Botanice` Est Scientia Naturalis Quae Vegetabilium Cognitiorem Tradit
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Number 67 April 2, 2020 A CHECKLIST AND PRELIMINARY KEY TO THE LIVERWORTS OF NEW MEXICO Karen S. Blisard 25 Oxbow Drive, Silver City NM 88061 Email: [email protected] Russell Kleinman Department of Natural Sciences Western New Mexico University, Silver City NM 88061 Email: [email protected] ABSTRACT A checklist and preliminary key are presented for the 89 species of liverworts that have been reported from New Mexico. It is hoped that this information will serve to stimulate interest in these fascinating little plants. INTRODUCTION The bryophyte lineage is comprised of mosses, liverworts, and hornworts. Liverworts, also known as hepaticae due to the superficial resemblance of some of them to the mammalian liver, would seem to be poorly adapted to life in the arid southwest. However, this is only partially true, and this report describes the surprising diversity of liverworts in New Mexico. There are three basic morphologies of liverworts: leafy, simple thalloid, and complex thalloid (Hicks 1992; Schofield 1985, Vanderpooten & Goffinet 2009). Although leafy liverworts superficially resemble mosses, they can be distinguished by visual and microscopic examination. The leaves of leafy liverworts are basically in the same plane as the stem, whereas in mosses, the leaves are arranged radially around the stem. Liverwort leaves virtually never have a mid-vein (costa); mid-veins are found in a majority of moss leaves. Liverwort leaves are often divided into multiple lobes; this characteristic is rare in mosses. Most leafy liverworts have microscopic oil bodies within their cells, which can be important in species identification; mosses never contain oil bodies. Although oil bodies can be transient and are best examined in fresh specimens, they often persist for months in specimens collected from arid environments. Thalloid liverworts are comprised of elongate thalli instead of leaves and stems. Simple thalloid liverworts have an undifferentiated thallus which is quite thin, often only one cell thick. Simple thalloid liverworts are very uncommon in New Mexico. Complex thalloid liverworts have thick fleshy thalli whose cross-sections show differentiated layers with air pores. They are generally drought tolerant (Schuster 1992a), and some are widespread in New Mexico. METHODS We reviewed the existing literature on New Mexico liverworts (Arsene 1933; Bird 1960; Evans 1922; Frye and Clark 1937-1947; Guerke 1971; Hong 1989; Hong 1992; Ireland et al. 1981; Little 1937; Little 1942; Prior 1969; Romig 2012; Shields 1954; Standley 1915; Standley 1916; Stark & Casstetter 1982, Whittemore 1995; Worthington 2001). Bryophyte databases (Southwest Environmental Information Network, Consortium of North American Bryophyte Herbaria) were searched. We standardized the nomenclature but did not confirm the identity of reported specimens. The taxonomy of hepaticae currently is in a state of flux. The taxonomy used here is basically that of Doyle and Stotler, 2006, with the exception that the Lophoziaceae are listed as their own family, instead of being included with the Scapaniaceae. The Lophoziaceae are not complex bilobed liverworts, whereas the Scapaniaceae are. Using species descriptions in the literature, as well as specimens collected by us (most deposited in the Dale (Continued on page 2) Botanice` est Scientia Naturalis quae Vegetabilium cognitiorem tradit. — Linnaeus Page 2 (Continued from page 1) A. Zimmerman Herbarium, SNM, at Western New Mexico University), we developed a dichotomous key to the liverworts reported from New Mexico. A few species which have not been reported from New Mexico, but are expected, have been included. It is noted that this key is at best preliminary, and readers are encouraged to report questions or corrections to the authors. Pictures or photographs are not presented here. Photographs of the species we have collected are in Allred et al. (2018) and at our website www.gilaflora.com. RESULTS A preliminary checklist of the liverworts of New Mexico and the Gila National Forest was presented at the 4th Symposium on the Natural History of the Gila in 2012 (Blisard and Kleinman 2015). At that time, approximately 73 species of liverworts were known from New Mexico. Since then, reference to additional species have been found in the literature, and four species new to the state have been found. A few species have been removed from the list, because they are not currently recognized as distinct taxa (ie Lophozia confertifolia Schiff.). We were able to identify a total of 89 species in 23 families for which there is relatively good evidence that these taxa were collected in New Mexico. These are listed in the checklist (Table 1), along with documentation (literature references, or specimen identity of those taxa listed in digital databases). We have not confirmed the identity of specimens which were not collected by us. Since the publication of the previous checklist (Blisard and Kleinman 2015), we have collected three species new to the state. Geocalyx graveolens (Schrad.) Nees was collected at El Malpais National Monument. Riccia atromarginata Levier was collected at Carlsbad Caverns National Park, and also within the city of Carlsbad (Kleinman et al, 2020). It is interesting to note that both of these specimens were collected after a period of seasonal flooding, in areas which had been visited on several other occasions. It is characteristic of R. atromarginata that it is inconspicuous when dry, but more obvious when wet. Fossombronia pusilla (L.) Nees was finally found in a reproductive state, so could be identified to species from the spores. In addition, a fourth species new to the state (Frullania mexicana Lindenb.) was reported by Attwood (2017). Finally, dichotomous keys to the liverwort families (Table 2) and the liverwort species (Table 3) reported from New Mexico are presented. We have included selected species which are not known to occur in New Mexico, but might be expected here; they are marked with an asterisk (*). At the end are tables of characteristics which are helpful in the identification of the genera Frullania (Table 5), Lophozia (Table 6), Scapania (Table 7), and Riccia (Table 8). DISCUSSION This checklist shows considerable diversity in the liverwort population of New Mexico. This diversity is perhaps not surprising in view of the size of our state (121,697 square miles) and the rich diversity of habitats. Habitats include Chihuahuan desert scrubland, found in arid regions at lower elevations (860-1676 m, 2840-5500 ft). At moderate elevations (1676-2133 m, 5500-7000 ft), pinon-juniper-oak forests and mixed conifer forests predominate. At higher elevations, Ponderosa pine forest (2133-2743m, 7000-9000 ft) and spruce-fir forests (above 2743 m, 9000 ft) occur. Riparian habitats are located adjacent to water courses, and bogs are found in isolated areas. Liverwort diversity in New Mexico is higher than that reported from nearby states: we found 89 species in New Mexico by literature review and our own collections. In contrast, 46 species have been reported from Nevada (Brinda et al. 2007), and 60 from Utah (Flowers 1961). The 142 species reported from California (Doyle and Stotler 2006) may represent that state’s habitat diversity as well as a greater number of bryologists studying California liverworts. It seems likely that as more people study liverworts in New Mexico (and indeed, the entire southwest), more species will be identified. REFERENCES Allred,K., R. Kleinman, and K. Blisard, 2018. Preliminary Guide to the Bryophytes of New Mexico. Lulu.com. 120 pp. Arsene, Frére G., 1933. Hépatiques du Nouveau-Mexique (U.S.A.) déterminées par Miss Caroline Coventry Haynes. Ann. Crypt. Exot. 6:150- 160. Attwood, J.J., 2017. Notes on two species of Frullania subg. Chonathelia (Frullaniaceae) from the southwestern United States. Evansia 34:104- 109. Bird, C.D., 1960. Notes on the Current List of New Mexican Hepaticae. The Bryologist 63:114-115. Blisard, K.S. and R. Kleinman, 2015. A Checklist of the Liverworts of New Mexico and a Preliminary Assessment of the Liverworts of the Gila National Forest. The New Mexico Botanist, special issue #4, 21-27. Botany is the natural science that transmits the knowledge of plants. — Linnaeus (Continued on page 3) Page 3 (Continued from page 2) Brinda, J.C., L.R. Stark, J.R. Shevock, and J.R. Spence. 2007. Annotated Checklist of the Bryophytes of Nevada, with Notes on Collecting History in the State. The Bryologist 110:673-705. Consortium of North American Bryophyte Herbaria (NABH) website: http://www.bryophyteportal.org/portal/index.php Doyle, W.T. and R.E. Stotler, 2006. Contributions toward a Bryoflora of California III. Keys and Annotated Species Catalogue for Liverworts and Hornworts. Madrono, 53:89-197. Evans, A.W., 1922. Notes on North American Hepaticae – IX. The Bryologist 25:25-33. Flowers, S. 1961. The Hepaticae of Utah. University of Utah Biological Series, 12:1-108. Frye, T.C. and L. Clark, 1937-1947. Hepaticae of North America. 5 parts. Univ. Washington Press Flora of the Gila National Forest website: http://www.gilaflora.com Guerke, W.R. 1971. Notes on Hepaticae from Louisiana and New Mexico. The Bryologist 74:202-204. Heinrichs, J., H-P. Kreier, K. Feldberg, A.R. Schmidt, R-L. Zhu, B. Shaw, A.J. Shaw, and V. Wissemann, 2011. Formalizing Morphologically Cryptic Biological Entities: New Insights from DNA Taxonomy, Hybridization, and Biogeography in the Leafy Liverwort Porella platyphylla (Jungermanniopsida, Porellales) American Journal of Botany 98:1252-1262. Hentschel, J., M.J. von Konrat, T. Pocs, A. Schaefer-Verwimp, A.J. Shaw, H. Schneider, and J. Heinrichs. 2009. Molecular Insights into the Phylogeny and Subgeneric Classification of Frullania Raddi (Frullaniaceae, Porellales). Molecular Phylogenetics and Evolution 52:142 -156. Hicks, M.L. 1992. Guide to the Liverworts of North Carolina. Duke University Press. Hong, W.S. 1989. The Genus Frullania in North America West of the Hundredth Meridian. The Bryologist 92: 363-367. Hong, W.S. 1992. Plagiochila in Western North America.