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ENDANGERED SPECIES RESEARCHEndang Species Res

Vol. 20: 205216, 2013doi: 10.3354/esr00498

Published online May 31

INTRODUCTION

The genus Fagus(Fagaceae) includes 10 species inthe northern hemisphere (Denk 2003, Fang & Lecho-wicz 2006). Fossil records of Fagus in Asia andEurope date from the Miocene and the Pliocene (24to 1.6 million years ago), and in North America fromthe Eocene (45 million years ago) (Huntley et al.1989). Continental drift and climate change have

drastically decreased the distribution of Fagus(Fon-seca et al. 2011, Wilson et al. 2011).

Currently, there are 7 species in eastern Asia:Fagus engleriana, F. hayatae, F. longipetiolata and F.lucida in China; F. crenata and F. japonica in Japan;and F. multinervisin Korea (Horikawa 1972). Thereare also 2 species in Europe and western Asia, F. ori-entalisand Fagus sylvatica (Rose et al. 2009, Milad etal. 2011); and 1 in North America, F. grandifolia (in

Inter-Research 2013 www.int-res.com*Corresponding author. Email: [email protected]

Current distribution and coverage of Mexicanbeech forests Fagus grandifoliasubsp. mexicanain Mexico

Ernesto Ch. Rodrguez-Ramrez1,, Arturo Snchez-Gonzlez1,*,Gregorio ngeles-Prez2

1Laboratorio de Sistemtica Vegetal, Centro de Investigaciones Biolgicas, Universidad Autnoma del Estado de Hidalgo,Ciudad Universitaria, Carretera Pachuca-Tulancingo Km. 4.5, Mineral de la Reforma, Hidalgo 42184, Mexico

2Postgrado Forestal, Colegio de Postgraduados, Montecillo 56230, Texcoco, Estado de Mxico, Mexico

ABSTRACT: Fagus grandifolia subsp. mexicana (Fagaceae) is a taxon endemic to Mexico and iscurrently considered to be in danger of extinction. It dominates the canopy at the sites where itgrows, forming the plant association known as Mexican beech forest. The objectives of this studywere to (1) determine the area currently occupied by beech forests in Mexico, based on a litera-ture review; (2) generate maps showing the distribution and area occupied by the less knownbeech forests in Mexico (which are located in the state of Hidalgo) based on field observations andusing geographic information systems; and (3) propose measures that can be taken to protect andmanage this plant association. The results show that the beech forests of Mexico currently coveran area of 155.54 ha, and several fragments have recently disappeared. The largest patches of

beech forest are located at 5 sites in the state of Hidalgo and occupy a total area of 106.79 ha(73.9%). Each of the sites is different in size, connectedness, and degree of fragmentation and dis-turbance. The Mexican beech forests urgently require management and conservation programs,as some of them will otherwise soon disappear due to changes in land use, logging, and climatechange. To preserve these forests the following measures are suggested in the short term:increase connectedness between beech forest patches, create core areas, reforest with native spe-cies, create seed concentrations, regulate the consumption of beechnuts by humans, and includethis plant association in the National System of Natural Protected Areas.

KEY WORDS: Mexican beech forest Fagus grandifolia subsp. mexicana Forest conservation Forest management Mexico

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Endang Species Res 20: 205216, 2013

eastern Canada and the USA). The intraspecifictaxon at imminent risk of extinction, F. grandifoliasubsp. mexicana (Martnez) A. E. Murray (Valencia &Flores-Franco 2006), is endemic to Mexico (Miranda& Sharp 1950, Williams-Linera et al. 2003, SEMAR-

NAT 2010, Gonzlez-Espinosa et al. 2011).The first specimens of Fagus collected in Mexico

were described as a new species, Fagus mexicanaMartnez. The larger fruits and cuneate base of theleaves were the main characteristics that Martnez(1940) used as a criterion to distinguish it from F.grandifolia. However, in later taxonomic treatments,the taxon was classified as F. grandifolia var. mexi-cana (Martnez) Little (Little 1965), and recently it hasbeen classified as Fagus grandifolia subsp. mexicana(Martnez) A. E. Murray (Valencia & Flores-Franco2006). Although there is still no conclusive evidence

to determine whether it is a species, subspecies or va-riety, it is clear that this taxon has a disjunct distribu-tion and that its populations in Mexico are at graverisk of extinction (Rowden et al. 2004, Tllez-Valdset al. 2006, Premoli et al. 2007, Frankham et al. 2012).In this study, it is treated as a subspecies, as proposedin the most recent publications.

The 11 small populations of Fagus grandifoliasubsp. mexicana recorded to date are restricted tomontane cloud forest in the Sierra Madre Oriental inthe states of Hidalgo, Nuevo Len, Puebla, San LuisPotos, Tamaulipas, and Veracruz (Ern 1976, Rowdenet al. 2004, Montiel-Oscura 2011). Individuals of thissubspecies dominate the canopy at the sites wherethey grow, forming a plant association known asMexican beech forest that has specific environmentalrequirements. They flourish at altitudes of 1400 to2000 m above sea level (asl) on high, steep, north-fac-ing slopes with little variation in climate. The annualaverage temperature ranges between 14.8 and 15.6Cand the total annual precipitation exceeds 1741 mm(Ehnis 1981, Peters 1992, lvarez-Aquino et al. 2004).

Due to its restricted distribution and narrow rangeof suitable habitat, Fagus grandifolia subsp. mexi-cana (Williams-Linera et al. 2003, Fang & Lechowicz

2006, Tllez-Valds et al. 2006) has been designatedas a taxon in danger of extinction by the Mexicanlegislature (SEMARNAT 2010) and has been in-cluded in the Red List of Mexican cloud forest trees(Gonzlez-Espinosa et al. 2011). Several other spe-cies characteristic of the beech forest canopy andsub-canopy are also included in Mexican risk cate-gories (SEMARNAT 2010, Gonzlez-Espinosa et al.2011) and/or international registers of threatenedspecies (CITES 2010), e.g. Clethra mexicana, Cya-thea fulva, Dicksonia sellowiana, Magnolia schiede-

ana, Pinus patula, Podocarpus matudae, and Quer-cus laurina (Ehnis 1981, Prez-Rodrguez 1999,Williams-Linera et al. 2003).

Several authors consider that the information aboutthe distribution, coverage, and conservation status of

the beech forests in Mexico is still incomplete(Martnez 1940, Miranda & Sharp 1950, Fox & Sharp1954, Alcntara & Luna-Vega 2001, Williams-Lineraet al. 2003). In this sense, the working hypothesis ofthe present study is that the current area and distri-bution of beech forest in Mexico is much more exten-sive than has been postulated to date, since there arelarge unexplored, well-conserved areas within themontane cloud forest where climate conditions areideal for this plant association.

MATERIALS AND METHODS

Compilation of bibliographic data

A literature search was conducted to discoverwhich parts of Mexico are known to contain beechforests and which of these already have adequateinformation about the area and current status of theseforests. According to several of the sources consulted(Alcntara & Luna-Vega 2001, Williams-Linera et al.2003, Godnez-Ibarra et al. 2007), the state of Hidalgocontains Mexicos largest Fagus grandifolia subsp.mexicana forests, but the least amount of informationis available about them (except for the forest locatedin La Mojonera in the municipality of Zacualtipn dengeles). We therefore chose the state of Hidalgo asa suitable place to gather local data to complementexisting national data on the distribution, area, andcurrent status of beech forests.

Study area

The study was carried out in the montane cloud for-est in the Sierra Madre Oriental mountain range in

the eastern part of the state of Hidalgo, Mexico(20 19 to 2038N, 98 14 to 9836W). The studysites (El Gosco, El Reparo, La Mojonera, Medio Monte,and Tutotepec) are isolated patches imbedded in thecloud forest, and the canopy is dominated by Fagusgrandifolia subsp. mexicana (Fig. 1). This plant asso-ciation is found between 1557 and 1997 m asl.

The climate where the Mexican beech forests growis Cf; it is humid temperate with year-round rains,characteristic of a mountain orobiome (Garca 1988,Peters 1995), it has a relatively high humidity of 60 to

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85% (Tinoco-Rueda et al. 2009), is frequently foggy,and has an annual average temperature of 12.7Cand a minimum low of 10C. The predominant soilis humic and vitric andosol (FAO-UNESCO 1988).The soil texture is sandy clay loam, with volcanic

glass in some places, and the pH ranges from 4 to 6(Peters 1995).

Fieldwork and herbarium review

Fieldwork was carried out in various regions of thestate of Hidalgo between 2010 and 2011. The beechforests were located based on: (1) specimens from theherbariums MEXU (Instituto de Biologa de la Uni-versidad Nacional Autnoma de Mxico), HGOM(Centro de Investigaciones Biolgicas de la Universi-

dad Autnoma del Estado de Hidalgo), and XAL(Instituto de Ecologa A.C.); (2) data from previousstudies (Miranda & Sharp 1950, Williams-Linera et al.2003, Rodrguez-Ramrez & Moreno 2010); (3) inves-tigation of towns or localities near the beech forestswhere environmental conditions were suitable forthis plant association (Williams-Linera et al. 2003,Rodrguez-Ramrez & Moreno 2010); and (4) showingphotographs of Fagus grandifolia subsp. mexicanatrees and botanical specimens (branches with leavesand inflorescences) to residents of villages near themontane cloud forests, asking them if they werefamiliar with the species.

Patch size estimation

The size of each forest patch was measured bywalking around its perimeter and recording geo-references (UTM coordinates, using a GPS map,

GARMIN, 6OCSx). Since the Mexican beechforests grow in rugged terrain, digital elevation mod-els that take into account the depth of slopes and/orravines were used to collect more reliable data on thearea, coverage, distribution, and degree of fragmen-tation of each forest (Maxwell 1982). Calculationswere carried out with the ArcView V.3.3 program(ESRI 2002). In addition, polygons were drawn ofeach forest patch using USGS Landsat 2012 images(http://glovis.usgs. gov/) with a 3 m level of resolu-tion. The Shape to KML extension of the ArcViewprogram was used to draw the outlines of each patch

on Google Earth V.6 (2011 and 2012) color photos.

Selecting areas for conservationin each forest patch

In order to draw the core areas, the polygons ofeach forest patch were used, with buffers of differentdiameters, using the Patch analyst 3.1 extension ofthe ArcView V.3.3 program with the create coreareas option (Elkie et al. 1999, ESRI 2002, Girvetz &Greco 2007). The core area of each forest was arbi-trarily defined as a minimally disturbed site ormicro-environment located away from gaps or roads(to avoid the edge effect) and with high moistureuptake (due to the presence of water bodies: rivers,streams, or springs). We believe that the selectedcore areas are those with the best environmentalconditions for conservation and management of thisplant association. In addition, the continuity of eachpatch was examined using USGS Landsat 2012images (http://glovis.usgs.gov/), as an essential fac-tor for increasing the probability of preserving envi-ronmental processes at each site (Forman 1995).

Measurement of fragmentation and connectedness

The PatchGrid extension for ArcView V.3.3(Riitters et al. 1995, ESRI 2002), with the spatial sta-tistics by regions option (Elkie et al. 1999), was usedto estimate the degree of fragmentation and connect-edness of each beech forest. The size metrics, edgemetrics, shape metrics, core area metrics, and inter-spersion metrics statistics were used. In order to findthe number of forest patches at each site, the number

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Fig. 1. Fagus grandifolia subsp. mexicana. Current distribu-tion of Mexican beech forest in Mexico


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of patches (NumP) and mean patch size (MPS) statis-tics were calculated. The perimeter of each patchwas estimated using the total edge (TE) and edgedensity (ED) indexes, and the shape of each patchwas determined using the mean shape index (MSI)

and area-weighted mean shape index (AWMSI).Spatial configuration, adjacency, and degree of

interspersion between closely located forest patcheswere obtained using the interspersion and juxtaposi-tion index (IJI), which indicates how evenly forestpatches are distributed. The lower (closer to zero) thevalue of the index, the more randomly the patchesare distributed in the landscape, while higher values(closer to 100) mean that patches are more evenlydispersed (Forman & Godron 1981, Elkie et al. 1999).The distance between nearest patches was estimatedusing the mean nearest neighbor distance index

(MNN; Riitters et al. 1995, Gelet et al. 2010).

RESULTS

Size and current distribution ofMexican beech forest

Based on bibliographic information and field ex-ploration in the state of Hidalgo, 14 areas were foundin Mexico where these forests originally existed.However, the Mexican beech forest plant associationcurrently grows in only 11 of these areas. At the re-maining 3 (Chucuyul-Chiconquiaco, Hueytemalco-Xiutetelco, and Xilitla), it has disappeared or only afew individuals are left, as a result of human activi-ties (changes in land use and deforestation). No ref-erences were found regarding the size of the beechforest at Ojo de Agua de los Indios (Table 1).

The data obtained indicate that there are cur-rently beech forests in the states of Hidalgo, NuevoLen, Tamaulipas, and Veracruz, with a total area of144.54 ha, and that they have virtually disappearedfrom the states of Puebla and San Luis Potos.

Main characteristics of Mexican beech forestin Hidalgo

Information provided by the residents of nearbyvillages has been very useful for locating, describing,and formulating management and conservation pro-posals for these forests. The results indicate thatbeech forests are present in 5 areas and 3 municipal-ities in the state of Hidalgo, and that they are all dif-ferent in terms of size, connectedness, degree of frag-

mentation, and disturbance (Fig. 2). The 5 forests(listed below) are known as Haya (beech) in SanBartolo Tutotepec and Zacualtipn de ngeles andas Tototlcal in Tenango de Doria.

(1) La Mojonera (El Hayal), Zacualtipn de nge-

les municipality (Tables 1 & 2). This is one of thelargest, best studied, and least disturbed beechforests in Mexico. Peters (1992) and Williams-Lineraet al. (2003) state that its area is 45 ha, but in the pres-ent study it was estimated to be 42.5 ha (Table 1). Itlies between 1780 and 1950 m asl, with a slopeexceeding 20. This site has Fagus grandifolia subsp.mexicana seedlings, but they are not as numerous asin other beech forests in Hidalgo. The forest islocated in a temperate orobiome (Peters 1992) typicalof montane environments, with a summer rainy sea-son and temperatures ranging between 11 and 18C.

(2) El Reparo site, Zacualtipn de ngeles munici-pality (Tables 1 & 2). This beech forest is the mostcontinuous and has the lowest degree of disturbance.It is located 2.4 km from the La Mojonera forestbetween 1966 and 1987 m asl, with an area of11.55 ha and pronounced slopes (>40). This smallforest is very important to the residents of the villageof El Reparo because they use it for their water sup-ply, and are therefore trying to preserve it. The tem-perature ranges between 11 and 17C, and the oro-biome is similar to that at La Mojonera, except that inthe sheltered areas there are a large number of treeferns (Cyathea fulva and Dicksonia sellowiana) andseveral bodies of water (streams and rivers). ManyFagus grandifolia subsp. mexicana juveniles wereobserved, and evidence of anthropogenic distur-bance was only present at the edge of the forest nearthe road.

(3) El Gosco site, Tenango de Doria municipality(Tables 1 & 2). This is the smallest beech forest in Hi-dalgo (4.5 ha) and the one with the most evident dis-turbance from illegal logging. The orobiome is similarto that of the La Mojonera forest. This forest is locatedbetween 1557 and 1864 m asl in rugged terrain(slopes >40), with temperatures ranging from 10 to

17C. The forest is severely fragmented, but somepatches grow in steep ravines, out of human reach, sothey are barely disturbed. The finding of this forestadds a record to the known distribution of Fagusgrandifolia subsp. mexicana in the state of Hidalgo.

(4) Medio Monte (Las Hayas) site, San BartoloTutotepec municipality (Tables 1 & 2). This is one ofthe largest (34.25 ha) and least disturbed beechforests. It has a similar orobiome to that of El Reparoand Tutotepec, which are the 3 forest sites with theleast or no evidence of human disturbance. This

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State Locality/municipality Area Elevation Latitude Longitude Status Source(ha) (m)

Hidalgo Medio Monte/San Bartolo 34.25 18001944 202450N, 981424W 1 Present studyTutotepec

Tutotepec/San Bartolo 13.99 19091943 202439.14N, 981652.2W 1 Present studyTutotepec

El Gosco/Tenango de Doria 4.5 15571864 201937.8N 981457.1W 3 Present studyLa Mojonera/Zacualtipn 42.5 17801950 20380.33N 983651.8W 1 Present studyde ngeles

El Reparo/Zacualtipn 11.55 19661987 203805.8N 983513.4W 1 Present studyde ngeles

Nuevo Len Agua Fria/Aramberri 26 1830 2402N, 9942W 1 Montiel-Oscura (2011)Puebla No reference/ 1450 195332.1N 971949.1W 4 Williams-Linera et al.

Hueytemalco-Xiutetelco (2000, 2003)San Luis Xilitla/Xilitla 2122N 9993W 4 Williams-Linera et al.

Potos (2003)Tamaulipas Casa de Piedra, El Cielo 3 1500 230357.8N 99123.8W 3 Williams-Linera et al.

Biosphere Reserve/ (2003)Gmez FariasOjo de Agua de los 1500 2303N 9912W 3 Williams-Linera et al.Indios, El Cielo Biosphere (2003)Reserve/Ocampo

Veracruz Mesa de la Yerba/ 4.05 1900 193337.2N 97019.8W 3 Williams-Linera et al.Acajete (2003)

Acatln Volcano crater/ 4.13 1840 194046.9N 96519.8W 2 Williams-Linera et al.Acatln (2000, 2003)

Acatln Volcano top/ 0.57 1900 194057.5N 965115.3W 2 Williams-Linera et al.Acatln (2000, 2003)

Chucuyul/Chiconquiaco 1750 1946N 9648W 4 Williams-Linera et al.(2003)

Total area 144.54

Table 1. Fagus grandifolia subsp. mexicana. Current distribution and area of Mexican beech forests. 1: good; 2: stable; 3:under threat of extinction; 4: extinct

Fig. 2. Size and distribution of Mexican beech Fagus grandifolia subsp. mexicana forests in Hidalgo State (light green areasin A to E). The circles represent the proposed core areas. Inset: the sampling area in Hidalgo


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forest grows between 1800 and 1944 m asl(Rodrguez-Ramrez & Moreno 2010), in shallowravines (42).

Proposed core areas

In order to counter the effects on Mexican beechforests of disturbance caused by human activities(changes in land use, climate change, and logging), itis proposed that the connected areas be extended

through the creation of core areas.We suggest that 5 core areas beestablished for the Medio Monte siteand the Tutotepec site based on thearea, state of conservation, and exis-

tence of less perturbed sites in these2 forests (Table 2). For the La Mojon-era site, 3 core areas are proposedbased on its area and state of conser-vation. For the El Reparo site, a sin-gle core area is proposed, as this for-est covers 11.5 ha, is continuous, andshows little evidence of disturbance(Fig. 2). Two core areas are sug-gested for the El Gosco beech forest,at the 2 sites least affected by humanactivity, although both are naturally

protected by the inaccessibility ofthe terrain in the ravines where theforest patches are located (Fig. 2).

Fragmentation and connectedness

The beech forests with the largestnumber of patches (NumP) areMedio Monte (8), El Gosco (7), and

Tutotepec (7), while the El Reparo beech forest is1 single continuous patch (Table 2). Analysis of thefragmentation, spatial heterogeneity, configuration,and structure of these Mexican beech forests showedthat the largest MPS in these forests in Hidalgo is atthe El Reparo site (since this forest is a single patch),and the second largest is at the La Mojonera site,with a MPS of 10.78 ha. The smallest MPS (0.65 ha) isat the El Gosco site, where the beech forest is thesmallest in the state and also the most affected byhuman activity.

The analysis of patch shape complexity (ED)showed that the spatial heterogeneity of the land-scape mosaic was highest at the El Gosco site andsecond highest at the Tutotepec site (Table 2). It is

likely that higher patch border complexity is relatedto illegal logging (at El Gosco) or to the type of spatialarrangement (at Tutotepec).

Consistent with the results reported in the previousparagraph, the estimated AWMSI for the El Goscosite was low (1.1), which supports the idea thathuman disturbance causes patch shape simplifica-tion. The higher AWMSI values for the beech forestsat the other 4 sites are characteristic of irregularlandscapes, possibly more natural and/or lessaffected by human activity (Table 2).

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LocalityLa Mojonera El Reparo Tutotepec El Gosco Medio Monte

Slope (degrees)Min. 1.8 3 0.45 16.1 0.45Max. 37.8 37.0 24.9 43.8 21.5

Size metricsNumP 4 1 7 7 8MPS (ha) 10.78 11.46 1.99 0.65 4.10

Edge metricsED (m) 350.86 352.49 274.42 1073.08 922.25

Shape metricsAWMSI (m) 4.12 3.32 4.56 1.10 4.50

MNN (m) 6.46 0 8.69 12.18 38.53Interspersion metricsIJI (%) 98.5 97.2 80.3 30.4 99.1

Proposed core areas (radius; m)1 108.55 310.8 114.71 124.27 227.292 216.36 86.62 111.19 218.413 216.36 86.62 83.444 68.79 83.445 36.94 51.53

Table 2. Fagus grandifolia subsp. mexicana. Size, characteristics, rates of change,and proposed core areas of Mexican beech forests in the state of Hidalgo, Mex-ico. Numbers listed under Proposed core areas represent circles in respectiveareas shown in Fig. 2. NumP: no. of patches; MPS: mean patch size; ED: edgedensity; AWMSI: area-weighted mean shape index; MNN: mean nearest neigh-bor distance; IJI: interspersion juxtaposition index; (): data not estimated


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Configuration is measured by the IJI, which washigh (>80%) at sites less affected by human activity(El Reparo, Tutotepec, La Mojonera, and MedioMonte), indicating that the patches there are moreevenly distributed and have not undergone signifi-

cant changes. In contrast, the IJI value was low(30.4%) for the El Gosco beech forest (Table 2).

The Medio Monte beech forest had the highestMNN value (38.53), as it is made up of 8 patches,closer together than the average distance betweenpatches at the other sites. The El Reparo site has azero MNN value because it is a single continuousforest (Table 2).

DISCUSSION

Size and current distribution ofMexican beech forests

It is estimated that only one-fifth of the worldsoriginal forests are in a favorable state of conserva-tion, these being what have been termed forestfrontiers (Bryant et al. 1997). Moreover, some 10%of the Earths tree species are considered to beendangered and will probably become extinct ifprotective measures are not now put into place(Gonzlez-Espinosa et al. 2011, 2012, Ponce-Reyeset al. 2012). In the particular case of Mexicos mon-tane cloud forests, some 60% of the tree species inthese forests are included in some category of riskaccording to Mexican legislation (SEMARNAT2010), and there is scant knowledge of the density,coverage, or distribution of populations of most ofthese species.

The results of the present study provide informa-tion on basic aspects needed for implementation ofMexican beech forest management and conservationprograms. Previous studies have concluded thatthere are only 10 remnants of Fagus grandifoliasubsp. mexicana forest in the world, and that 3 ofthese are disappearing or have already disappeared.

It was also estimated that they cover an area of lessthan 60 ha (Williams-Linera et al. 2000, 2003, Row-den et al. 2004). However, we found that these forestsoccupy a total area of 144.54 ha, i.e. more than dou-ble the earlier estimate.

The discovery in the field of a new beech forest sitein Hidalgo (El Gosco; present study), and recent datafrom 2 further sites, one at Agua Fria, Nuevo Len,and another at El Reparo, Hidalgo (Montiel-Oscura2011), have increased the known area covered by40.05 ha. Additionally, references to the existence of

beech forests in Tutotepec, Hidalgo (Williams-Lineraet al. 2003), enabled us to locate, describe, and meas-ure 2 further forests, which together cover 48.24 ha,considerably increasing the known area of beech for-est in Mexico.

The beech forests of Mexico were separated fromthose of eastern North America during the Pleis-tocene. Since then, they have been isolated fromtheir northern counterparts (currently by >880 km)and have been growing under specific environmen-tal conditions for 100s of 1000s of years (Little 1965,Peters 1992, Fang & Lechowicz 2006, Premoli et al.2007, Montiel-Oscura 2011). The evidence showsthat, in the past, they underwent contraction ratherthan expansion (Messier et al. 2011) and, in the pres-ent day, rising temperatures and falling moistureavailability in the environment caused by global

warming (Tllez-Valds et al. 2006), as well as frag-mentation and disappearance of the montane cloudforests where small islands of beech forests grow, arerapidly reducing their coverage and distribution(Price et al. 2011).

Although the area occupied by some of the Mexi-can beech forests has remained relatively unchangedin recent decades, for example, near the Acatln Vol-cano, Veracruz (Williams-Linera et al. 2003), and LaMojonera, in the state of Hidalgo (Alcntara & Luna-Vega 2001, Rowden et al. 2004, present study), otherforests survive with a very small number of beechtrees or have disappeared altogether (Williams-Lin-era et al. 2003). The taxon Fagus grandifolia subsp.mexicana is currently classified as endangered, duemainly to uncontrolled human activity such aschanges in land use, forest fires, and illegal logging(Prez-Rodrguez 1999, Rowden et al. 2004, Tllez-Valds et al. 2006, SEMARNAT 2010).

Fragmentation in the beech forest of Hidalgo state

The largest Mexican beech forests are in the stateof Hidalgo, where their combined area is 106.79 ha

(73.9% of the national total). The various forest sitesexhibit differing degrees of fragmentation and dis-turbance; the largest (54.5 ha) and least fragmentedin the entire country are those in the municipality ofZacualtipn de ngeles (La Mojonera and ElReparo), as previously suggested by other authors(Prez-Rodrguez 1999, Alcntara & Luna-Vega2001, Williams-Linera et al. 2003). The data ofWilliams-Linera et al. (2003), Rowden et al. (2004),and the present study show that the Mexican beechforests in San Bartolo Tutotepec (Tutotepec and

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Medio Monte), which cover approximately 48.24 ha,are the least affected by human activity.

The beech forest at the El Gosco site (4.5 ha) in themunicipality of Tenango de Doria shows a highdegree of deterioration, due mainly to clandestine

logging, and will likely soon disappear if measures,such as protecting the zone, are not immediatelytaken to preserve it.

There are significant differences between theMexican beech forests studied in terms of the num-ber, shape, and size of the patches (Table 2). It islikely that the effect of disturbances (logging, fires,and diseases) will have different impacts on the com-position, structure, and ecological processes in eachof the forests (Gang 1998, Messier et al. 2011). Frag-mentation of the forests analyzed here does not nec-essarily imply a high degree of disturbance. For

example, MNN values were high for Medio Monte,which means that the patches are relatively close toeach other, which allows genetic flow between them(Gelet et al. 2010). Additionally, the degree of group-ing, measured by the value of IJI, indicates that thereare connections between patches in all the forestsstudied.

Knowledge about the coverage, degree of frag-mentation, and distribution of the beech forests inMexico is fundamental for establishing managementand conservation programs (Alexandrov & Dakov2010, Ghalachyan & Ghulijanyan 2010). Neverthe-less, there is still little information about otherequally important aspects of these forests, such asspecies diversity and spatial and temporal dynamics(Alcntara & Luna-Vega 2001, Williams-Linera et al.2003, Godnez-Ibarra et al. 2007, Rodrguez-Ramrez& Moreno 2010). Such information can only beobtained from longer term studies (Pagiola et al.2003).

Considering that some patches of these Mexicanbeech forests will soon disappear (Williams-Linera etal. 2003, Rowden et al. 2004, Premoli et al. 2007; ElGosco site in the present study), it is necessary thatmanagement plans and protective measures for the

short term be proposed and implemented.

Conservation and management proposals forMexican beech forests

A viable short-term option is to utilize informa-tion from prior studies on other tree species in thesame genus (Yilmaz 2010), or in other genera butwith similar characteristics or issues to those ofFagus grandifolia subsp. mexicana (Robinson et al.

2009) that could be used as models for manage-ment and conservation of beech forests and of thissubspecies. Some relevant examples are F. multin-ervis, endemic to Ulleung Island, Korea, in whichgenetic studies have helped to preserve the

species through maintenance of high heterozygos-ity (Tomoshi et al. 2006); F. sylvatica, for whichregeneration programs have been implemented inareas suffering deforestation in central Europe(Geler et al. 2007); and F. japonica (in Japan), F.multinervis (in Korea), and F. engleriana (inChina), for which areas have been prioritized forconservation to enable these species to recovernaturally (Peters 1992, Ohkubo et al. 1996). It ispossible that similar methods can help preservethe largest genetic reserve of F. grandifolia subsp.mexicana in Mexico. Molecular level data indicate

that genetic variation in this subspecies is positivelyrelated to population size (Rowden et al. 2004, Pre-moli et al. 2007). Since the mortality rate of F.grandifolia subsp. mexicana seedlings is very highduring the first year (Godnez-Ibarra et al. 2007),the persistence of populations depends largely onthe implementation of programs to reduce theimpact of human activities.

Land ownership

These beech forests are part of community assetsor ejido properties, which means that they are ownedcommunally by the people living in nearby villages(Hardin 1968, Pazos 1991, Congreso de los EstadosUnidos Mexicanos 1992, Gutirrez-Lacayo et al.2002). As a result, all and each of the ejido membersbenefit directly from exploitation of the naturalresources from these forests (lumber, seeds, landuse), and the forests are being consumed rapidly andwithout regulation (the tragedy of the commonssensu Hardin 1968).

This negative dynamic can, however, be reversedif peoples awareness is raised and they are edu-

cated (through workshops and information meet-ings) about the increased benefits they can obtainfrom proper management of natural forest re-sources. Some of the environmental services pro-vided by Mexican beech forests can be useddirectly (mainly edible fungi, seeds, and wood),while indirect services include carbon capture,watershed protection, animal habitats, and recre-ation. A piece of good news is that in mid-2012, thecommunal owners of the largest Mexican beechforest (the La Mojonera Ejido), began taking con-

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crete steps to conserve and manage their forest,with help from Mexican public universities andagencies.

Reforestation of Fagusforests with native species

Mexico has reforestation programs that are basedon detailed analyses determining what tree speciesare suitable for each particular environment. Forexample, native tree species (Pinus spp., Quercusspp.) are used for reforestation, ideal for the soil typeand characteristics where they will be planted (Barryet al. 2010).

In the specific case of Mexican beech forests, theproposal is to increase connectedness betweenpatches through reforestation (Angelsen & Wertz-

Kanounnikoff 2009, Yilmaz 2010) using the moststructurally significant native tree species in theseforests; Liquidambar styraciflua, Magnolia schiede-ana, Pinus patula, Quercus laurina, and Symploccuslimoncillo(Williams-Linera et al. 2003, Rowden et al.2004). The natural coexistence of these species couldfacilitate the growth of Fagus grandifolia subsp. mex-icana seedlings between patches, minimizing theborder effect (Guevara & Laborde 2008, Bezaury-Creel & Gutirrez 2009) and creating concentrationsof seeds.

Four further important recommendations are:(1) Take into account the opinions and empirical

knowledge of the residents living near the areas tobe reforested, since they are generally the ones whorecognize which native species have the best chanceof becoming established (Bennet 1999, Wunder 2006,Barry et al. 2010).

(2) Implement mycorrhizal fungi inoculation tech-niques in reforestation, to increase tree seedling sur-vival probabilities (Rodrguez-Ramrez 2009, Mon-toya et al. 2010).

(3) Choose suitable tree species for reforestation,considering the characteristics, stage of succession,and degree of disturbance of each forest or forest

patch (Morin et al. 2011). For example, in the patchesthat require immediate reforestation (e.g. El Gosco,in Tenango de Doria), species that improve soil qual-ity are suggested (Angelsen & Wertz-Kanounnikoff2009, Morin et al. 2011).

(4) Implement reducing emissions from deforesta-tion and forest degradation (REDD+) programs atthe state level that will help ejido members reducepressure on the natural resources they obtain frombeech forests by enabling them to obtain a good pricefor their agricultural products. The environment and

the populations will benefit in the medium- to longterm from the reduction in greenhouse gas emissions(CONAFOR 2010).

Beechnut consumption

One of the most critical problems for the survival ofFagus grandifolia subsp. mexicana populations isrelated to the trees life cycle. Beech trees produceseeds synchronously in certain years, called mastyears, and the rate of seedling establishment aftergermination is low (Ehnis 1981, lvarez-Aquino &Williams-Linera 2002, Godnez-Ibarra et al. 2007). Inmast years (which occur every 5 to 8 yr), local resi-dents collect the seeds (beechnuts) to eat and/or selllocally, which could decrease the natural regenera-

tion of trees.To increase the viability of Fagus seeds and therate of seedling establishment, it is necessary togradually regulate the practice of seed harvesting,considering the size, degree of disturbance, and frag-mentation of each forest. Ejido members and resi-dents of villages near the beech forests must behelped to understand the importance of regulatingand controlling Fagus seed harvesting, which willbenefit the sustainable use of forest resources and, inconsequence, development in local communities(Blyth et al. 1995, Saunders et al. 1995).

A practice recommended to increase the popula-tion viability of Fagus grandifolia subsp. mexicana isto produce seedlings raised from seed in nurseries.Further experimental protocols could be used toincrease germination and seedling survival (Arriagaet al. 1994), for example, inoculating seeds or seed-lings with native fungi (Rodrguez-Ramrez 2009,Montoya et al. 2010).

During field trips undertaken in February 2012, itwas observed that Fagus grandifolia subsp. mexi-cana trees were flowering synchronously in the 5beech forests in Hidalgo; the forest floor was abun-dantly carpeted with beech flowers and pollen. Later,

during June and July, the first young fruits wereobserved, and by mid-September 100s of 1000s ofseeds were germinating. It was impossible to walk inthe woods without crushing tiny Fagusseedlings atevery step.

Although in this mast year the majority of seedlingsdid not survive (Godnez-Ibarra et al. 2007), it isencouraging to know that the Mojonera forest ejidoresidents are collecting seeds for the tree nurserythat they began to build in late September 2012(pers. comm. from ejido members) to ensure the sur-

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vival of more seedlings, promote forest regeneration,and increase the connectivity of the forest fragmentsby planting trees at suitable sites.

Only by means of a long-term demographic studycan the consequences of harvesting the beechnuts be

unraveled (lvarez-Aquino & Williams-Linera 2002,Godnez-Ibarra et al. 2007). Many demographic dataon long-lived plants show that even harvesting>90% of the seeds of a population does not lower theintrinsic rate of population growth (Cleavitt et al.2008, Barna et al. 2009).

Incorporation into the national system ofNatural Protected Areas

The results of the present study show that the Mex-

ican beech forests are at imminent risk of extinction,as they have a total area of


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Editorial responsibility: Hans Juergen Boehmer,

Bonn, Germany

Submitted: May 31, 2012; Accepted: January 18, 2013

Proofs received from author(s): May 7, 2013
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