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Inicio Revista Mexicana de Biodiversidad Diversity and distribution of mosses in the state of Hidalgo, Mexico
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Vol. 85. Núm. 1.
Páginas 84-97 (Marzo 2014)
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Vol. 85. Núm. 1.
Páginas 84-97 (Marzo 2014)
DOI: 10.7550/rmb.35761
Open Access
Diversity and distribution of mosses in the state of Hidalgo, Mexico
Diversidad y distribución de musgos en el estado de Hidalgo, México
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Claudio Delgadillo, José Luis Villaseñor, Ángeles Cárdenas, Enrique Ortiz
Departamento de Botánica, Instituto de Biología, Universidad Nacional Autónoma de México, Apartado postal 70-233, 04510 México, D. F., Mexico.
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Tablas (3)
Table 1. Moss taxa (371) in the state of Hidalgo, supported by herbarium specimens at MEXU. Neovolcanic Belt (NVB) and Eastern Sierra Madre (ESM) species are indicated by an X
Table 2. Moss taxa listed in Sharp et al. (1994), not supported by specimens at MEXU. Endemic taxa are indicated by “+”
Table 3. Summary of number of species listed in Table 1. NVB=Neovolcanic Belt; ESM=Eastern Sierra Madre
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Abstract

Through field work, bibliographic information and herbarium collections, a preliminary list of mosses for the state of Hidalgo was compiled. Records for 355 species were supported by 3 068 herbarium specimens at MEXU; when varieties are included along with taxa unsupported by local herbarium specimens, the number of taxa reaches 420. A collecting effort analysis indicates that 74.5% of the moss flora has been surveyed, that is, 56 taxa remain to be added. Species distribution modeling using 20 climatic variables from the WorldClim database for 150 species produced a map of potential distribution using a 5-minute cell network; it shows that the species potential richness is higher in central and southeastern Hidalgo, although most collections were obtained in southern and northwestern stations. Because large portions of the state land area are underexplored for mosses, no biodiversity hotspots are recognized. The Caribbean element is best represented in the Eastern Sierra Madre, but the confluence of the latter with the Neovolcanic Belt does not seem to show other major floristic differences between them, despite their geographical proximity.

Key words:
richness patterns
collecting effort
potential distribution
Maxent
Resumen

Se presenta una lista preliminar de musgos del estado de Hidalgo basada en trabajo de campo, bibliografía y colecciones de herbario. Con base en 3 068 ejemplares en MEXU se registran 355 especies, pero esta cifra se incrementa a 420 si se incluyen las variedades y los taxa sin registros en los herbarios locales. El análisis del esfuerzo de recolecta señala que se ha registrado el 74.5% de la riqueza estatal, es decir, 56 taxa menos del valor esperado. Los modelos de distribución potencial de 150 especies, usando 20 variables climáticas de la base de datos de WorldClim y una división en celdas de 5minutos, indica que la riqueza potencial de musgos es más alta en el centro y sureste del estado, a pesar de que la mayoría de las colecciones provienen de sitios del sur y noroeste. Como partes importantes del centro del estado todavía están poco exploradas, no se reconocen zonas de alta diversidad. No se han detectado diferencias en patrones florísticos, excepto en el elemento del Caribe que está mejor representado en la sierra Madre Oriental que en el Eje Neovolcánico, a pesar de la cercanía geográfica de las 2 áreas.

Palabras clave:
patrones de riqueza
esfuerzo de colecta
distribución potencial
Maxent
Texto completo
Introduction

Mosses of the state of Hidalgo have been collected and studied since the beginning of the XX century. Among the early collections are those obtained by Cyrus G. Pringle through several visits to the state (Davis, 1936). His moss collections were sent to J. Cardot (1909, 1910, 1911) for identification. In mid-XX century, Crum (1951) cited specimens by various collectors, including A. J. Sharp and his collaborators, referring to them by collector name and number; most of these specimens were deposited in MICH and other American herbaria. Sharp et al. (1994) included 281 moss taxa for Hidalgo, but the Moss Flora of Mexico did not cite specimens. In recent years, Alfaro and Castillo (1986) listed 169 species and varieties for Sierra de Pachuca; Cárdenas and Delgadillo (2009) cited specimens from localities bordering the Valley of Mexico that politically belong in Hidalgo; Delgadillo et al. (2011) listed 129 species and varieties from Los Mármoles National Park. The specimens derived from the last 3 contributions were deposited in the Bryophyte Collection at the National Herbarium (MEXU).

Despite the floristic information available, it seems that many areas in Hidalgo have not been explored for mosses, some sites are represented by many collections, and that a broader selection of sites should provide an adequate representation of the state’s moss flora. Because of its geological and geographical setting, especially at the point of contact between the Eastern Sierra Madre (ESM) and the Neovolcanic Belt (NVB), collections in the state of Hidalgo along with other nearby areas may be informative of the history of moss migration in this part of Mexico. In this contribution we offer a preliminary assessment of actual and potential species richness, and patterns of distribution; these may be in order to plan future field work in various parts of the state.

The state of Hidalgo in eastern Mexico has a surface area of nearly 21 000km2 (García and Falcón, 1984). Its rugged relief is dominated by the Eastern Sierra Madre that runs NW-SE; numerous sierras and isolated mountains are found in southern and western areas, several of them reaching more than 3 000m in elevation (cf. INFDM, 2005). Because of its geographical position, the state is also part of the Neovolcanic Belt area.

Materials and methods

Recently collected specimens and samples deposited in MEXU were examined along with records from the literature to produce a list of moss species from the state of Hidalgo. Major literature sources of floristic and geographical information were the updated electronic version of LATMOSS 2010 (Delgadillo, 2010) and Sharp et al. (1994) that complemented specimen data. The information for 3 068 moss specimens was compiled in a georreferenced database with records for 355 species that served to calculate cell width of the area of occupancy (AOO), according to IUCN criteria (IUCN, 2001). In this study, cell size is the longest axis between 2 collecting points divided by 10; the size of the grids (area of occupancy) was calculated by using the Conservation Assessment Tools designed for Arcview (Moat, 2007). The species cell width was averaged to obtain the width value applicable to all species; the value thus obtained, 8.4km, was transformed to arc minutes (about 5minutes). For further analysis, the state of Hidalgo was then divided into a network of 5-minute cells.

Collecting effort

The geographical data of the collecting records were used to produce a species accumulation curve (Gotelli and Colwell, 2001). Seventy-seven 5-minute cells with collecting records were used for the analysis. The asymptote of the accumulation curve (Fig. 1) is theoretically related to the number of species expected for the study area (Jiménez-Valverde and Hortal, 2003) and the number of cells is a measure of the collecting effort after randomly sorting these 50 times to produce a soft curve with EstimateS, version 8.2.0 (Colwell, 2009). The asymptote was estimated adjusting Clench’s equation to the accumulation curve (Soberón and Llorente, 1993; Colwell and Coddington, 1994) by the Simplex and Quasi-Newton method in the STATISTICA software (StatSoft, 2011); the predicted asymptotic value was used to estimate the precision of the inventory.

Figure 1.

Accumulation curve for moss species in the state of Hidalgo. The circles represent sampling units (5 arc-minute longitude/latitude cells). Curve parameters are indicated in the equation on the upper end.

(0,09MB).
Known species richness

The collecting data for 3 068 records were placed in the 5-minute cell network to identify the number of species per geographic unit and to produce a known species richness map (Fig. 2).

Figure 2.

Known diversity is given for each 5-minute latitude/longitude cell. Readings were made at each collecting point intersection. Color scales indicate the number of species recorded.

(0,16MB).
Species distribution modeling

Twenty variables were thought to have potential predictive value for the distribution of plant species. Nineteen bioclimatic variables were obtained from the WorldClim website (Hijmans et al., 2005; http://www.worldclim.org/bioclim.htm) while the values of the twentieth, altitude, were obtained from an elevation digital model in the WorldClim website; spatial resolution was 1km2 for all variables.

Maxent was used to prepare models of potential distribution of 150 species (Phillips et al., 2006; Phillips and Dudik, 2008) each with at least five collecting records that were considered sufficient to obtain a reliable model. With Maxent’s predetermined configuration, 75% of the records were used in model’s training and the remaining 25% for the validation of the model. Models generated by Maxent show a logistic probability of 0.000 to 1.000 that may be transformed in presence-absence Boolean area maps by applying thresholds, i.e., all pixel values higher than the selected threshold are classified as “1” while a “0” value is given to the remaining pixels. The optimum threshold value has not been adequately established in Maxent (Phillips et al., 2006), however, in this contribution the threshold used was the logistic value equivalent to a 10% omission error to maintain a high proportion of correctly predicted presence records, as applied in Pearson et al. (2007), Suarez-Seoane et al. (2008), and Kumar and Stohlgren (2009).

Potential richness

Distribution models for 150 species were placed in the 5-minute cell network to indicate the cells occupied by each species. A new richness map was produced with the models thus prepared and the records for 205 species without models supplemented the richness data for each cell. This information was used to prepare a map of potential richness (Fig. 3).

Figure 3.

Patterns of richness based on potential distribution models for 150 moss species and point records for the 205 remaining species. No varieties were included. Cell size is 5-minute latitude/longitude.

(0,4MB).
Results

Herbarium and literature records produced a list of 420 species for the state of Hidalgo (Tables 1, 2). The species accumulation curve suggests that the moss flora of the state of Hidalgo should contain about 476 species, as indicated by the asymptote (Fig. 1). Since the presence of 355 species has been documented, the level of completeness of the flora under study is 74.5%. However, computation did not consider the varieties recognized for several species so that the actual number of taxa would be 371 (Table 1). Sharp et al. (1994) recorded 49 additional taxa (Table 2) that were represented in herbarium collections elsewhere in the world and should be expected to complement holdings in MEXU. This means that there are still 56 taxa lacking to fulfill the model prediction. The 420 moss taxa known from the state (Tables 1, 2) represent a high floristic number for the area. To be sure, published data indicate that such neighboring states as Guanajuato harbor 114 moss species and varieties (Delgadillo and Cárdenas, 1996), México 268 (Sharp et al., 1994), and Querétaro 212 (Herrera et al., 2008); it seems that a higher number may be found once the central dry lands and other forested areas are explored.

Table 1.

Moss taxa (371) in the state of Hidalgo, supported by herbarium specimens at MEXU. Neovolcanic Belt (NVB) and Eastern Sierra Madre (ESM) species are indicated by an X

Taxon/Element  NVB  ESM 
Northern     
Anomobryum filiforme var. concinnatum (Spruce) Boul.   
Anomodon attenuatus (Hedw.) Huebener   
Anomodon rostratus (Hedw.) Schimp.   
Anomodon thraustus Müll. Hal.   
Atrichum angustatum (Brid.) Bruch and Schimp. 
Barbula indica (Hook.) Spreng.   
Bryum erythroloma (Kindb.) Syed   
Campyliadelphus chrysophyllus (Brid.) Kanda   
Campylophyllum sommerfeltii (Myr.) Hedenäs 
Campylopus fragilis (Brid.) Bruch and Schimp.   
Ceratodon purpureus subsp. stenocarpus (Bruch and Schimp.) Dixon 
Claopodium pellucinerve (Mitt.) Best   
Dicranella varia (Hedw.) Schimp.   
Dicranoweisia cirrata (Hedw.) Lindb.   
Dicranum flagellare Hedw. 
Diphyscium foliosum (Hedw.) Mohr   
Drepanocladus sordidus (Müll. Hal.) Hedenäs in W. R. Buck   
Encalypta ciliata Hedw.   
Entodon schleicheri (Schimp.) Demeter   
Fissidens dubius P. Beauv.   
Grimmia pilifera P. Beauv. 
Haplocladium angustifolium (Hampe and Müll. Hal.) Broth. 
Haplocladium microphyllum (Hedw.) Broth. 
Heterophyllium affine (Hook. ex Kunth) M. Fleisch. 
Hygroamblystegium fluviatile (Hedw.) Loeske   
Isopterygium tenerum (Sw.) Mitt.   
Kindbergia praelonga (Hedw.) Ochyra   
Mnium marginatum (With.) P. Beauv.   
Molendoa sendtneriana (Bruch and Schimp.) Limpr.   
Orthodontium gracile Schwägr. ex B.S.G.   
Oxyrrhynchium pringlei (Cardot) J. T. Wynns   
Plagiomnium cuspidatum (Hedw.) T. Kop.   
Platygyrium fuscoluteum Cardot 
Pylaisia polyantha (Hedw.) Schimp. 
Pylaisia selwynii Kindb.   
Rhodobryum roseum (Hedw.) Limpr. 
Rhynchostegium pulchellum (Hedw.) H. Rob. 
Rhynchostegium riparioides (Hedw.) Cardot 
Rhynchostegium serrulatum (Hedw.) A. Jaeger 
Rhytidium rugosum (Hedw.) Kindb. 
Sematophyllum marylandicum (Müll. Hal.) E. Britton   
Syntrichia fragilis (Taylor) Ochyra 
Syntrichia ruralis (Hedw.) Web. and Mohr   
Taxiphyllum deplanatum (Bruch and Schimp. ex Sull.) M. Fleisch.   
Thuidium delicatulum (Hedw.) Schimp. var. delicatulum 
Thuidium delicatulum var. radicans (Kindb.) Crum, Steere and Anders. 
Timmia megapolitana Hedw. var. bavarica (Hessl.) Brid.   
Tortella tortuosa (Hedw.) Limpr. 
Tortula acaulon (With.) R. H. Zander   
Trichostomum crispulum Bruch 
Trichostomum tenuirostre (Hook. and Taylor) Lindb. 
Weissia condensa (Voit) Lindb.   
Zygodon viridissimus (Dicks.) Brid.   
Meso-American     
Aloina hamulus (Müll. Hal.) Broth. 
Aloinella catenula Cardot   
Anomobryum plicatum Cardot   
Atractylocarpus flagellaceus (Müll. Hal.) Williams 
Atrichum oerstedianum (Müll. Hal.) Mitt. 
Bartramia potosica Mont.   
Brachymenium spirifolium (Müll. Hal.) A. Jaeger   
Brachythecium cirriphylloides McFarland 
Brachythecium conostomum (Taylor) A. Jaeger   
Brachythecium occidentale (Hampe) A. Jaeger 
Braunia andrieuxii Lorentz   
Braunia squarrulosa (Hampe) Müll. Hal. 
Bryoerythrophyllum recurvirostrum (Hedw.) Chen var. aeneum 
Bryum chryseum Mitt.   
Bryum procerum Schimp. 
Bryum richardsii Sharp 
Campylopus anderssonii (Müll. Hal.) A. Jaeger 
Campylopus reflexisetus (Müll. Hal.) Broth.   
Catagonium brevicaudatum Broth.   
Cryphaea apiculata Schimp.   
Cyclodictyon erubescens E. B. Bartram   
Cyclodictyon humectatum Cardot   
Cyrto-hypnum mexicanum (Mitt.) W. R. Buck and H. A. Crum   
Dicranum frigidum Müll. Hal. 
Didymodon hampei R. H. Zander   
Didymodon rigidulus var. subulatus(Thér. and E. B. Bartram) R. H. Zander 
Entodon jamesonii (Taylor) Mitt. 
Epipterygium immarginatum Mitt.   
Fissidens excurrentinervis Williams   
Flowersia campylopus (Schimp. ex Müll. Hal.) Griffin and W. R. Buck 
Globulinella globifera (Hampe) Steere ex Steere and Chapm.   
Herzogiella cylindricarpa (Cardot) Iwats.   
Horridohypnum mexicanum (Thér.) W. R. Buck 
Leptodontium viticulosoides (P. Beauv.) Wijk and Margad. var. exasperatum(Cardot) R. H. Zander 
Leskea angustata Taylor 
Leucodon cryptotheca Hampe 
Leucodon curvirostris Hampe 
Lindbergia mexicana (Besch.) Cardot 
Macromitrium fragilicuspis Cardot   
Meteorium teres Mitt.   
Mironia ehrenbergiana (Müll. Hal.) R. H. Zander   
Mironia stenotheca (Thér.) R. H. Zander   
Neckera chlorocaulis Müll. Hal. 
Neckera ehrenbergii Müll. Hal. 
Orthostichella pachygastrella (Müll. Hal. ex Aongstr.) B. H. Allen and Magill   
Orthotrichum bartramii Williams   
Orthotrichum pycnophyllum Schimp. ex Müll. Hal. 
Orthotrichum pycnophyllum var. verrucosum (Müll. Hal.) Lewinsky 
Physcomitrium subsphaericum Schimp.   
Platygyriella pringlei (Cardot) W. R. Buck   
Pohlia oerstediana (Müll. Hal.) Shaw   
Polytrichastrum tenellum (Müll. Hal.) G. Smith   
Pterobryopsis mexicana (Renauld and Cardot) M. Fleisch.   
Ptychomitrium serratum Bruch and Schimp. 
Rauiella lagoensis (Hampe) W. R. Buck   
Rhexophyllum subnigrum (Mitt.) Hilp. 
Rhynchostegium semiscabrum (E. B. Bartram) H. Rob.   
Rhynchostegium subrusciforme (Müll. Hal.) A. Jaeger   
Rozea andrieuxii (Müll. Hal.) Besch. Var. andrieuxii 
Rozea andrieuxii var. bourgeana (Besch.) W. R. Buck 
Sagenotortula quitoensis (Taylor in Hook.) R. H. Zander   
Schizymenium landii (Cardot) Shaw   
Schizymenium serratum (Cardot and Herz.) Shaw   
Sphaerotheciella pachycarpa (Schimp. Ex Besch.) Manuel   
Sphaerotheciella pinnata (Schimp.) Manuel 
Syntrichia amphidiacea (Müll. Hal.) R. H. Zander 
Syntrichia obtusissima (Müll. Hal.) R. H. Zander 
Thuidium delicatulum var. peruvianum (Mitt.) H. A. Crum 
Zygodon ehrenbergii Müll. Hal. 
Zygodon liebmannii Schimp. Ex Müll. Hal.   
Caribbean     
Archidium donnellii Austin   
Atrichum polycarpum (Müll. Hal.) Mitt. 
Bartramia brevifolia Brid.   
Brachymenium mexicanum Mont. 
Breutelia brittoniae Renauld and Cardot   
Breutelia inclinata (Hampe and Lorentz) A. Jaeger   
Breutelia subarcuata (Müll. Hal.) Schimp.   
Bryum limbatum Müll. Hal.   
Bryum pseudocapillare Besch.   
Campylopus albidovirens Herz.   
Campylopus tallulensis Sull. and Lesq. 
Caribaeohypnum polypterum (Mitt.) Ando and Hig.   
Chryso-hypnum salleanum (Besch.) W. R. Buck   
Cryphaea patens Hornsch. 
Ctenidium malacodes Mitt. 
Chryso-hypnum diminutivum (Hampe) W. R. Buck   
Daltonia longifolia Taylor   
Dicranum sumichrastii Duby 
Ditrichum rufescens (Hampe) Hampe   
Entodon beyrichii (Schwägr.) Müll. Hal. 
Entodon hampeanus Müll. Hal.   
Entosthodon obtusifolius Hook. F. in Hook. 
Erythrodontium longisetum (Hook.) Paris   
Fabronia ciliaris var. polycarpa (Hook.) W. R. Buck 
Fabronia ciliaris var. wrightii (Sull.) W. R. Buck 
Fabronia macroblepharis Schwägr.   
Fissidens crispus Mont. 
Fissidens elegans Brid.   
Fissidens polypodioides Hedw.   
Holomitrium arboreum Mitt.   
Homalia glabella (Hedw.) Schimp.   
Hypnum amabile (Mitt.) Hampe 
Isodrepanium lentulum (Wils.) Britt.   
Leptodontium viticulosoides var. sulphureum (Müll. Hal.) R. H. Zander 
Leucobryum albidum (Brid. ex P. Beauv.) Lindb. 
Leucobryum antillarum Schimp.   
Leucobryum polakowskii (Müll. Hal.) Cardot   
Leucodon julaceus (Hedw.) Sull.   
Leucoloma serrulatum Brid.   
Macromitrium cirrosum (Hedw.) Brid. 
Macromitrium guatemaliense Müll. Hal.   
Macromitrium longifolium (Hook.) Brid. 
Meteoridium remotifolium (Müll. Hal.) Man.   
Meteorium illecebrum Sull. 
Microcampylopus curvisetus (Hampe) Giese and J.-P. Frahm   
Neckera urnigera Müll. Hal.   
Papillaria deppei (Hornsch. ex Müll. Hal.) A. Jaeger   
Papillaria imponderosa (Taylor) Broth.   
Philonotis longiseta (Mx.) E. Britton   
Philonotis sphaericarpa (Hedw.) Brid.   
Phyllogonium fulgens (Hedw.) Brid.   
Pilopogon guadalupensis (Brid.) J.-P. Frahm   
Pilotrichella mauiensis (Sull.) A. Jaeger   
Pireella pohlii (Schwägr.) Cardot   
Plagiothecium drepanophyllum Renauld and Cardot   
Pleuridium mexicanum Cardot   
Pogonatum campylocarpum (Müll. Hal.) Mitt. 
Pogonatum tortile (Sw.) Brid.   
Pohlia richardsii Shaw   
Porotrichum korthalsianum (Dozy and Molk.) Mitt. 
Porotrichum longirostre (Hook.) Mitt. 
Porotrichum mutabile Hampe   
Pseudosymblepharis schimperiana (Paris) H. A. Crum 
Pterobryon densum Hornsch. 
Ptychomitrium lepidomitrium (Müll. Hal.) Schimp. in Besch. 
Rhodobryum beyrichianum (Hornsch.) Müll. Hal. ex Hampe 
Rhynchostegiopsis flexuosa (Sull.) Müll. Hal.   
Rhynchostegium scariosum (Taylor) A. Jaeger   
Rigodium toxarion (Schwägr.) A. Jaeger   
Schlotheimia jamesonii (Arnott) Brid.   
Schlotheimia rugifolia (Hook.) Schwägr.   
Sematophyllum cuspidiferum Mitt.   
Sematophyllum swartzii (Schwägr.) Welch and H. A. Crum 
Sphagnum meridense (Hampe) Müll. Hal.   
Splachnobryum obtusum (Brid.) Müll. Hal.   
Squamidium nigricans (Hook.) Broth.   
Syrrhopodon prolifer Schwägr.   
Thuidium tomentosum Schimp. 
Weissia jamaicensis (Mitt.) Grout   
Zelometeorium patulum (Hedw.) Manuel   
Zygodon campylophyllus Müll. Hal. 
Southern     
Braunia plicata (Mitt.) A. Jaeg. 
Bryum microimbricatum Ochi   
Bryum radiculosum Brid.   
Campylopus heterostachys (Hampe) A. Jaeger   
Erpodium beccarii Müll. Hal. ex Vent.   
Leptodontium capituligerum Müll. Hal. 
Orthotrichum aequatoreum Mitt. 
Rhacocarpus purpurascens (Brid.) Paris   
Wide distribution     
Aloina rigida (Hedw.) Limpr.   
Amphidium tortuosum (Hornsch.) Cufodontis   
Anacolia laevisphaera (Taylor) Flowers 
Andreaea rupestris Hedw.   
Anoectangium aestivum (Hedw.) Mitt. 
Anomobryum filiforme (Dicks.) Solms. in Rabenh. var. filiforme   
Anomodon tristis (Ces.) Sull. and Lesq.   
Aongstroemia orientalis Mitt. 
Barbella pendula (Sull.) M. Fleisch.   
Barbellopsis trichophora (Mont.) W. R. Buck   
Barbula arcuata Griff.   
Barbula bolleana (Müll. Hal.) Broth. 
Barbula convoluta Hedw.   
Brachymenium exile (Dozy and Molk.) Bosch and Sande Lac. 
Brachymenium systylium (Müll. Hal.) A. Jaeger 
Brachymitrion jamesonii Taylor 
Brachythecium plumosum (Hedw.) Schimp. 
Brachythecium ruderale (Brid.) W. R. Buck 
Braunia secunda (Hook.) Bruch and Schimp. 
Breutelia tomentosa (Brid.) A. Jaeg. and Sauerb.   
Bryoerythrophyllum campylocarpum (Müll. Hal.) H. A. Crum 
Bryoerythrophyllum inaequalifolium (Taylor) R. H. Zander   
Bryoerythrophyllum recurvirostrum var. recurvirostrum   
Bryum argenteum Hedw. 
Bryum billarderi Schwägr. 
Bryum capillare Hedw. 
Bryum muhlenbeckii Bruch and Schimp.   
Bryum pallescens Schleich. ex Schwägr.   
Bryum pseudotriquetrum (Hedw.) Gaertn., Meyer and Scherb.   
Bryum subapiculatum Hampe   
Campylopus flexuosus (Hedw.) Brid.   
Campylopus nivalis (Brid.) Brid. 
Campylopus pilifer Brid. 
Campylopus savannarum (Müll. Hal.) Mitt.   
Campylopus sinensis (Müll. Hal.) J.-P. Frahm   
Ceratodon purpureus (Hedw.) Brid. subsp. purpureus 
Crossidium crassinervium (De Not.) Jur.   
Cryphaea jamesonii Taylor   
Cyrto-hypnum minutulum (Hedw.) W. R. Buck and H. A. Crum   
Desmatodon convolutus (Brid.) Grout 
Didymodon australasiae (Hook. and Grev.) R. H. Zander 
Didymodon ferrugineus (Schimp. ex Besch.) M.O. Hill   
Didymodon revolutus (Cardot) Williams 
Didymodon rigidulus Hedw. var. gracilis (Schleich. ex Hook. and Grev.) R. H. Zander 
Didymodon rigidulus var. icmadophilus(Schimp. ex Müll. Hal.) R. H. Zander 
Didymodon rigidulus var. rigidulus 
Didymodon vinealis (Brid.) R. H. Zander   
Distichium capillaceum (Hedw.) Bruch and Schimp.   
Drepanocladus aduncus (Hedw.) Warnst.   
Entodon macropodus (Hedw.) Müll. Hal.   
Entosthodon muhlenbergii (Turner) Fife   
Eustichia longirostris (Brid.) Brid.   
Fabronia ciliaris (Brid.) Brid. var. ciliaris 
Fissidens asplenioides Hedw. 
Fissidens curvatus Hornsch.   
Fissidens pellucidus Hornsch.   
Fissidens submarginatus Bruch in Krauss   
Fissidens taxifolius Hedw.   
Fissidens weirii Mitt. var. hemicraspedophyllus(Cardot) Pursell   
Forsstroemia producta (Hornsch.) Paris 
Forsstroemia trichomitria (Hedw.) Lindb.   
Funaria hygrometrica Hedw. var. calvescens(Schwägr.) Mont. 
Funaria hygrometrica var. hygrometrica 
Grimmia longirostris Hook.   
Grimmia ovalis (Hedw.) Lindb.   
Grimmia trichophylla Grev.   
Groutiella tomentosa (Hornsch.) Wijk and Margad.   
Gymnostomum aeruginosum Sm. 
Hedwigia ciliata (Hedw.) P. Beauv. 
Hedwigidium integrifolium (P. Beauv.) Dixon   
Henicodium geniculatum (Mitt.) W. R. Buck   
Herpetineuron toccoae (Sull. and Lesq.) Cardot   
Homaliodendron flabellatum (Sm.) M. Fleisch.   
Hookeria acutifolia Hook. and Grev.   
Hymenostylium recurvirostrum (Hedw.) Dixon   
Hyophila involuta (Hook.) A. Jaeger 
Hypnum cupressiforme Hedw. var. cupressiforme 
Hypnum cupressiforme var. lacunosum Brid. 
Hypnum revolutum (Mitt.) Lindb.   
Hypopterygium tamarisci (Sw.) Brid. ex Müll. Hal.   
Leptobryum pyriforme (Hedw.) Wilson   
Leptodictyum riparium (Hedw.) Warnst.   
Leptodontium brachyphyllum Broth. and Thér.   
Leptodontium flexifolium (Dicks. ex With.) Hampe 
Leptodontium viticulosoides var. viticulosoides 
Leptohymenium tenue (Hook.) Schwägr.   
Lescuraea arizonae (R.S. Williams) P.S. Wilson and D.H. Norris 
Macrocoma orthotrichoides (Raddi) Wijk and Margad. 
Macrocoma tenuis (Hook. and Grev.) Vitt subsp. sullivantii 
Microbryum starkeanum (Hedw.) R. H. Zander   
Mittenothamnium reptans (Hedw.) Cardot 
Orthostichella rigida (Müll. Hal.) B. H. Allen and Magill 
Orthostichella versicolor (Müll. Hal.) B. H. Allen and Magill   
Orthotrichum anomalum Hedw.   
Orthotrichum diaphanum Schrad. ex Brid.   
Palamocladium leskeoides (Hook.) E. Britton 
Papillaria nigrescens (Sw. ex Hedw.) A. Jaeger 
Philonotis fontana (Hedw.) Brid.   
Philonotis glaucescens (Hornsch.) Broth.   
Philonotis marchica (Hedw.) Brid.   
Pilotrichella flexilis (Hedw.) Aongstr. 
Plagiomnium rhynchophorum (Hook.) T. Kop. 
Platygyriella densa (Hook.) W. R. Buck   
Pleuridium acuminatum Lindb.   
Pleurochaete squarrosa (Brid.) Lindb. 
Pogonatum oligodus (Müll. Hal.) Mitt.   
Pohlia cruda (Hedw.) Lindb.   
Pohlia elongata Hedw. 
Polytrichum commune L. ex Hedw.   
Polytrichum juniperinum Hedw. 
Porotrichum usagarum Mitt.   
Prionodon densus (Hedw.) Müll. Hal. 
Pseudocrossidium crinitum (Schultz) R. H. Zander   
Pseudocrossidium replicatum (Taylor) R. H. Zander 
Pylaisia falcata Schimp. 
Pylaisiadelpha tenuirostris (Bruch and Schimp.) W. R. Buck 
Pyrrhobryum spiniforme (Hedw.) Mitt.   
Racomitrium subsecundum (Hook. and Grev. ex Harv.) Mitt.   
Racopilum tomentosum (Hedw.) Brid. 
Rhabdoweisia fugax (Hedw.) Bruch and Schimp. 
Rhodobryum huillense (Welw. and Duby) Touw 
Schistidium apocarpum (Hedw.) Bruch and Schimp.   
Schistidium rivulare (Brid.) Podp.   
Sematophyllum adnatum (Mx.) E. Britton 
Sematophyllum galipense (Müll. Hal.) Mitt. 
Sematophyllum subpinnatum (Brid.) E. Britton 
Sematophyllum subsimplex (Hedw.) Mitt.   
Sphagnum palustre L. 
Sphagnum strictum Sull.   
Stereophyllum radiculosum (Hook.) Mitt.   
Symblepharis vaginata (Hook.) Wijk and Margad.   
Syntrichia chisosa (Magill, Delgad. and L. R. Stark) R. H. Zander   
Syntrichia pagorum (Milde) Amann   
Syntrichia papillosa (Wilson) Jur.   
Taxiphyllum taxirameum (Mitt.) M. Fleisch.   
Timmiella anomala (Bruch and Schimp.) Limpr. 
Tortella humilis (Hedw.) Jenn.   
Trachypus viridulus (Mitt.) Broth.   
Trematodon sp.   
Trichostomum brachydontium Bruch 
Weissia controversa Hedw. 
Zygodon obtusifolius Hook. 
Zygodon reinwardtii (Hornsch.) Braun   
Endemic     
Brachymenium saint-pierrei Thér.   
Didymodon incrassatolimbatus Cardot   
Entodon abbreviatus (Schimp.) A. Jaeger 
Grimmia involucrata Cardot   
Grimmia pulla Cardot   
Hennediella heteroloma (Cardot) R. H. Zander var. eckeliae R. H. Zander   
Homomallium sharpii Ando and Higuchi   
Jaffueliobryum arsenei (Thér.) Thér.   
Neckera angustifolia Müll. Hal.   
Oreoweisia delgadilloi H. Rob. and F. D. Bowers 
Pylaisiadelpha duellii H. A. Crum   
Synthetodontium pringlei Cardot   
Weissia semidiaphana (Thér.) R. H. Zander   
Chihuahuan     
Entosthodon apiculatopilosus (Cardot) Fife   
Homomallium mexicanum Cardot 
Weissia ligulifolia (E. B. Bartram) R. H. Zander   
Table 2.

Moss taxa listed in Sharp et al. (1994), not supported by specimens at MEXU. Endemic taxa are indicated by “+”

Taxon 
Acroporium longirostre (Brid.) W. R. Buck 
Aerobryopsis martinicensis (Broth.) Spessard-Schued. 
Anomobryum prostratum (Müll.Hal.) Besch. 
Aulacomnium palustre (Hedw.) Schwägr. 
Barbula orizabensis Müll.Hal. 
Brachymenium radiculosum (Schwägr.) Hampe 
Breutelia austroarcuata (Müll. Hal.) Paris 
Breutelia jamaicensis (Mitt.) A. Jaeger 
Bryoerythrophyllum recurvirostrum(Hedw.) Chen var. aeneum (Müll. Hal.) R. H. Zander 
Bryum apiculatum Schwägr. 
Bryum caespiticium Hedw. 
Bryum leptotorquescens Müll.Hal.ex Broth. 
Bryum microchaeton Hampe 
Calyptothecium duplicatum (Schwägr.) Broth. 
Chryso-hypnum salleanum (Besch.) W. R. Buck 
Cryphaea filiformis (Hedw.) Brid. 
Cyclodictyon albicans (Hedw.) Kuntze 
Daltonia tenuifolia Mitt. 
+Dicranella barnesii Cardot 
Dicranella hilariana (Mont.) Mitt. 
Dicranella lindigiana (Hampe) Mitt. 
+Dicranum lophoneuron Müll. Hal. 
Didymodon tophaceus (Brid.) Lisa 
Didymodon umbrosus (Müll. Hal.) R. H. Zander 
Entodon serrulatus Mitt. 
Entosthodon bonplandii (Hook.) Mitt. 
Epipterygium mexicanum (Besch.) Broth. 
Fissidens weirii Mitt.var. weirii 
Grimmia elongata Kaulf. 
Helicodontium capillare (Hedw.) A. Jaeger 
Heterocladium macounii Best 
Hyophiladelphus agrarius (Hedw.) R. H. Zander 
Luisierella barbula (Schwägr.) Steere 
Macromitrium punctatum (Hook. and Grev.) Brid. 
Neckeropsis undulata (Hedw.) Reichardt 
Philonotis cernua (Wilson) Griffin and W. R. Buck 
Philonotis uncinata (Schwägr.) Brid. 
Pilotrichella nudiramulosa Müll.Hal. 
Plaubelia sprengelii (Schwägr.) R. H. Zander var. stomatodonta (Cardot) R. H. Zander 
Pohlia papillosa (Müll. Hal. ex Jaeger) Broth. 
Pohlia pseudobarbula (Thér.) H. A. Crum ex A. J. Shaw 
Porotrichodendron lindigii (Hampe) W. R. Buck 
Pylaisiadelpha deplanatula (Cardot) W. R. Buck 
Rhachithecium perpusillum (Thwaites and Mitt.) Broth. 
Schoenobryum concavifolium (Griff.) Gangulee 
Streptopogon cavifolius Mitt. 
Streptopogon matudianus H. A. Crum 
Syntrichia bogotensis (Hampe) Mitt. 
Trachypodopsis serrulata (P. Beauv.) M. Fleisch. var. crispatula (Hook.) Zant. 

The list of species based on herbarium specimens (Table 1) also cites the taxa represented in the major mountain systems of the state, i.e., the Neovolcanic Belt with 245 taxa, and the Eastern Sierra Madre with 275. There are no strong differences in the number of taxa between mountain systems, but the Caribbean taxa are distinctly higher in the ESM (Table 3). Besides this, salient features of the moss flora include the large group of widely distributed and the Meso-American taxa. The endemic species, 12 in total, are not restricted to Hidalgo or to a topographic feature of the state and represent less than 3% of the entire moss flora. Because of their small number, it is also remarkable the presence of members of the Chihuahuan element.

Table 3.

Summary of number of species listed in Table 1. NVB=Neovolcanic Belt; ESM=Eastern Sierra Madre

Element  NVB  ESM  Total 
Northern  40  34  53 
Meso-American  53  50  70 
Caribbean  37  71  81 
Southern 
Wide distribution  98  107  143 
Endemic  10  13 
Chihuahuan 
Total  245  275  371 

The species listed in table 2 may follow the same patterns of distribution as those listed in table 1, but should be added when their state distribution is confirmed.

Figure 2 shows the number of known species per cell in the state of Hidalgo. The distribution of cells with data also indicate the extent of field work thus far conducted; there are numerous collections from SE and NW areas, followed in order of importance, by the NE and SW areas. These were obtained along major highways and forested areas along the way. The empty cells in the map represent dry lands or scattered peaks, inaccessible areas, and major cities and industrial areas. The potential distribution map for 150 species (Fig. 3) suggests that many of the empty cells in figure 2 may contain rich moss floras, especially in central and southeastern parts of the state. However, potential species richness values become smaller toward the northeastern lowlands and, in the southwest, toward the lower areas of the Neovolcanic Belt.

Discussion

The known moss flora of the state of Hidalgo is comparatively larger than that of adjacent states. Part of the size differences are undoubtedly due to insufficient bryological exploration in various states in central Mexico. The dry continental area north of the NVB and to the west of the ESM may indeed contain a reduced moss flora, but full diversity evaluations in Hidalgo require ample exploration in its central region. The size of the moss flora, with 420 species and varieties, is potentially more diverse as suggested in preceding paragraphs, but not as rich as that of Veracruz which includes more than 500 taxa (Delgadillo, 2011).

The presence of 2, NVB and ESM, groups of species of presumed different derivation might suggest a state flora with higher moss diversity or with peculiar geographical affiliation in the mountain areas. Neither hypothesis is confirmed by the results summarized in table 3; except for the values of the Caribbean element, the number of species in the NVB and in ESM is similar, perhaps due to the close proximity of the mountain ranges. The Caribbean species are an important constituent of the tropical floras of Mexico and these, along with the Meso-American and endemic taxa give a neotropical character to the flora of Hidalgo.

With respect to diversity, the number of species in Hidalgo is higher than in neighboring states, but no hotspots are readily identified with current data, although there are areas (e.g., between Zimapán and Jacala in the northwest, and in Sierra de Pachuca) where actual or potential species richness is higher (Figs. 2, 3). These areas, however, have been well collected and may not qualify for designation as hotspots.

The potential distribution map identifies areas of concentration of species; because of the habitat interrelationships between mosses and vascular plants, the map in figure 3 would be similar to the distribution of certain vascular plant communities. At present, it is unknown whether their distribution is comparable, but if field observations confirm this, mosses would be an additional criterion to justify conservation of diverse areas.

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