by two people who directly searched for fruit bod-ests, the soil type is Andisol, suborder Vitrands (nomenclature follows the U.S. ies for 2 h outside the permanent plot.Soil Taxonomy of the United States Department of Agriculture) with the clearpresence of organic matter. The linear distance between forests is ca. 50 km, Taxonomic identiﬁcation—All the fruit bodies encountered in the ﬁeldand both are surrounded by a matrix of montane cloud forest, with some patches were photographed and put into waxed paper bags for transport to the labora-of cattle pastures. We did not ﬁnd any other potential ectomycorrhizal hosts, tory. Taxonomic identiﬁcation was based on macroscopic morphologicaleither within or surrounding the permanent sampling plots of either forest. descriptions and color changes with chemical reagents (KOH 3–10%, FeSO4To characterize any environmental differences between the two forests, we 10%, and NH4OH 3–70%). Also, microscopic samples of dehydrated materialassessed ﬁve variables at 20 randomly selected sampling sites in the permanent were examined to characterize the size (length and width) and shape of micro-plots (see below) where boletes were sampled: arboreal cover, distance to the scopic structures such as basidiospores, basidia, and cystidia. All this informa-nearest tree, diameter at breast height (dbh) of the nearest tree, slope, and litter tion was used to identify specimens with the taxonomic keys available anddepth. At each sampling site, these variables were measured four times, once in expert opinion. Dehydrated specimens were deposited in the Mycological Col-the direction of each cardinal point, and the mean value was calculated at each lection of the Universidad Autónoma del Estado de Hidalgo (M-UAEH), andpoint for each variable. The two forests have statistically signiﬁcant differences voucher information is provided in Appendix 1. A taxonomic description of
May 2010] Rodríguez-Ramírez and Moreno—Boletes in Mexican beech forests 895each species and a dichotomous taxonomic key is available in Rodríguez- Table 1. Boletaceae species collected in two Mexican beech forests in theRamírez (2009). state of Hidalgo, Mexico. Key codes indicate fruit body abundance, shown in Fig. 3. Data analysis—To include only standardized samples for the alpha diver-sity analyses, we used only data collected in the permanent plots, but for the Species Keybeta diversity analysis, we used data collected with both procedures (permanent Boletellus russellii (Frost) E. J. Gilbert (1931) Oplots and opportunistic sampling). Boletus hypocarycinus Singer (1945) R Before analyzing alpha diversity, we assessed the completeness of the Boletus miniato-olivaceus Frost (1874) Sbolete inventories in each forest as the proportion of observed species richness Boletus rubropunctus Peck (1904) Arelative to maximum expected richness. Expected richness was calculated using Boletus sp. 1 Qtwo nonparametric richness estimators, ICE and ACE, which are based on inci- Leccinum albellum (Peck) Singer (1945) Tdence and abundance data, respectively (Colwell, 2006). These estimators were Leccinum eximium (Peck) Singer (1973) Dcalculated with the program EstimateS version 8.0.0 (Colwell, 2006). Giventhat total number of fruit bodies collected in each forest was markedly different, Leccinum rugosiceps (Peck) Singer (1905) Pwe compared cumulative species richness using rarefaction to standardize sam- Leccinum sp. 1 Nples. Rarefaction curves based on the number of fruit bodies collected, with Leccinum tablense Halling & G. M. Mueller (2003) Hstandard errors, were calculated with the software Species Diversity and Rich- Phlebopus sp. 1 Lness version 3.0.2 (Henderson and Seaby, 2002). Species abundance structure Phylloporus leucomycellinus Singer & M. H. Ivory (1978) Vwas plotted in rank–abundance graphs. The Shannon diversity and Pielou even- Phylloporus sp. 1 Kness indexes were calculated with 95% conﬁdence intervals obtained by boot- Pulveroboletus cramesinus (Secr. ex Watling) M. M. Moser ex Singer Mstrap resampling using the Species Diversity and Richness software (Henderson (1966)and Seaby, 2002). Retiboletus retipes (Berk. & M. A. Curtis) Manfr. Binder & Bresinsky I To assess beta diversity between the two sampled forests, we drew Venn (2002)diagrams with the number of species and genera in three groups: those present Strobilomyces confusus Singer (1945) Uonly in La Mojonera, those present only in Medio Monte, and those shared by Tylopilus felleus (Bull. ex Fr.) Karsten (1818) Gboth forests. As a measure of beta diversity, we calculated the complementarity Tylopilus rubrobrunneus Mazzer & A. H. Smith (1976) Bof the two forests, using the index described by Colwell and Coddington (1994), Tylopilus tabacinus (Peck) Singer (1896) Eat the genus and species levels. Then, to test statistical differences in similarity Tylopilus vinosobrunneus Hongo (1979) Jbetween the two forests, we performed a nonparametric one-way analysis of Xanthoconium separans (Peck) Halling & Both (1998) Fsimilarity (ANOSIM; Clarke and Warwick, 1994). The null hypothesis of the Xerocomus sp. 1 CANOSIM was that there are no statistical differences in species composition Pulveroboletus ravenelii (Berk. & M. A. Curtis) Murrill (1909)abetween the two forests, i.e., mean similarity between pairs of samples within a Boletellus betula (Schwein.) E. J. Gilbert (1931)aforest is not different from the similarity between pairs of samples from differ- Boletus pallidus Frost (1874)aent forests. Then, to search for temporal or spatial groups of samples according Boletus roseolateritius Bessette, Both & Dunaway (2003)ato their similarity in species composition, we constructed single linkage cluster a Species collected only during opportunistic sampling outside permanentdendrograms. To see the inﬂuence of using presence/absence or fruit body sampling plots (see Materials and Methods).abundance data, we calculated both the ANOSIM and cluster analysis usingtwo similarity measures: qualitative and quantitative Sørensen coefﬁcients.These two analyses were performed using the PRIMER ver. 5.0 program (H′ = 2.269, J′ =0.734), but not signiﬁcantly so, given that their(Clarke and Gorley, 2001). 95% conﬁdence intervals overlap. Thus, the rank–abundance graphs are similar for the permanent plots of both forests (Fig. 3), where the most abundant species is Boletus rubropunc- RESULTS tus, which accounted for 35.79% of the total fruit bodies in La Mojonera and 24.64% in Medio Monte. Tylopilus rubrob- We found 484 fruit bodies from 26 bolete species in the runneus was also very abundant at both sites, while T. tabaci-Mexican beech forests sampled (Table 1), ﬁve of which are nus was abundant at La Mojonera but rare at Medio Monte. Theprobably new species, and thus new records for Mexico (to bedescribed elsewhere). Within the permanent plots, we found333 fruit bodies from 20 bolete species in La Mojonera forestand 144 fruit bodies from 14 species in Medio Monte. With op-portunistic sampling, we found four additional species repre-sented by seven fruit bodies, along with many species that wehad also found in the permanent plots (Table 1). For the perma-nent plots of both forests, the ICE richness estimator predicteda higher maximum number of species (26 species for LaMojonera and 16.26 species for Medio Monte) than the ACEestimator (21 and 14.64 species for La Mojonera and MedioMonte, respectively). Thus, according to the incidence-basedestimator, the species inventory within the permanent plot atLa Mojonera is 77% complete, and the inventory of MedioMonte is 86% complete; while for the abundance-based estima-tors both inventories are >95% complete. Even after we standardized the sampling effort to a total of144 fruit bodies per forest, rarefaction curves showed a signiﬁ-cantly higher cumulative richness in the permanent plot atLa Mojonera (20 species) than at Medio Monte (14 species, Fig. 2. Rarefaction curves for boletes in the two Mexican beechFig. 2). Ecological diversity and evenness at La Mojonera forests studied, for the species collected in permanent plots. The bars areare also higher (H′ = 2.328, J′ = 0.753) than at Medio Monte standard errors.
896 American Journal of Botany [Vol. 97 DISCUSSION At present, Mexican beech is restricted to isolated popula- tions in the Sierra Madre Oriental mountain range, probably because of historical events such as the retreat and expansion of its distribution during glacial and interglacial periods in the Pliocene and Pleistocene (Williams-Linera et al., 2003). These events may have shaped the biodiversity associated with these relicts of Mexican beech forests, including that of the ectomyc- Fig. 3. Rank–abundance plots of bolete ensembles collected in perma- orrhizal fungi. Our results support the idea that independent ofnent plots in the two Mexican beech forests studied. Species codes are the isolation of forests, there is a single bolete ensemble with agiven in Table 1. Relative species abundance (ni/N) was plotted on a loga- common history in the two forests studied.rithmic scale against the species-rank ordered by species from those with The community structure of the boletes is similar in the twothe most fruit bodies to those with the fewest. forests in terms of diversity and evenness, with Boletus rubro- punctus and Tylopilus rubrobrunneus the most abundant spe-rarest species at both sites was Strobilomyces confusus, with cies. Beta diversity was low, and ANOSIM detected noonly one fruit body in each forest. signiﬁcant difference in sample similarity within and between For beta diversity between forests, the complementarity forests. This result can be considered robust given that the com-index is only 9% at the genus level (Fig. 4), given that 10 genera pleteness of our bolete inventories is high, whereas undersam-were shared and only one genus was exclusive to La Mojonera pling would result in lower observed similarity values compared(Xerocomus). At the species level, the complementarity value with the true similarity values from complete species invento-reaches 42.30% (Fig. 4) because 15 species were shared be- ries (Chao et al., 2005). However, time intervals between sam-tween the two forests, nine of which (Boletus hypocarycinus, pling events, as well as environmental and phenologicalBoletus sp. 1., B. betula, B. pallidus, B. pallidoroseus, Lecci- conditions may inﬂuence fruit body detectability in samplesnum tablense, Leccinum sp. 1., Tylopilus vinosobrunneus, and (Unterseher et al., 2005; Osono and Takeda, 2006).Xerocomus sp. 1) were found exclusively at La Mojonera and Baselga et al. (2007; Baselga, 2010) explained how betatwo (L. eximium and Phylloporus leucomycellinus) at Medio diversity may be caused by two different phenomena: true spe-Monte only. cies turnover and nestedness. Nestedness occurs when the biota Bolete samples from within the same forest were not signiﬁ- of sites with smaller numbers of species are subsets of the biotacantly more similar than samples from between forests based at richer sites, reﬂecting a nonrandom process of species loss.on either the abundance data (ANOSIM: R = 0.15; P = 0.052) In contrast, spatial turnover implies the replacement of someor the incidence data (R = 0.15; P = 0.051). Neither the cluster species by others as a consequence of environmental sorting oranalysis from the indexes of similarity nor the abundance of spatial and historical constraints (Baselga, 2010 and referencesboletes revealed clear temporal or spatial groupings of therein). In this study, though low, the beta diversity of boletessamples. that we found resulted from the rate of species turnover, be- cause some species were replaced by different species in both forests, so the forest’s species compositions are not subsets of each other. La Mojonera forest clearly harbors more bolete spe- cies richness than Medio Monte does. This richness may be related to the current environmental conditions at La Mojonera. Bolete diversity might be responding to the steeper slope and greater degree of arboreal cover than recorded for Medio Monte. However, more research on the particular responses of ectomy- corrhizal fungi to forest structure and microclimate are needed to understand the symbiosis. For example, ectomycorrhizal species richness may be positively related to forest size (Newton and Haigh, 1998; Peay et al., 2007) and soil type (Gehring et al., 1998). In La Mojonera, the forest is considered one of the most important populations of Mexican beech in terms of its conservation status because it is structurally well developed and regenerating, as indicated by its seedling and sapling densities (Williams-Linera et al., 2003). This Fagus population at La Mojonera is probably the largest and most ge- netically heterogeneous population in Mexico (Pérez, 1999). Plant and landscape ecology studies are needed to characterize the current status of the relict Mexican beech forests. Also, an assessment of the threats to its conservation is needed, given that at present there are no laws or programs to protect Fagus Fig. 4. Venn diagrams with a schematic representation of bolete betadiversity components: the total number of taxa (genera and species) found and the species associated with it.in only one of the two forests (exclusive) and the number of species shared Although a signiﬁcant proportion of ectomycorrhizal fungiby both Mexican beech forests. The percentage of complementarity in might exhibit host speciﬁcity (Newton and Haigh, 1998; Ishidaspecies composition between forests was calculated using the index de- et al., 2007), fortunately from a conservation perspective, sev-scribed by Colwell and Coddington (1994). eral of the bolete species that we report in this study have other
May 2010] Rodríguez-Ramírez and Moreno—Boletes in Mexican beech forests 897hosts in addition to the Mexican beech. Boletus rubropunctus— Henderson, P. A., and R. M. H. Seaby. 2002. Species diversity and rich-the most abundant species—has been thoroughly studied in ness III, version 3.0.2. Pisces Conservation, Lymington, Hampshire,several Quercus hosts (Smith and Pﬁster, 2009). Boletellus UK. Ishida, T. A., K. Nara, and T. Hogetsu. 2007. Host effects on ecto-betula, B. miniato-olivaceus, B. russellii, Strobilomyces con- mycorrhizal fungal communities: Insight from eight host species infusus, Tylopilus eximium, T. felleus, Leccinum rugosiceps, Reti- mixed conifer–broadleaf forests. New Phytologist 174: 430–440.boletus retipes, and Pulveroboletus ravenelii have been found Mueller, G. M., J. P. Schmit, S. M. Huhndorf, L. Ryvarden, T. E.in temperate subhumid forests of Hidalgo, where Pinus, Quercus, O’Dell, D. J. Lodge, P. R. Leacock, et al. 2004. Recommendedand Alnus are the dominant tree genera (Rodríguez-Ramírez, protocols for sampling macrofungi. In G. M. Mueller, G. F. Bills, and2007). M. S. Foster [eds.], Biodiversity of fungi: Inventory and monitoring The 26 species reported in this paper, and the other ﬁve spe- methods, 168–171. Elsevier Academic Press, San Diego, California, USA.cies that have been reported for this vegetation type (Leccinum Newton, A. C., and J. M. Haigh. 1998. Diversity of ectomycorrhizaltalamancae, L. chromapes, L. griseum, Boletus stramineum fungi in Britain: A test of the species–area relation, and the role ofand B. zelleri; Rodríguez-Ramírez, 2007), give us a current to- host speciﬁcity. New Phytologist 138: 619–627.tal of 31 bolete species associated with Mexican beech forests Ortiz-Santana, B., D. J. Lodge, T. J. Baroni, and E. E. Both. 2007.in the state of Hidalgo. Considering the taxonomic and ﬁeld Boletes from Belize and the Dominican Republic. Fungal Diversitylimitations that constrain mycologists, the results we present 27: 247–416.are an important contribution to the knowledge of fungi occur- Osono, T., and H. Takeda. 2006. Fungal decomposition of Abies needle and Betula leaf litter. Mycologia 98: 172–179.ring with this particular host. Peay, K. G., T. D. Bruns, P. G. Kennedy, S. E. Bergemann, and M. Garbelotto. 2007. A strong species–area relationship for eukary- LITERATURE CITED otic soil microbes: Island size matters for ectomycorrhizal fungi. Ecology Letters 10: 470–480.Baselga, A. 2010. Partitioning the turnover and nestedness components Pérez, P. M. 1999. Las hayas de México: Monografía de Fagus gran- of beta diversity. Global Ecology and Biogeography 19: 134–143. difolia spp. mexicana. Universidad Autónoma Chapingo, Chapingo,Baselga, A., A. Jiménez-Valverde, and G. Niccolini. 2007. A mul- México. tiple-site similarity measure independent of richness. Biology Letters Rodríguez-Ramírez, E. Ch. 2007. 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898 American Journal of BotanyAppendix 1. Voucher information of Boletaceae species collected at two Fagus grandifolia var. mexicana forest in the state of Hidalgo, Mexico. All dehydrated specimens are deposited in the Mycological Collection of the Universidad Autónoma del Estado de Hidalgo, Mexico.Taxon; Voucher specimen; Collection locale.Boletellus russellii (Frost) E. J. Gilbert (1931); M-UAEH749, M-UAEH782, Tutotepec. Pulveroboletus cramesinus (Secr. ex Watling) M.M. Moser M-UAEH783; La Mojonera, Zacualtipán de Ángeles. Boletus hypo- ex Singer (1966); M-UAEH779, M-UAEH780; La Mojonera, Zacualtipán carycinus Singer (1945); M-UAEH754; La Mojonera, Zacualtipán de de Ángeles. Retiboletus retipes (Berk. y M.A. Curtis) Manfr. Binder y Ángeles. B. miniato-olivaceus Frost (1874); M-UAEH781; La Mojonera, Bresinsky (2002); M-UAEH769, M-UAEH770; Medio Monte, San Bartolo Zacualtipán de Ángeles. B. rubropunctum Peck; M-UAEH751, Tutotepec, and La Mojonera, Zacualtipán de Ángeles. Strobilomyces M-UAEH762; Medio Monte, San Bartolo Tutotepec, and La Mojonera, confusus Singer (1945); M-UAEH169; Medio Monte, San Bartolo Zacualtipán de Ángeles. Boletus sp. 1; M-UAEH752; La Mojonera, Tutotepec. Tylopilus felleus (Bull. ex Fr.) Karsten (1818); M-UAEH760; Zacualtipán de Ángeles. Leccinum albellum (Peck) Singer (1945); La Mojonera, Zacualtipán de Ángeles. T. rubrobrunneus Mazzer y A. H. M-UAEH766; Medio Monte, San Bartolo Tutotepec. L. eximium Smith (1976); M-UAEH745, M-UAEH746, M-UAEH755, M-UAEH756, (Peck) Singer (1973); M-UAEH767, M-UAEH768; Medio Monte, San M-UAEH757, M-UAEH758, M-UAEH759; Medio Monte, San Bartolo Bartolo Tutotepec. L. rugosiceps (Peck) Singer (1905); M-UAEH771, Tutotepec, and La Mojonera, Zacualtipán de Ángeles. T. tabacinus (Peck) M-UAEH772; La Mojonera, Zacualtipán de Ángeles. L. sp.1; Singer (1896); M-UAEH742, M-UAEH743, M-UAEH784, M-UAEH785; M-UAEH753; La Mojonera, Zacualtipán de Ángeles. L. tablense Halling La Mojonera, Zacualtipán de Ángeles, and Medio Monte, San Bartolo y G. M. Mueller (2003); M-UAEH750, M-UAEH761; La Mojonera, Tutotepec. T. vinosobrunneus Hongo (1979); M-UAEH774, M-UAEH775, Zacualtipán de Ángeles. Phlebopus sp.1; M-UAEH763, M-UAEH764; M-UAEH776; La Mojonera, Zacualtipán de Ángeles. Xanthoconium La Mojonera, Zacualtipán de Ángeles. Phylloporus leucomycellinus separans (Peck) Halling y Both (1998); M-UAEH777, M-UAEH778; Singer y M.H. Ivory (1978); M-UAEH748; Medio Monte, San Bartolo Medio Monte, San Bartolo Tutotepec. Xerocomus sp.1; M-UAEH744; La Tutotepec. Phylloporus sp.1; M-UAEH773; Medio Monte, San Bartolo Mojonera, Zacualtipán de Ángeles.