Delgadillo-Moya and Peña-Retes: Grimmia (Grimmiaceae, Bryophyta) in southern Mexico (Oaxaca and Chiapas)
Vol. , Num. 126, Año. 2018
Recibido: 2018 06 14
Aceptado: 2018 09 14



Introduction

The geographical distribution of most species of Grimmia Hedw. (Grimmiaceae, Bryophyta) in the New World is well known due to field work and taxonomic revisions at continental and global scales (Greven, 1999, 2003; Muñoz, 1999; Muñoz and Pando, 2000; Delgadillo, 2015). However, despite this upgraded information, there are still questions on the origin of the discontinuous range of various species. Most of the 26 species recognized for the Neotropics are unknown from the area between central Mexico and northern South America (e.g., Grimmia donniana Sm. and G. fuscolutea Hook.). This geographical discontinuity is even larger for those species that occur in northern Mexico and northern/southern South America (G. anodon Bruch & Schimp., G. laevigata (Brid.) Brid., and G. pulvinata (Hedw.) Sm.). Their apparent absence in southern Mexico and Central America may be due to a deficient collection record or to the lack of suitable habitats. At least six species are known from Central America (Muñoz and Allen, 2002; Delgadillo, 2015).

Models of potential distribution for Grimmia indicated that some species were expected to occur in southern Mexico (Delgadillo et al., 2012), but only G. longirostris Hook., G. pilifera P. Beauv., and G. trichophylla Grev. have been recorded for this area. The study of more than 900 specimens from Mexico (Delgadillo, 2015) showed that the Neovolcanic Belt is the southernmost limit for most Mexican species of Grimmia, suggesting that the genus and its alpine species reached the Neovolcanic Belt area by migration from North America (Delgadillo, 1987). Hypothetically, the discovery of other species in the highlands of Oaxaca, and in central or southern Chiapas would suggest an alternate southern origin of Grimmia in Mexico. This contribution reports on recent field exploration in Oaxaca and Chiapas, updates the geographical distribution of the genus and the species in southern Mexico, and evaluates the hypothesis of its geographical derivation.

Materials and Methods

Field work was conducted in the highlands of Oaxaca and Chiapas in 2013, 2015 and 2018. Emphasis was placed on the search of Grimmia to validate previous models (Delgadillo et al., 2012) and offer support to an alternate hypothesis regarding a southern origin of Mexican species. Records of the specimens collected were added to a database originally prepared for a revision of the genus in the Neotropics (Delgadillo, 2015). Localities visited for Grimmia are given in Table 1 along with a summary of identified specimens; the latter were deposited in the Bryophyte Collection at MEXU.

Table 1:

Specimens of Grimmia Hedw. collected in Oaxaca and Chiapas. AC=Ángeles Cárdenas; CD=Claudio Delgadillo; EH=Enrique Hernández; PP=Paola Peña; AJS=Aaron J. Sharp. All specimens were deposited in the Bryophyte Collection at MEXU.

Species State Locality/Specimens Coordinates
Grimmia longirostris Hook. Oaxaca Cerro Corral de Piedra: AC 4294; CD 4852 17°10'N - 96°39'W
Camino a Sta. María, ca. Mitla: CD 7553 16°55'N - 96°17'W
Grimmia pilifera P. Beauv. Oaxaca Ixtlán: EH 155 17°18'N - 96°29'W
Luvina: EH 687 17°30'N - 96°32'W
East of La Cumbre: AJS 2608 17°11'N - 96°36'W
Grimmia trichophylla Grev. Oaxaca Cerro Corral de Piedra: CD 4854 17°10'N - 96°39'W
Cerro Pelón: CD 7546 17°34'N - 96°30'W
El Cuartel: CD 7554 17°10'N - 96°37'W
Tres Ocotes: CD 7723 16°11'N - 96°26'W
Ozolotepec: CD 7730, 7733 16°07'N - 96°13'W
Grimmia trichophylla Grev. Chiapas Cerro Mozotal: CD 7841, 7855; PP 404, 419 15°26'N - 92°20'W

Identification of specimens was made using the keys and descriptions of publications cited in the introduction.

Results

According to our Grimmia database, the species are frequent in alpine grasslands, coniferous or Quercus forests. It is a group of high elevation elements representing a broad spectrum of environmental conditions present in xerophytic scrubby vegetation, dry pinyon pine forests, pine-oak-juniper woodlands, tropical deciduous forests, and grasslands. In the northern states Grimmia is found at elevations of 500-600 m a.s.l., while in central Mexico some species may reach 4600 m. Samples of Grimmia from Oaxaca and Chiapas were obtained at 2950-3270 m a.s.l., but the altitudinal range for the genus there may be as low as 1944 m a.s.l. Exploration in Pinus forests yielded G. longirostris, G. pilifera, and G. trichophylla in the state of Oaxaca. The last one is the only species of the genus known from Chiapas. The specimens listed in Table 1 and those cited by Delgadillo and Cárdenas (1989) from the same area represent the first records of the genus for Chiapas. All of them were collected in a disturbed Pinus forest on top of the Cerro Mozotal.

While the pine forest of the summit of Cerro Mozotal seems the habitat where Grimmia would be expected to occur, the accompanying moss flora is usually part of other plant communities. Braunia imberbis (Sm.) N.J. Dalton & D.G. Long, Polytrichastrum tenellum (Müll. Hal.) G.L. Sm., and Racomitrium subsecundum (Hook. & Grev. ex Harv.) Mitt. & Wilson are alpine/subalpine species; Caribaeohypnum polypterum (Mitt.) Ando & Higuchi, Dendropogonella rufescens (Schimp.) E. Britton, and Prionodon spp. are examples from cloud forests (Table 2).

Table 2:

Mosses from the summit of Cerro Mozotal associated with Grimmia trichophylla Grev. and their main habitat in Mexico. Species marked with an asterisk (*) are new records for Chiapas. CD=Claudio Delgadillo; DEB=Dennis E. Breedlove; PP=Paola Peña. All specimens were deposited in the Bryophyte Collection at MEXU.

Species Specimens Habitat
Anoectangium aestivum (Hedw.) Mitt. CD 7862 Coniferous forests
Anomobryum julaceum (Schrad. ex G. Gaertn., B. Mey. & Schreb.) Schimp. PP 424a Coniferous forests
Bartramia longifolia Hook. PP 423 Cloud forest/subalpine
Braunia imberbis (Sm.) N.J. Dalton & D.G. Long CD 7840 Alpine/subalpine
Breutelia subarcuata (Müll. Hal.) Schimp. CD 7833 Coniferous forests
Bryum billarderi Schwägr. DEB 68819; PP 410 Pine forests and oak forests
Campylopus pilifer Brid. CD 7852 Pine-oak forests
Caribaeohypnum polypterum (Mitt.) Ando & Higuchi CD 7836 Cloud forest
Dendropogonella rufescens (Schimp.) E. Britton CD 7847 Cloud forest
*Dicranum scoparium Hedw. CD 7834, 7838 Coniferous forests
Didymodon rigidulus Hedw. var. icmadophilus (Schimp. ex Müll. Hal.) R.H. Zander PP 424b, CD 7860 Oak forests
Entodon hampeanus Müll. Hal. PP 409 Cloud forest
Herzogiella cylindricarpa (Cardot) Z. Iwats. CD 7851 Coniferous forests
*Leptodontium capituligerum Müll. Hal. PP 407 Coniferous forests
Leptodontium flexifolium (Dicks.) Hampe CD 7844, 7846 Alpine
Loeskeobryum brevirostre (Brid.) M. Fleisch. CD 7857 Cloud forest
Macromitrium longifolium (Hook.) Brid. PP 420, 425 Cloud forest
Mittenothamnium reptans (Hedw.) Cardot PP 421a Cloud/Coniferous forests
Pilotrichella flexilis (Hedw.) Ångstr. PP 422, CD 7849 Cloud forest
Plagiothecium drepanophyllum Renauld & Cardot PP 411 Coniferous forests
Polytrichastrum tenellum (Müll. Hal.) G.L. Sm. CD 7839 Alpine/subalpine
Prionodon fuscolutescens Hampe CD 7848 Cloud forests
Prionodon luteovirens (Taylor) Mitt. CD 7858 Cloud forests
*Racomitrium subsecundum (Hook. & Grev. ex Harv.) Mitt. & Wilson CD 7837, 7854 Alpine/subalpine
Rozea andrieuxii (Müll. Hal.) Besch. CD 7845 Coniferous forests
Thuidium assimile (Mitt.) A. Jaeger PP 421b Coniferous forests

Discussion

The number of species of Grimmia detected in southern Mexico is surprisingly low, particularly in the states of Oaxaca and Chiapas that exhibit topographic elevations and types of vegetation that seem regular habitats for several species elsewhere. According to their known altitudinal range in Mexico and their potential distribution (Delgadillo et al., 2012), at least eight species would be expected in the area, but field work revealed only three of them (Table 1). Grimmia elongata Kaulf., G. mexicana Greven, and G. ovalis (Hedw.) Lindb. have been reported from northwestern Guatemala; G. longirostris and G. trichophylla have a broader range in Central America. There are no records of G. fuscolutea, G. pulla Cardot, and G. torquata Drumm. All three species form spores or gemmae that would enable them to spread into available habitats, but their consistent absence in southern Mexico suggests that other environmental or historical variables control their establishment.

The peculiar habitat of G. trichophylla in Chiapas is of interest; high altitude pine forests are the usual vegetation for this taxon. However, on the Cerro Mozotal it dwells with other alpine or subalpine mosses and species from deciduous or tropical derivation. Graham (1973) has shown that members of the arborescent temperate element migrated into Mexico since the Eocene and moved progressively southwards; Abies Mill., Picea A. Dietr., Alnus Mill., Fagus L., Juglans L., Liquidambar L., and others reached southern Mexico by the mid-Miocene. This fact may explain the presence of temperate mosses in Chiapas, i.e., species that moved along with the vascular plant flora; the tropical taxa perhaps derive from montane forests from lower elevations. The presence of the alpine and subalpine mosses (Table 2), including some species of Grimmia, may be related to high elevation habitats in Chiapas that may have become available by the late Tertiary (cf. López Ramos, 1983) and were filled by new arrivals.

Field and herbarium work resulted in more than 900 Mexican specimens for study (Delgadillo, 2015). These specimens and bibliographic information show that most species of Grimmia are unknown from southern Mexico and Central America. The lower altitude of the Central American mountains, relative to those in the Neovolcanic Belt and the northern Andes, would not facilitate the spread of Grimmia from the north or from the south. However, since the distribution of most species of that genus does not extend southwards beyond the Neovolcanic Belt one must conclude that the original hypothesis of a northern origin in Mexico is the most suitable explanation. The broad continental ranges of G. longirostris and G. trichophylla do not give definite support to a northern or southern origin, but the northern option is preferred because the Isthmus of Panama is considered a barrier, not a corridor, for high montane elements (Simpson and Neff, 1985). According to these authors, even a 1200 m lowering of the vegetational belts would not provide continuous cool high forest areas between North and South America. Present distribution of these two species is perhaps associated with lower mountain habitats unlike other species in the Mexican highlands. Undoubtedly, additional geological, geographic, and floristic information is required to understand the history of the area; a sample of our incomplete knowledge is illustrated by the new records for the Chiapas moss flora as given in Table 2.

Author contributions

CDM and PPR participated in field work and specimen identification. CDM prepared the manuscript assisted by PPR.

Acknowledgments

Thanks are extended to Francisco Hernández Najarro, Instituto de Historia Natural de Chiapas, for field and site information following earthquakes in Chiapas. An anonymous reviewer made valuable comments to the original manuscript.

Literature cited

1

Delgadillo, M. C. 1987. Moss distribution and the phytogeographical significance of the Neovolcanic Belt of Mexico. Journal of Biogeography 14(1): 69-78. DOI: https://dx.doi.org/10.2307/2844787

2

Delgadillo, M. C. 2015. Grimmia (Grimmiaceae, Bryophyta) in the Neotropics. Instituto de Biología, Universidad Nacional Autónoma de México. México, D.F., México. 88 pp.

3

Delgadillo M. C., and Á. Cárdenas S. 1989. Phytogeography of high-elevation mosses from Chiapas. The Bryologist 92(4): 461-466. DOI: https://dx.doi.org/10.2307/3243666

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Delgadillo M., C., J. L. Villaseñor and E. Ortiz. 2012. The potential distribution of Grimmia (Grimmiaceae) in Mexico. The Bryologist 115(1): 12-22. DOI: https://dx.doi.org/10.1639/0007-2745-115.1.12

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Graham, A. 1973. History of the arborescent temperate element in the northern Latin American Biota. In: Graham, A. (ed.). Vegetation and vegetational history of northern Latin America. Elsevier Scientific Publishing Company. Amsterdam, The Netherlands. Pp. 301-314.

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Greven, H. C. 1999. A synopsis of Grimmia in Mexico, including Grimmia mexicana, sp. nov. The Bryologist 102(3): 426-436. DOI: https://dx.doi.org/10.2307/3244231

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Greven, H. C. 2003. Grimmias of the World. Backhuys Publishers. Leiden, The Netherlands. 247 pp.

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López Ramos, E. 1983. Geología de México. Tomo III, 3ª ed. México, D.F., México. Pp. 263-264.

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Muñoz, J. 1999. A revision of Grimmia (Musci, Grimmiaceae) in the Americas. 1: Latin America. Annals of the Missouri Botanical Garden 86(1): 118-191. DOI: https://dx.doi.org/10.2307/2666219

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Muñoz, J. and B. H. Allen. 2002. Grimmia. In: Allen, B. H. (ed.). Moss flora of Central America. Part 2. Encalyptaceae-Orthotrichaceae. Monographs in Systematic Botany from the Missouri Botanical Garden 90: 221-236.

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Muñoz, J. and F. Pando. 2000. A world synopsis of the genus Grimmia (Musci, Grimmiaceae). Monographs in Systematic Botany from the Missouri Botanical Garden 83: 1-133.

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Simpson, B. B. and J. L. Neff. 1985. Plants, their pollinating bees, and the Great American Interchange. In: Stehli, F. G. and S. D. Webb (eds.). The Great American Biotic Interchange. Plenum Press. New York, USA. Pp. 427-452. DOI: https://dx.doi.org/10.1007/978-1-4684-9181-4_16

To cite as:

1 Delgadillo-Moya, C. and P. Peña-Retes. 2018(2019). Grimmia (Grimmiaceae, Bryophyta) in southern Mexico (Oaxaca and Chiapas). Acta Botanica Mexicana 126: e1408. DOI: 10.21829/abm126.2019.1408

Funding

2 The Departamento de Botánica, Instituto de Biología, Universidad Nacional Autónoma de México, provided funds for field and laboratory work.



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