CALIFORNIA STATE UNIVERSITY, NORTHRIDGE A PHOTOGRAPHIC KEY TO THE MOSS FAMILY ORTHOTRICHACEAE IN CALIFORNIA A thesis submitted in partial fulfillment of the requirements For the degree of Master of Science in Biology By Nickte M. Méndez December 2016 Copyright by Nickte M. Méndez 2016 ii The thesis of Nickte M. Méndez is approved: ____________________________________ ______________ Dr. Jeanne M. Robertson Date ____________________________________ ______________ Dr. Robert E. Espinoza Date ____________________________________ ______________ Dr. Paul S. Wilson, Chair Date California State University, Northridge iii Dedication For Amaya iv Acknowledgements First and foremost I want to thank my advisor Paul Wilson, for his encouragement, patience, and sometimes brutal but well-intentioned honesty. He is a great advisor who wants his students and advisees to be successful. Thank you to Dr. Jeanne Robertson and Dr. Robert Espinoza for serving on my committee and for your support throughout this process. I would also like to thank A. Heinrich, L. Coleman, S. Morley, S. Khimji, and N. Uelman for keying specimens through an earlier draft. Your suggestions are appreciated and useful in revising the key. I was very fortunate to get to speak to Dan Norris who provided me with insight into Orthotrichum at the very beginning of my work; thank you Dan. I would like to thank Kim Kersh at the Herbaria of the University of California, Berkeley, and Jim Shevock at the California Academy of Sciences for loans of specimens. I would also like to thank Ricardo Garilleti and Dale Vitt for their help with species identification. To Brent Mishler and Ken Kellman, thank you for your words of encouragement. Thank you Karol Marhold of The International Association for Plant Taxonomy and Scott Schuette of The American Bryological and Lichenological Society for permission to use images from their journals in my key. To my labmates Katherine Gould, Dani Amoroso, and Lena Ayala Coleman, thank you for the advice and support. I will miss our weekly lunches. To my parents, Clemente and Bertha Méndez, thanks for your love and support. Hector, thank you for letting me go first. Thanks for being my rock and holding me together when I was falling v apart. To our daughter Amaya, you were the catalyst that drove me to want to be better. Thanks for being so understanding at such a young age. One day I hope you learn to love plants, especially mosses, as much as I do. vi Table of Contents Copyright Page ii Signature Page iii Dedication iv Acknowledgements v List of a Tables viii Abstract ix Introduction 1 Methods 8 Identifying Orthotrichum reported and expected from California 13 Key 17 Discussion 76 References 84 Appendix: Useful Specimens Examined 91 vii List of a Table Table 1: Orthotrichaceae known and expected from California 3 viii Abstract A PHOTOGRAPHIC KEY TO THE MOSS FAMILY ORTHOTRICHACEAE IN CALIFORNIA By Nickte M. Méndez Master of Science in Biology Orthotrichum (Orthotrichaceae) is a species-rich genus of mosses in California. The last key to Californian Orthotrichaceae was produced in 2004 and included 35 species. Since that time, new discoveries have been made, and new identification tools are being posted on the site called the California Moss eFlora. As a model for the eFlora, I created an updated key to Orthotrichaceae in California with photographs at each couplet. I included 47 species known from California and adjacent states. My key will be useful in verifying the thousands of Orthotrichum specimens currently housed in herbaria. Species distribution maps produced after verification of those specimens could allow for tests of hypotheses regarding the biogeography of Orthotrichum in California. ix Introduction Orthotrichum is the most species-rich genus of mosses in California. Hedwig (1789) initiated the genus with three species: O. anomalum, O. crispum, and O. striatum. Up to the present, nearly 600 names have been proposed, and 166 have not been synonymized (www.tropicos.org search in 2016; Goffinet et al. 2007). Species within Orthotrichum grow on trees or rocks, occurring mainly below the elevation at which snow accumulates throughout the winter (Vitt 1973). Most species of Orthotrichum are monoicous. They are usually found with sporophytes, which are born at the ends of ascending stems (they are acrocarpous). Gametophytes are also able to reproduce through leaf fragmentation (Goffinet & Shaw 2008), and a few species produce specialized asexual propagules (gemmae). Generally, if gemmae are abundant, sporophytes are not present (Goffinet & Shaw 2008, Mishler 1988). The group is known by its distinctive calyptra, which completely covers the developing sporophyte and which is often hairy (Lewinsky 1993). It has been suggested that Orthotrichum has largely radiated in the Mediterranean-climate regions of the Mediterranean Basin, California, and to a lesser extent the Andes (Carter et al. 2016). Vitt’s (1973) A revision of the genus Orthotrichum in North America, north of Mexico thoroughly dealt with older literature and is thus the starting point for recognizing species new to science and for reporting range extensions. A number of recent regional treatments of Orthotrichum have improved taxonomic resolution of the species. Flora Briofítica Ibérica, Volume V contains detailed information on 35 species and five additional varieties along with line drawings illustrating most characters used in identification (Guerra et al. 2014). Nationalnyckeln till Sveriges Flora och Fauna, covering Scandinavia, presents a photograph key to 26 Orthotrichum species (Hedenäs et al. 2014). The Moss Flora of Mexico covers 15 species, most of which are illustrated (Sharp et al. 1994). Flore des Bryophytes du Québec-Labrador provides helpful illustrations and descriptions to 13 species (Faubert et al. 2014). Lastly, Flora of North America treats 49 terminal taxa (Vitt in Zander & Eckel 2014). Orthotrichum is in the Orthotrichaceae. The Orthotrichaceae as a whole is distributed worldwide and includes over 500 species, most of which are epiphytic or epilithic. The family is known for two very large genera: Orthotrichum is rich mainly in the northern hemisphere, whereas Macromitrium radiated in the southern hemisphere and the tropics with only one species extending into North America. Aside from the two large genera, the Orthotrichaceae includes up to 20 smaller genera. In California, in addition to Orthotrichum, we have three species of Ulota and one species of Zygodon (as well as three species of Amphidium, not treated here, sometimes placed in the Rhabdoweisiaceae). The most recent key to Orthotrichaceae in California was published in 2004 by Norris and Shevock. Since then, a number of new species have been described, range extensions have been reported, and taxonomic revisions have resulted in name changes (Table 1: Norris et al. 2004; Garilleti et al. 2006, 2011; Medina 2008, 2012; Medina et al. 2013; Vigalondo et al. 2016). Ten species have been newly reported in California: Orthotrichum acuminatum, O. anodon, O. columbicum, O. confusum, O. cucullatum, O. franciscanum, O. mazinpakanum, O. norrisii, O. persimile, and O. shawii. 2 Identification of all mosses in California, not just Orthotrichaceae, is now being presented online as The California Moss eFlora (http://ucjeps.berkeley.edu/ CA_moss_eflora/). Initially, the eFlora pulled together the keys of Norris and Shevock (2004), California Mosses (Malcolm et al. 2009), Norris’s unpublished descriptions, and a specimen database. The site is curated by bryologists from UC Berkeley and CSU Northridge, with contributions from other professional and amateur bryologists. For my thesis, I revised the key to the Orthotrichaceae for the California Moss eFlora. My key presents photographs at each couplet. It treats 47 species rather than the 35 of Norris and Shevock (2004). And, I have reworked the sequence and wording based on specimens that I examined and on treatments of Orthotrichum from elsewhere (Méndez 2015). My thesis ends with a discussion developing hypotheses on the origin of endemic species in the Orthotrichaceae in California. Table 1. Orthotrichaceae known and expected from California. Information largely extracted from Vitt (2014) with additions from publications on novelties and from examining the specimen database of the Consortium of North American Bryophyte Herbaria. Species and authors Range* and elevation Substrate Orthotrichum acuminatum ⱡ H. w Eu, e Africa, w N Am, (0-)150 On tree trunks Philib., 1881 Rev. Bryol. 8: 28- to 1850 (-2100) m 31 Orthotrichum affine Schrader nw N Am, Eu, As, n Af 0..3000 m Deciduous trees, shaded rocks ex Bridel 1801 Muscol.. Recent. In CA mainly CaR, MP, NW, SN (rare) 2(2): 22 Orthotrichum alpestre Greenland, nw N Am, Eu, As, Rock, base of trees, moist Hornschuch ex Bruch & New Zealand..2100..3000 m crevices of large boulders, moist Schimper 1849 Byol . Europ. 3: In CA mainly in CaR, SN, SNE pine and deciduous forests 3 75 Orthotrichum anodon ⱡ F. Lara, Endemic to Los Angeles Co. Quercus trunks R. Garilleti & Mazimpaka 2006 Bryologist 109: 188 Orthotrichum anomolum Greenland, N Am, Mexico, Eu, Rock, limestone bases, bases of Hedwig, S 1801 Frond., 162 Asia n Af 0..3000 m trees, dead branches, dry rock faces in canyons Orthotrichum bolanderi w N Am (Calif. & Wash.), Mex, c Xerophytic rock surfaces, Sullivant, Icon 1874 Musc., Am (Guatemala) 100..200 m especially sandstone suppl., 64 In CA mainly CW, NW, SN, SW Orthotrichum columbicum ⱡ R. w N Am, Eu Tree trunks, branches, and Medina, F. Lara, B. Goffinet, R. shrubs in humid coniferous Garilleti, Mazimpaka 2012 forests, boulders in shaded Taxon 61: 1180-1198 habitats Orthotrichum confusum ⱡ R. Endemic to n CA Tree trunks, branches, and Medina, F. Lara, B. Goffinet, R. shrubs in humid coniferous Garilleti, Mazimpaka 2012 forests, boulders in shaded Taxon 61: 1180-1198 habitats Orthotrichum consimile Mitten, B.C., w N Am, Eu Tree trunks, branches, and J. 1864 Linn. Soc., Bot. 8:24. In CA mainly CW, NW shrubs in humid coniferous forests, boulders in shaded habitats Orthotrichum coulteri R.