SciELO - Scientific Electronic Library Online

 
vol.30 número1Cambios en la diversidad y estructura arbórea de un bosque templado bajo dos tratamientos silvícolas en Durango, MéxicoTendencias mundiales en el modelado de la distribución de especies arbóreas índice de autoresíndice de materiabúsqueda de artículos
Home Pagelista alfabética de revistas  

Servicios Personalizados

Revista

Articulo

Indicadores

Links relacionados

  • No hay artículos similaresSimilares en SciELO

Compartir


Revista Chapingo serie ciencias forestales y del ambiente

versión On-line ISSN 2007-4018versión impresa ISSN 2007-3828

Rev. Chapingo ser. cienc. for. ambient vol.30 no.1 Chapingo ene./abr. 2024  Epub 03-Dic-2024

https://doi.org/10.5154/r.rchscfa.2023.05.029 

Scientific articles

Variation in moss biomass as an effect of the use and management of sites in a forest of Abies religiosa (Kunth) Cham. & Schltdl. in Mexico

Jaydine Pen1 
http://orcid.org/0000-0003-4061-6080

Arturo Sánchez-González1  * 
http://orcid.org/0000-0002-3190-8789

José F. Juárez-López1 
http://orcid.org/0000-0003-2291-1588

Claudio A. Hernández-Becerra2 
http://orcid.org/0000-0003-3986-1302

1 Universidad Autónoma del Estado de Hidalgo, Centro de Investigaciones Biológicas, Laboratorio de Ecología de Poblaciones. Ciudad del Conocimiento, km 4.5 carretera Pachuca-Tulancingo. C. P. 42184. Mineral de la Reforma, Hidalgo, México.

2 Comisión Nacional de Áreas Naturales Protegidas, Parque Nacional El Chico. Carretera Casas Quemadas-Mineral del Chico km 7.5. C. P. 42120. Mineral del Chico, Hidalgo, México.


Abstract

Introduction:

Mosses are pioneer organisms known for their high-water storage capacity and biomass. Consequently, they play a significant structural and functional role in terrestrial ecosystems.

Objective:

The aim of the present study was to analyze whether there is variation in moss biomass and water storage capacity among moss species because of site use and management in a fir forest at Parque Nacional El Chico, Hidalgo, México

Materials and methods:

A total of ten sites were selected, distributed across an altitudinal range of 2 781 to 2 981 m: three in the core zone of the park, three in the buffer zone (tourism without moss extraction), and four in the zone with moss extraction. Plots and subplots were established at each site to estimate the biomass and water storage capacity (WSC) of moss species. Furthermore, the coverage of the herbaceous and shrub strata at the sites was determined.

Results and discussion:

Thuidium delicatulum (Hedw.) Schimp. var. delicatulum predominated in biomass across all sites. The highest moss biomass (0.140 g∙cm-2) was found in an extraction site (Esquillero), while the highest Water Storage Capacity (WSC) (0.254 g∙cm-2) was estimated in ecotourism-oriented sites (La Orozca). In sites with moss extraction, coverage of the herbaceous and shrub strata was minimal, and moss biomass increased.

Conclusions:

Use (conservation, ecotourism) and management (extraction) of the forest affects the biomass and WSC values of mosses and the percentage of understory cover (herbaceous and shrubs).

Keywords: bryophytes; moss extraction; fir forest; Parque Nacional El Chico; Thuidium delicatulum.

Resumen

Introducción:

Los musgos son organismos pioneros caracterizados por su alta capacidad de almacenamiento de agua y biomasa, por lo que desempeñan un papel estructural y funcional importante en los ecosistemas terrestres.

Objetivo:

Analizar si existe variación en la biomasa y capacidad de almacenamiento de agua en las especies de musgos como efecto del uso y manejo de sitios en un bosque de oyamel del Parque Nacional El Chico, Hidalgo, México

Materiales y métodos:

Se seleccionaron 10 sitios distribuidos en un intervalo altitudinal de 2 781 a 2 981 m: tres en la zona núcleo del parque, tres en la zona de amortiguamiento (turismo sin extracción de musgo) y cuatro en la zona con extracción de musgo. En cada sitio se establecieron parcelas y subparcelas para estimar la biomasa y capacidad de almacenamiento de agua (CAA) de las especies de musgos. Asimismo, se determinó la cobertura del estrato herbáceo y arbustivo de los sitios.

Resultados y discusión:

En todos los sitios predominó por biomasa Thuidium delicatulum (Hedw.) Schimp. var. delicatulum. La biomasa más alta de musgos (0.140 g∙cm-2) se encontró en un sitio de extracción (Esquillero), mientras que la mayor CAA (0.254 g∙cm-2) se estimó en sitios destinados al ecoturismo (La Orozca). En los sitios con extracción de musgos, la cobertura de los estratos herbáceo y arbustivo fue escasa y la biomasa muscinal incrementó.

Conclusiones:

El uso (conservación, ecoturismo) y manejo (extracción) del bosque afecta los valores de biomasa y CAA de los musgos y el porcentaje de cobertura del sotobosque (herbáceas y arbustos).

Palabras clave: briofitas; extracción de musgo; oyamel; Parque Nacional El Chico; Thuidium delicatulum.

Highlights:

  • Sites where moss extraction occurs exhibit the highest moss biomass.

  • Ecotourism sites show the highest water storage capacity among mosses.

  • All sites were dominated by Thuidium delicatulum (Hedw.) Schimp. var. delicatulum.

  • Moss-extraction sites experience limited coverage of herbaceous and shrub vegetation.

Introduction

Bryophytes (hornworts, liverworts, and mosses) constitute the second most diverse group of plants, comprising approximately 13 000 species (Goffinet et al., 2008). Since they lack effective mechanisms for water content regulation (poikilohydry), environments suitable to their development and reproduction are those that are humid, with moderate temperatures and indirect exposure to sunlight (Vanderpoorten & Goffinet, 2009).

Bryophytes play an important role in maintaining moisture in terrestrial ecosystems, since they can store a high volume of water, about 50 % of the total intercepted rainwater (Delgadillo-Moya, 2014; Vanderpoorten & Goffinet, 2009). Mosses, especially, contribute significantly to plant biomass in several ecosystems; moreover, they are pioneer organisms, because they favor soil formation and water retention in the system.

Mosses are highly sensitive to changes in environmental conditions, which is related to their anatomical (e.g., poorly differentiated tissues and one-cell-thick phyllids) and physiological (poikilohydria) characteristics. Climate change and disturbance (e. g. deforestation, fragmentation and land use change) can affect the abundance and distribution of mosses, because increased temperature favors water loss by evapotranspiration, which decreases metabolic activity and causes tissue damage (Oishi, 2018; Siwach et al., 2021; Toro Manriquez et al., 2020).

Regarding direct uses, mosses are used for air pollution monitoring, because they absorb particles and chemical compounds from the atmosphere that they store in their tissues (Siwach et al., 2021). Additionally, these bryophytes serve as a non-timber forest resource of economic importance in Mexico, especially during the Christmas season (Acatitla Pluma et al., 2020); however, the intensive extraction and sale of mosses prevent the regeneration of their populations and cause environmental deterioration of the soil and forest, which could represent a risk for the survival of these species (Anastacio Martínez et al., 2017; Hernández-Rodríguez & Delgadillo-Moya, 2021).

Delgadillo-Moya (2014) mentions that information on floristic diversity, ecological importance and potential uses of mosses in Mexico is still scarce. In the particular case of the state of Hidalgo, there have been some studies focused on the richness and distribution of this group of plants, but most of the territory has not been explored (Delgadillo et al., 2014). The present research had the following objectives: 1) to analyze whether there is variation in the biomass and water storage capacity of moss species in sites for conservation, ecotourism and extraction in the oyamel fir forest of Parque Nacional El Chico, Hidalgo, Mexico, and 2) to estimate whether the coverage of the herbaceous and shrub stratum differs between sites according to the conditions of use. The aforementioned to provide information on the effect of moss extraction in this oyamel fir forest.

Materials and Methods

Study area

The Parque Nacional El Chico (PNC) is located in the Sierra de Pachuca, within the Transmexican Volcanic Belt. The predominant climate in the area is C(m)(w)b(i’)gw’’; that is, temperate subhumid with summer rainfall, mean annual temperature between 12 and 18 °C and total annual precipitation of 1 479.5 mm (Comisión Nacional de Áreas Naturales Protegidas [CONANP], 2005; Razo-Zárate et al., 2013). The dominant vegetation in the PNC consists of temperate sub-humid forests of Abies (62.9 %), %), Quercus-Abies (10.55 %), Juniperus (4.67 %), Quercus (3.97 %), Abies-Quercus (3.93 %), Quercus-Pinus (2.25 %), Pinus (1.87 %), Cupressus (1.16 %) and Pinus-Quercus (0.87 %); in addition, there is secondary grassland-scrub (2.18 %) and, in the rest of the area, aquatic, ruderal and arvense, rupicolous and xerophytic vegetation. In general, these vegetation types develop among numerous rocky formations, small valleys, lagoons and rivers (CONANP, 2005).

Moss sampling sites and units

Sampling sites in the oyamel fir forest of the PNC, dominated by Abies religiosa (Kunth) Cham. & Schltdl., were chosen based on the following criteria: 1) three sites without moss extraction, located in the core zone (CZ), 2) three sites without moss extraction, but with ecotourism activities (adjacent to the CZ) and 3) four sites where moss extraction is allowed (in the periphery of the PNC). The sampling sites are distributed in an altitudinal range between 2 780 and 2 900 m (Figure 1).

Figure 1 Location of the sampling sites, within (core zone) and on the periphery (ecotourism and moss extraction zone) of Parque Nacional El Chico, Hidalgo, Mexico. 

At each sampling site, two 10 m x 10 m plots were established randomly, separated by a distance of 10 to 15 m from each other, according to the BRYOLAT methodology (Gabriel et al., 2014). Each plot was divided into 25 squares (2 m x 2 m), to subsequently select three of them at random. Each square was divided into micro squares of 5 cm x 5 cm, to carry out the sampling in three of them randomly selected. Therefore, sampling was performed in 18 microplots at each site: one site, two plots, three squares per plot, three microplots per square, i. e. 1 × 2 × 3 × 3 = 18.

Estimation of biomass and water storage capacity

Biomass and water storage capacity (WSC) of mosses were estimated in the following order: (1) separation of samples per site, (2) determination of the species present in each sample, (3) estimation of dry weight or biomass (DB), for which samples were placed on newspaper and allowed to dry for one week at room temperature, and (4) estimation of saturated or wet biomass or weight (WB), for which samples were hydrated for 30 minutes and placed in a mesh for one hour in order to preserve their integrity and remove excess water by runoff (Oishi, 2018). Weights were determined using a triple beam balance or an analytical balance, according to the amount of biomass. The WSC of the species/samples was determined using the formula described by Oishi (2018): WSC (g∙100 cm-2) = WB - DB.

Coverage of the herbaceous and shrub strata

In each locality, two randomly selected plots measuring 10 m x 10 m were established to determine the percentage of coverage for the herbaceous and shrub strata, following the scale proposed by Braun-Blanquet (Mueller-Dombois & Ellenberg, 1974), modified by Mejía Canales et al. (2018). In this scale, 5 = 75 to 100 %, 4 = 50 to 74.9 %, 3 = 25 to 49.9 %, 2 = 10 to 24.9 %, and 1 = less than 9.9 %. For this purpose, a visual estimation was made of the approximate percentage of the total ground area covered by the herbaceous or shrub stratum, without taking into account the overlap between individuals (Fehmi, 2010) or the taxonomic identity of species in both strata. The coverage percentages of the two plots in each locality were averaged to obtain a single value on the Braun-Blanquet scale.

Tree canopy cover

Tree canopy cover was estimated using a convex spherical densiometer. For this purpose, five measurements were made in each sampling plot (two per site), in each corner and in the central part; in each position the canopy cover was estimated considering the four cardinal points, as indicated in the manual of use of the densiometer (Lemmon, 1956). From these 10 measurements (five for each plot), the average canopy coverage per sampling site was calculated. The results were considered an indirect measure of the amount of light that moss species receive in the forest (Baudry et al., 2014).

Identifying species

Moss specimens were identified to family and genus level using the taxonomic keys of Gradstein et al. (2001) and to species level using the keys of Sharp et al. (1994). The nomenclature of species and taxonomic authors was updated based on information from the WFO project website (2023).

Data analysis

The values of dry biomass, wet biomass, and water storage capacity per sampling site showed no normal distribution. Therefore, the data obtained for each sampling site were compared using the non-parametric Mann-Whitney test. This test compares two independent samples (pairs of sampling sites) when the data are not normally distributed, generating a test statistic U and a P-value, which are used to determine whether the two groups (pairs of sampling sites) are significantly different. The analysis was conducted using the PAST software version 4.03 (Hammer et al., 2001), which automatically estimates a Bonferroni adjustment or correction of the P-value based on the number of comparisons made between sites independently. This is done to avoid errors in the interpretation of the results (Matamoros & Ceballos, 2017).

Results

Moss species with the highest biomass in the oyamel fir forest

We identified five species of mosses with structural importance in the 10 sampling sites, but Thuidium delicatulum (Hedw.) Schimp. var. delicatulum had the highest biomass values. Two other varieties were determined in this species: Thuidium delicatulum var. peruvianum and T. delicatulum var. radicans, but they showed low biomass values. The other most representative moss species in terms of biomass value were Bryum argenteum Hedw., Ceratodon purpureus (Hedw.) Brid., Cyrto-hypnum mexicanum (Mitt.) W. R. Buck & H. A. Crum and Helicodontium capillare (Hedw.) A. Jaeger (Table 1). The general characteristics of the identified species and varieties with data on their habitat (vegetation type), altitude, climate and geographic distribution are described in Annex 1.

Table 1 Record of moss species (1 = present, 0 = absent) according to sampling site: Core zone (A = Cedros, B = Los Shishis, C = Los Conejos), tourist zone (D = Camino hacia Conejos, E = Peña de la Orozca, F = Orozca) and moss extraction zone (G = Esquillero, H = Esquillero, I = Cruz Blanca, J = Viga Tirada) in an Abies religiosa forest at Parque Nacional El Chico, Hidalgo. 

Species Sampling sites
A B C D E F G H I J
Bryum argenteum Hedw. 0 0 0 0 0 0 0 1 0 1
Ceratodon purpureus (Hedw.) Brid. 0 1 0 0 0 0 0 0 0 0
Cyrto-hypnum mexicanum (Mitt.) W. R. Buck & H. A. Crum 0 0 1 1 0 1 0 1 1 1
Helicodontium capillare (Hedw.) A. Jaeger 1 0 0 0 0 0 0 0 0 0
Thuidium delicatulum (Hedw.) Schimp. var. delicatulum 1 1 1 1 1 1 1 1 1 1
T. delicatulum var. peruvianum (Mitt.) H. A. Crum 1 0 0 0 0 1 1 0 1 0
T. delicatulum var. radicans (Kindb.) H. A. Crum, Steere & L. E. Anderson 0 0 0 0 0 0 0 1 0 0

Water storage capacity, wet biomass and dry biomass

Table 2 indicates that the species T. delicatulum var. delicatulum predominated widely over the others due to its presence and structural values: dry biomass, wet biomass, and WSC (water storage capacity). Figure 2 shows the results of the Mann-Whitney test, which indicates that dry biomass, wet biomass, and WSC of the mosses vary significantly (P < 0.05) among most of the analyzed sites. The core zone sites (A, B, and C) were similar to each other in the three analyzed variables but had lower values of biomass and WSC than most of the sampling sites in the ecotourism zones (E and F) and moss extraction zones (I and J). Regarding the ecotourism zone sites (D, E, and F), only site D differed from the others in the three analyzed variables; however, they did show statistically significant differences (P < 0.05) compared to most of the core zone sites and moss extraction zone sites. On the other hand, the moss extraction sites (G, H, I, and J) differed from each other in the three analyzed variables; sites G and H with low values of dry biomass, wet biomass, and WSC differed significantly from sites I and J. When comparing the extraction zone sites with the core zone and ecotourism zone sites, it was observed that two of them (I and J) had significantly higher values in the three analyzed variables (Figure 2).

Figure 2 The Mann-Whitney test was conducted for I) water storage capacity (yellow color), II) wet biomass (green color), and III) dry biomass (blue color) of mosses identified in a forest of Abies religiosa. Whiskers indicate the maximum and minimum values of the variables at each site, and the vertical line within the boxes represents the median, indicating where 50 % of the data lies. The letters at the bottom of the figure (x-axis) represent the names of the sites: A: Cedros, B: Los Shishis. C: Los Conejos, D: Camino hacia Conejos, E: Peña Orozca, F: La Orozca, G: Esquillero 7, H: Esquillero 8, I: Cruz Blanca and J: Viga Tirada. Each site was compared with each of the other nine sites; the letters at the top of each box correspond to the sites that exhibited statistically significant differences (P < 0.05) when compared with the site positioned on the x-axis according to the Bonferroni test. For example, in the first graph, sites A, B, C, and D had no significant differences between them, so no letters are included at the top of the corresponding boxes. In contrast, sites E and F (x-axis) had significant differences compared to sites C and D. 

Table 2 Record of dry biomass (DB), wet biomass (WB), wet biomass to dry biomass ratio (WB/DB) and water storage capacity (WSC) per moss species in an Abies religiosa forest at Parque Nacional El Chico, Hidalgo. 

Taxon DB (g∙cm-2) WB (g∙cm-2) WB/DB (%) WSC (g∙cm-2)
Bryum argenteum 22.35 126.44 5.7 1.041
Ceratodon purpureus 24.7 124.08 5 0.994
Cyrto-hypnum mexicanum 35.2 185.61 5.3 1.503
Helicodontium capillare 5.6 34.33 6.1 0.287
Thuidium delicatulum var. delicatulum 585.12 4 455.51 7.6 36.445
T. delicatulum var. peruvianum 35.46 170.31 4.8 1.349
T. delicatulum var. radicans 3.6 23.6 6.6 0.2

On the other hand, Table 3 indicates that the percentage of coverage of herbaceous and shrubby species, estimated using the Braun-Blanquet scale, differed among the sampling sites based on the type of disturbance or management (conservation, ecotourism, and moss extraction). The core zone and ecotourism sites at PNC exhibited well-developed herbaceous and shrubby layers (50 % or more coverage), but the shrubby layer was less prominent in the core zone (25 % or more coverage). In contrast, in the moss extraction zone sites, the coverage of both herbaceous and shrubby layers was low (25 % or less), leading to the forest floor being covered by plants with a creeping habit; specifically, mosses. It is important to mention that in all sampling sites, the dominant tree species was A. religiosa, and the canopy coverage was relatively uniform (between 92.5 % and 98 %).

Regarding the herbaceous and shrub strata, in all sites, the most common species and with the highest contribution to the percentage of cover were: Acaena elongata L., Ageratina glabrata (Kunth) R. M. King & H. Rob., Alchemilla procumbens (Rose) Rydb., Baccharis conferta Kunth, Ribes ciliatum Humb. & Bonpl. ex Roem. & Schult., Roldana angulifolia (DC.) H. Rob. & Brettell, Salvia elegans Vahl, Sigesbeckia jorullensis Kunth and Symphoricarpus microphyllus Kunth. It should be emphasized that the percent cover of herbs and shrubs was estimated at the stratum level, not at the species level.

Table 3 Cover of the herbaceous, shrub, and canopy strata; water storage capacity (WSC), and biomass (WSC/dry biomass) of mosses, at each sampling site in a forest of Abies religiosa at Parque Nacional El Chico, Hidalgo. The coverage of shrub and herbaceous strata was qualitatively determined through visual observation, considering the space occupied by each stratum. 

Site Strata cover (%) Elevation (m) WSC (g∙cm-2) Biomass (g∙cm-2)
Herbaceous Shrub Canopy
A. Cedros 50-74.9 25-49.9 97 2 871 0.171 0.053
B. Los Shishis 50-74.9 50-74.9 93 2 919 0.161 0.051
C. Los Conejos 75-100 50-74.9 93 2 822 0.115 0.599
D. Camino Conejos 50-74.9 25-49.9 96 2 822 0.096 0.049
E. La Peña Orozca 50-74.9 25-49.9 96 2 850 0.245 0.044
F. La Orozca 50-74.9 25-49.9 94 2 851 0.254 0.042
G. Esquillero 1 10-24.9 <9.9 93 2 957 0.087 0.140
H. Esquillero 2 <9.9 <9.9 98 2 957 0.064 0.073
I. Cruz Blanca <9.9 <9.9 95 2 891 0.233 0.054
J. Viga Tirada <9.9 <9.9 94 2 901 0.245 0.042

Discussion

In all examined sites, T. delicatulum var. delicatulum stood out as the most significant moss species in terms of biomass and WSC values. The widespread presence and structural importance, reflected in higher biomass, across all sites suggest that this taxon possesses a notable capacity for adaptation and phenotypic plasticity compared to other coexisting moss species within the altitude gradient of 2 822 to 2 957 m and the analyzed vegetation types (Anastacio Martínez et al., 2021). Consistent with these findings, Cárdenas (1999) notes that this moss variety thrives in mature forests or at different stages of succession with varying degrees of disturbance, where trees of the genera Abies, Juniperus, and Quercus dominate, mainly; that is, they grow in heterogeneous microenvironmental conditions either under the shade of the canopy or with exposure to radiation (soil, trunks, roots or branches), in dry or humid places, in an altitudinal interval that ranges between 2 830 and 2 900 m. In any case, all identified mosses share characteristics: wide geographic distribution on the planet, presence and structural importance in various types of vegetation and at different stages of succession, which indicates that they can survive in a wide range of environmental conditions (Hernández-Rodríguez et al., 2021; Sharp et al., 1994).

The results clearly show the impact of tourism and extraction activities on the most structurally important moss species in the oyamel fir forest of Parque Nacional El Chico, Hidalgo, since there are significant differences in the wet biomass, dry biomass and WSC of mosses between sampling sites. These differences are related to the type of forest management or use (extraction, tourism or conservation); however, it was also found that, in some sites, variables were independent of the type of forest use or management, so it is likely that other factors at local (exposure, slope, soil type, light input and floristic composition) and regional (temperature and humidity) scales influence the structure of the moss stratum (Fojcik et al., 2019; Stefanska-Krzaczek et al., 2022; Toro Manriquez et al., 2020). In this sense, several authors indicate that the presence and abundance of moss species is related to other plant groups and to multiple environmental factors such as solar radiation, canopy cover, humidity, and soil properties (Oishi, 2018; Siwach et al., 2021; Toro Manriquez et al., 2020).

In the present study, biomass and WSC values of mosses and percent cover of herbaceous and shrub strata in the oyamel fir forest of Parque Nacional El Chico differed markedly between managed/used (moss extraction/ecotourism) and unmanaged (core zone) sites. This is consistent with results in other studies indicating that bryophyte richness, distribution, and biomass are closely associated with the composition and structural values of coexisting vascular plant species (Fojcik et al., 2019; Hernández-Rodríguez et al., 2021; Stefanska-Krzaczek et al., 2022; Toro Manriquez et al., 2020).

The cover of the herbaceous and shrub stratum is negatively related to the total cover or biomass of bryophytes, due to competition for water, light and growth substrate (Fojcik et al., 2019). In the study area, the understory structure was evidently modified in two of the sites where moss is extracted, there the cover of herbaceous and shrub strata was drastically reduced and the biomass and WSC of mosses increased. However, at two remaining sites, biomass and WSC were not significantly different from the values estimated at the core zone sites. In these cases, it is possible that the upper tree layer dominated by A. religiosa in all sampling sites, by remaining intact, is generating a buffering effect by providing partial protection against desiccation and the passage of enough light for moss growth and reproduction (Hernández-Rodríguez et al., 2021; Toro Manriquez et al., 2020). In the synecological analysis conducted by Hernández-Álvarez et al. (2021), the species composition of the herbaceous and shrub stratum of the oyamel fir forest showed no difference between sites; the species identified were characteristic of this type of vegetation, so the differences between sites were at the structural level (cover).

In general, the results indicate that there are evident changes in understory structure (herbs and shrubs) and in biomass and WSC of mosses, among sites with different use and management. Therefore, a detailed study of forest structure and quantification of local environmental factors (light, temperature, humidity, exposure and slope) under different management conditions (Gabriel et al., 2014; Hernández-Rodríguez et al., 2021; Herrera-Paniagua et al., 2018; Zepeda-Gómez et al., 2014) and over time, through a monitoring program, are necessary. In this way, it would be possible to define more precisely what the near future of the PNC’s oyamel fir forest, ecotourism activities, and moss extraction will be if they continue as they have been doing so far.

The extraction of mosses from forests in Mexico should be a controlled activity. Most species of this group of plants are intensively extracted and the most affected ones such as T. delicatulum (Acatitla Pluma et al., 2020; Anastacio Martínez et al., 2017) are not yet included in any risk category within the Mexican Official Standard NOM-059-2010 (Secretaría del Medio Ambiente y Recursos Naturales [SEMARNAT], 2010) because there is a lack of quantitative biological information on the current status of their populations.

Conclusions

Moss species in the oyamel fir forest of Parque Nacional El Chico have high water store capacity (WSC) and wide coverage (biomass) at all sampling sites. Dry biomass, wet biomass and WSC values differed among sites, in general, depending on forest use and management (conservation, ecotourism or extraction). At moss extraction sites, herbaceous and shrub plant species cover was low compared to ecotourism and conservation sites, which probably favored the increase in moss biomass. The canopy cover could be maintaining the radiation and humidity conditions necessary for moss growth in some of the sites with use and management. Therefore, it is important to implement a monitoring program and moss management and conservation measures to maintain the basic ecological structure of the oyamel fir forest.

Acknowledgments

We thank the authorities of Parque Nacional El Chico for the facilities granted to carry out this study. In addition, we are grateful for the valuable comments and suggestions of three anonymous reviewers who contributed to substantially improve the final version of this manuscript.

Referencias

Acatitla Pluma, O., Villamil Carrera, C. y Martínez-Pérez, J. L.2020.La importancia comercial de los musgos.Madera y Bosques, 26(3), 10.21829/myb.2020.2632031 [ Links ]

Anastacio Martínez, N. D., Franco-Maass, S., Valtierra Pacheco, E. y Nava Bernal, G.2017.El proceso de extracción y comercialización del musgo (Thuidium delicatulum) en el Estado de México.Ciencia Ergo Sum, 24(1), 44‒61.https://www.redalyc.org/articulo.oa?id=10449880005Links ]

Baudry, O., Charmetant, C., Collet, C. y Ponette, Q.2014.Estimating light climate in forest with the convex densiometer: Operator effect, geometry and relation to diffuse light.European Journal of Forest Research, 133, 101-110.10.1007/s10342-013-0746-6 [ Links ]

Cárdenas, S.1999.Los musgos pleurocárpicos del Valle de México.Tropical Bryology, 16, 109‒116.https://www.scielo.org.mx/scielo.php?script=sci_nlinks&ref=6831567&pid=S1870-3453201400010001200007&lng=esLinks ]

Comisión Nacional de Áreas Naturales Protegidas (CONANP)2005.Programa de conservación y manejo: Parque Nacional El Chico, México.https://www.conanp.gob.mx/que_hacemos/pdf/programas_manejo/PN_Chico.pdfLinks ]

Delgadillo-Moya, C.2014.Biodiversidad de Bryophyta (musgos) en México.Revista Mexicana de Biodiversidad, 85, 100‒105.10.7550/rmb.30953 [ Links ]

Delgadillo, C., Villaseñor, J. L., Cárdenas, Á. y Ortiz, E.2014.Diversidad y distribución de musgos en el estado de Hidalgo, México.Revista Mexicana de Biodiversidad, 85(1), 84‒97.10.7550/rmb.35761 [ Links ]

Fehmi, J. S.2010.Confusion among three common plant cover definitions may result in data unsuited for comparison.Journal of Vegetation Science, 21(2), 273‒279.10.1111/j.1654-1103.2009.01141.x [ Links ]

Fojcik, B., Wierzgoń, M. y Chmura, D.2019.Response of bryophytes to disturbances in managed forests. A case study from a Polish forest.Cryptogamie, Bryologie, 40(10), 105‒118.10.5252/cryptogamie-bryologie2019v40a10 [ Links ]

Gabriel, R., Coelho, M. M., Henriques, D. S., Borges, P. A., Elias, R. B., Kluge, J. y Ah-Peng, C.2014.Long-term monitoring across elevational gradients to assess ecological hypothesis: a description of standardized sampling methods in oceanic islands and first results.Arquipelago - Life and Marine Sciences, 31, 45‒67.https://islandlab.uac.pt/fotos/projectos/1440666688.pdfLinks ]

Goffinet, B., Buck, W. y Shaw, A.2008.Morphology, anatomy, and classification of the Bryophyta. En Shaw, B. y Goffinet, A. (eds.). Bryophyte biology. Cambridge University Press. [ Links ]

Gradstein, R. S., Churchill, S. P. y Salazar-Allen, N.2001.Guide to the bryophytes of Tropical America (vol. 86). New York Botanical Garden. [ Links ]

Hammer, Ø., Harper, D. A. y Ryan, P. D.2001.PAST: Paleontological Statics software package for education and data analysis.Paleontología Electrónica, 4(1), 9.https://palaeo-electronica.org/2001_1/past/past.pdfLinks ]

Hernández-Álvarez, A. G., Reyes-Ortiz, J. L., Villanueva-Díaz, J. y Sánchez-González, A.2021.Variación en la estructura del bosque de Abies religiosa (Pinaceae), en diferentes condiciones de manejo y disturbio.Acta Botanica Mexicana, 128, 10.21829/abm128.2021.1752 [ Links ]

Hernández-Rodríguez, E. y Delgadillo-Moya, C.2021.The ethnobotany of bryophytes in Mexico.Botanical Sciences, 99(11), 13‒27.10.17129/botsci.2685 [ Links ]

Hernández-Rodríguez, E., Escalera-Vázquez, L. H., García-Ávila, D., Montoro Girona, M. y Mendoza, E.2021.Reduced-impact logging maintain high moss diversity in temperate forests.Forests, 12(4), 10.3390/f12040383 [ Links ]

Herrera-Paniagua, P., Martínez, M. y Delgadillo-Moya, C.2018.Patrones de riqueza y de asociación al hábitat y microhábitat de los musgos del Área Natural Protegida Sierra de Lobos, Guanajuato, México.Revista Mexicana de Biodiversidad, 89(4), 1002‒1011.10.22201/ib.20078706e.2018.4.2455 [ Links ]

Lemmon, P. E.1956.A spherical densiometer for estimating forest overstory density.Forest Science, 2(4), 314-320.10.1093/forestscience/2.4.314 [ Links ]

Matamoros, P. R. A. y Ceballos, M. A.2017.Errores conceptuales de estadística más comunes en publicaciones científicas.Revista CES Medicina Veterinaria y Zootecnia, 12(3), 211‒229.10.21615/cesmvz.12.3.4 [ Links ]

Mejía Canales, A., Franco-Maass, S., Endara Agramont, A. R. y Ávila Akerberg, V.2018.Caracterización del sotobosque en bosques densos de pino y oyamel en el Nevado de Toluca.Madera y Bosques, 24(3), 10.21829/myb.2018.e2431656 [ Links ]

Mueller-Dombois, D. y Ellenberg, H.1974.Aims and methods of vegetation ecology. John Wiley and Sons. [ Links ]

Oishi, Y.2018.Evaluation of the water storage capacity of bryophytes along an altitudinal gradient from temperate forests to the alpine zone.Forests, 9(7), 10.3390/f9070433 [ Links ]

Razo-Zárate, R., Gordillo-Martínez, A., Rodríguez-Laguna, R., Maycotte-Morales, C. y Acevedo-Sandoval, O.2013.Escenarios de carbono para el bosque de oyamel del Parque Nacional El Chico, Hidalgo, México.Revista Latinoamericana de Recursos Naturales, 9(1).https://revista.itson.edu.mx/index.php/rlrn/article/view/207Links ]

Secretaría de Medio Ambiente y Recursos Naturales (SEMARNAT)2010.Norma Oficial Mexicana NOM-059-SEMARNAT-2010, Protección ambiental- Especies nativas de México de flora y fauna silvestres- Categorías de riesgo y especificaciones para su inclusión, exclusión o cambio. Lista de especies en riesgo. Diario Oficial de la Federación.https://dof.gob.mx/nota_detalle_popup.php?codigo=5173091Links ]

Sharp, A. J., Crum, H. y Eckel, P. M.1944.The moss flora of Mexico (vol. 69). New York Botanical Garden. [ Links ]

Siwach, A., Kaushal, S. y Baishya, R.2021.Effect of mosses on physical and chemical properties of soil in temperate forests of Garhwal Himalayas.Journal of Tropical Ecology, 37(3), 126‒135.10.1017/S0266467421000249 [ Links ]

Stefanska-Krzaczek, E., Swacha, G., Zarnowiec, J., Raduła, M., Kącki, Z. y Staniaszek-Kik, M.2022.Central European forest floor bryophytes: Richness, species composition.Ecological Indicators, 139, 1‒12.10.1016/j.ecolind.2022.108954 [ Links ]

Toro Manriquez, M., Ardiles, V., Promis, Á., Huertas Herrera, A., Soler, R., Lencinas, M. y Martínez Pastur, G.2020.Forest canopy-cover composition and landscape influence on bryophyte communities in Nothofagus forests of southern Patagonia.PLoS ONE, 15(11), 10.1371/journal.pone.0232922 [ Links ]

Vanderpoorten, A. y Goffinet, B.2009.Sampling of bryophytes. En Vanderpoorten, A. y Goffinet, B. (eds.). Introduction to bryophyte biology (2nd ed., pp. 33‒45). Cambridge University Press. [ Links ]

WFO 2023.WFO Plant List. Snapshots of the taxonomy.https://wfoplantlist.org/Links ]

Zepeda-Gómez, C., Ávila-Pérez, P., Díaz-García, U. S., Alanís-Martínez, Y., Zarazúa-Ortega, G. y Amaya-Chávez, A.2014.Diversidad de musgos epifitos de la zona metropolitana del valle de Toluca, México.Revista Mexicana de Biodiversidad, 85(1), 108‒124.10.7550/rmb.35456 [ Links ]

Annex 1. General characteristics of the moss species identified in the sampling sites of an Abies religiosa forest at Parque Nacional El Chico, Hidalgo. The description and information on the habitat and geographic distribution of the species were obtained from Sharp et al. (1994) and WFO (2023).

Bryum argenteum Hedw.: small, scarcely shiny, and julaceous plants. Caulids very thin, either simple or branched. Phyllids pressed against the caulid when dry but extend when wet. They are ovate, with a hyaline apex, somewhat acuminate and apiculate; the margins are entire, flat, or slightly reflexed. The costa terminates below the apex or may be excurrent, and the basal laminal cells are hexagonal or rectangular. The species is cosmopolitan. In Mexico, it is distributed throughout almost the entire country and is found on soil, rocks, cement walls, tiles, tree bark, and in open to semi-shaded sites.

Ceratodon purpureus (Hedw.) Brid.: The synonym is Ceratodon stenocarpus Bruch & Schimp. The plants are small to medium-sized and can exhibit colors ranging from dark green and yellowish-green to reddish tones. Caulids are erect, simple, or sparingly branched, with roots at the base. Phyllids are slightly appressed or twisted when dry, becoming erect when wet. They are lanceolate and gradually acuminate, with strongly reflexed or revolute margins, irregularly serrulate near the apex. The costa (midrib) is thick, percurrent to slightly excurrent. The upper and middle cells are square with slightly thick and smooth walls, while the basal cells are short and rectangular with fairly thin walls. This species is cosmopolitan. In Mexico, it is found in different habitats due to its broad ecological tolerance.

Cyrto-hypnum mexicanum (Mitt.) W. R. Buck & H. A. Crum: Thin plants of light green or yellowish color. Creeping caulids, irregularly pinnate or bipinnate; numerous paraphilia, two to five cells long, very short or linear, ending in a conical cell with one to two very small papillae. Phyllids strongly curved when dry, spreading when wet, broadly ovate, acuminate, and pleated near the base; margins sometimes narrowly recurved; yellowish costa ending near the apex; cells with thin walls and unipapillose on both surfaces. The species is mainly distributed in Mexico with some representatives in South America and grows in humid places on tree bark, trunks, stumps, and rocks.

Helicodontium capillare (Hedw.) A. Jaeger: Small plants on flat mats of opaque green color. Caulids with numerous branches, mostly short, slender and terete when dry. Phyllids spreading when wet, oblong-ovate-lanceolate to ovate and acute to short acuminate; margins crenulate to serrulate at top; costa occupying 2/3 to 3/4 of phyllid length; lamellar cells rhomboidal to hexagonal, smooth, becoming longer toward base of phyllid; wing cells square arranged in four to six rows. When plants grow on habitats such as rocks or decaying logs, the apex of the phyllids is usually large and the costa relatively short. Its main area of distribution is tropical America. In Mexico it grows on soil and humus in humid forests.

Thuidium delicatulum (Hedw.) Schimp var. delicatulum : Plants are robust green or yellowish brown. Two to three pinnate and somewhat leafy caulids; paraphilia abundant, usually papillose at cell ends. Phyllids of main axis erect, adpressed when dry, and erect and spreading when moist, triangular-ovate, not plicate; margins revolute, papillose-serrulate; apex acuminate; costa terminating before apex; distal lamellar cells irregularly oblong-hexagonal, unipapillose abaxially and papillae usually curved and somewhat bifurcate. Phyllids of the secondary branches are similar to those of the main branches, but smaller. Its distribution is mainly Pantropical, but it can be found in the Holarctic region. In Mexico it grows mainly in moist, shady soil, humus, rocks, logs or stumps, less frequently on the bark at the base of trees or even on trunks in particularly humid places.

Thuidium delicatulum var. peruvianum (Mitt.) H. A. Crum: Robust, dull green, yellow or brown plants. One to three creeping to ascending caulids, usually curved and pinnate. Main axis phyllids about 2 mm long, strongly plicate, abruptly narrowed to an often-falcate acumen; papillae simple and curved. Perichaeal phyllids ciliate. The species is distributed mainly in tropical America. In Mexico it is found on soil and rocks.

Thuidium delicatulum var. radicans (Kindb.) H. A. Crum, Steere & L. E. Anderson. Robust, dull green, yellow or brown plants. One to three creeping to ascending caudils, usually curved and pinnate. Main axis phyllids 1-1.5 mm long, ending in a slender tip composed of two to eight cells in a single row. The perichaeal phyllids are usually not ciliated, rarely present cilia, but are not numerous and not well developed. It is a cosmopolitan species. In Mexico it is found growing mainly on humus and soil, although it can also grow on other substrates.

Received: May 18, 2023; Accepted: January 29, 2024

*Corresponding author: arturosg@uaeh.edu.mx; tel.: +52 771 717 2000 ext. 6676.

Creative Commons License This is an open-access article distributed under the terms of the Creative Commons Attribution License