California State University, Northridge a Photographic

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

Nickte M. Méndez

December 2016

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

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

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

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.

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

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. Endemic to CA Tree trunks and branches, Medina, F. Lara, B. Goffinet, R. In CA mainly CR, SN occasionally rock in Garilleti & Mazimpaka 2013 sclerophyllous and deciduous Taxon 62: 1133-1152 forests

Orthotrichum cucullatum ⱡ R. Endemic to CA Tree trunks and branches, Medina, F. Lara, B. Goffinet, R. occasionally rock in Garilleti & Mazimpaka 2013 sclerophyllous and deciduous Taxon 62: 1133-1152 forests

Orthotrichum cupulatum B.C. w N Am, Mexico (Nuevo Calcareous cliffs and large Hoffman ex. Bridel 1801 León), Eu, As, n Af 100..3000 m boulders in coniferous forests, Muscol. Recent 2(2): 25 In CA mainly CaR, NW, SN, SNE on dry cliff faces in canyons

Orthotrichum diaphanum B.C. sw N Am, S Am, Eu, n As, Base and trunks of trees < 2 m, Schrader ex. Bridel 1801 Africa (Algeria), Atlantic Islands usually exposed areas, rock Muscol. Recent, 2(2): 29 100..1000 m (rare) In CA mainly CW, SW

Orthotrichum euryphyllum w N Am (Ca, Or, Wa) 0..500 m Calcareous springs in Artemisia Venturi, Bot 1890 Centralbl. In CA mainly CaR, MP scrub

44: 417

Orthotrichum flowersii Vitt w N Am, Eu 500..2000 m Base of trees, trunks of 1971 Bryologist 74: 159 In CA mainly DMoj, SN, SNE, SW deciduous trees, xeric and exposed habitats

Orthotrichum franciscanum ⱡ R. Endemic to Ca Tree trunks and branches, Medina, F. Lara, B. Goffinet, R. occasionally rock in Garilleti & Mazimpaka 2013 sclerophyllous and deciduous Taxon 62: 1133-1152 forests

Orthotrichum hallii Sullivant & B.C., w N Am 200..3000 m Limestone, calcareous Lesquereux in W.S. Sullivant, In CA mainly SN, SNE sandstone, rare on granite, Icon 1874 Musc., suppl., 63 quartzite, or basalt, deciduous tree trunks (rare), vertical canyon walls, shaded cliff faces

Orthotrichum holzingeri w N Am 1000..2000 m Seasonally wet rock in small Renault & Cardot, Contr 1895 In CA mainly SN streams of dry montane forests U.S. Natl. Herb. 3:270

Orthotrichum kellmanii D. H. Endemic to Santa Cruz Co. Dry sandstone boulders in dense Norris, Shevock & Goffinet chaparral 2004 Bryologist 107: 210

Orthotrichum laevigatum J, E. nw N Am, Eu, e As (Japan) Non-calcareous boulders, base Zetterstedt, Öfvers 1862 Kongl. 500..3000 m of deciduous trees Vetensk.-Akad. Förh 19: 363 In CA mainly CaR, MP, NW, SN, SNE

Orthotrichum lyellii Hooker & nw N Am, Mexico, Eu, n Af Trees, covering trunks up to 10 Taylor 1818 Muscol. Brit., 76 10..1000 m m, boulders In CA mainly CaR, CW, DMoj, NW, SN

Orthotrichum macounii Aust., Bull. Torrey Bot. Club 6: 343. 1879

Orthotrichum mazimpakanum ⱡ Endemic to s CA (Riverside Co, Tree bases (Quercus R. Garilleti, J. R. Shevock, D. H. Santa Barbara Co., & Humboldt chrysolepis), in Quercus agrifolia Norris & F. Lara 2011 Bryologist Co. woodlands 114: 346-355

Orthotrichum norrisii ⱡ F. Lara, Endemic to Orange Co. Tree trunks and branches, R. Medina & Garilleti 2008 occasionally rock in Bryologist 111: 670 sclerophyllous and deciduous forests

Orthotrichum obtusifolium B. C., n N Am, Eu, As 10..2000 m Trunks and bases of deciduous

Bridel 1801 Muscol. Recent. In CA mainly CW, MP, NW, SN trees ; Populus, Juniperus, or 2(2): 23 aspen

Orthotrichum pallens Bruch ex n N Am, Mexico, Eu 300..3000 m Trunks and branches of trees, Bridel 1827 Bryol. Univ. 1: 788 In CA mainly SNE especially calcareous or granitic rock

Orthotrichum papillosum In CA mainly CaR, CW, NW, SN On moist, diffusely lit tree trunks and branches

Orthotrichum pellucidum w N Am, Greenland, Eu Calcareous or siliceous boulders, Lindberg, Öfvers 1867 Kongl. In CA mainly SN cliff faces Vetensk.-Akad. Förh. 23: 549

Orthotrichum persimile ⱡ R. Endemic to n Ca (coast) 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 pilosissimum § R. Endemic to Nevada (Nye Co., Rock and tree roots and bases Medina, F. Lara, Mazimpaka, J. Mineral Co., Esmeralda Co., R. Shevock & R. Garilleti 2011 Eureka Co.) 1530..2300 m Bryologist 114: 316-324

Orthotrichum praemorsum w N Am Xerophytic rock surfaces (rock Venturi 1890 Bot. Centralbl. 44: In CA mainly SNE lava) and tree trunks 418

Orthotrichum pulchellum B. C., w N Am 0..1000 m Tree trunks, twigs and branches Brunton in J. E. Smith et al. In CA mainly NW of deciduous forests and 1807 Engl. Bot., 25 coniferous trees

Orthotrichum pumilum Swartz U.S. and Canada 10..1000 m Trunks, lower branches, base of 1801 Monthly Rev. 34: 538 In CA mainly NW, SN, SW deciduous trees, rarely in rock crevices

Orthotrichum pylaisii Bridel nw N Am, Eu, n As 100..3000 m On rock near oceans or alpine or 1826 Bryol. Univ. 1: 722 In CA mainly CaR, NW, SN arctic zones, on Salix or Alnus, bone, in or near bird rookeries, granitic boulders in coniferous forests

Orthotrichum rivulare Turner B.C., nw N Am, Eu 0..1000 m Exposed tree roots, base of 1804 Muscol. Hibern. Spic., 96 In CA mainly CaR, CW, NW, SN trees, siliceous boulders

Orthotrichum rupestre Greenland, B. C., w N Am, S Am, Non-calcareous boulders and Schleicher ex Schwägrichen Eu, e As, n, e Af, Atl Islands, Pac cliff faces, base of trees 1816 Sp. Musc. Frond. Suppl. Islands, Aus 100..3000 m 1(2): 27 In CA mainly CaR, CW, MP, NW, SN, SNE

Orthotrichum shawii ⱡ Wilson in w N Am (Ca), Eu Tree trunks W. P. Schimper 1864 Musci Eur. In CA mainly Nov. 1-2(1): 1

Orthotrichum shevockii Endemic to s CA (Kern Co, Mono Dry granite rocks Lewinsky-Haapasaari & D. H. Co., Alpine Co.) 1100..1600 m Norris 1998 Bryologist 101: 435 In CA mainly SN, SNE

Orthotrichum speciosum Nees Greenland, nw N Am, Eu, n As, n Coniferous and deciduous trees in J. Sturm et al. 1819 Deutschl. Af 10..2000 m and rocks Fl. 2(17): 5 In CA mainly NW, SN.

Orthotrichum spjutii D. H. Endemic to w N Am (Ca & Nv) Deciduous trees, moist, diffusely Norris & Vitt 1993 nova lit rocks Hedwigia 56: 260

Orthotrichum striatum Hedwig B. C., w N Am, Eu, As, n Af Deciduous trees, trunks of Acer 1801 Sp. Musc. Frond., 163 10..1000 m and Alnus In CA mainly NW, SN

Orthotrichum texanum In CA mainly CaR, NW Non-calcareous boulders and cliff faces, base of trees

Orthotrichum underwodii F. Endemic to CA (SF Co. & Trinity Trees Lara, Garilleti & Mazimpaka Co.) 2001 Bryologist 104: 268 In CA mainly CW

Ulota megalospora Venturi nw N Am Tree branches and trunks 1890 Bot. Centralbl 44:389 In CA mainly NW

Ulota obtusiuscula Müller Hal. w N Am Trees, epiphytic in temperate & Kindberg g 1892 in J. Macoun In CA mainly NW rainforests and N. C. Kindberg, Cat. Canad. Pl., Musci, 82

Ulota phyllantha Bridel 1818 w N Am, w & e Can, S Am, Eu, Af Tree branches and trunks along Muscol. Recent., suppl. 4: 113 In CA mainly CW, NW coast, associated with salt spray

Zygodon rupestris W. P. w N Am, Eu, Jap On trunks of trees, rock Schimp. Ex Lor. 1865 In CA mainly CW, NW, SN * based on herbarium records that have not been updated to current species concepts. ⱡ Species discovered since Norris and Shevock (2004). § not yet recorded from California but found in an adjacent state.

Methods

I started with Norris and Shevock’s (2004) key to the Orthotrichaceae. Much of their wording remains. Couplets were added, reordered, and revised to intercalate species recently found in California and to improve the structure of the logic. Most importantly, I added photos of key characters adjacent to each couplet.

I studied the characters that make each species distinct. I dissected 235 specimens

(see Appendix). Most of the specimens were from the herbaria of University of California

Berkeley (UC), California State University Northridge (SFV), and California Academy of

Sciences (CAS). When possible I used specimens verified by the contemporary experts

Dale Vitt, Francisco Lara, or Ricardo Garilleti. Specimens were keyed through other treatments (e.g., Guerra et al. 2014; Hedenäs et al. 2014; Vitt 2014). I was unable to identify two species that had not been previously recorded from California. Dale Vitt and

Ricardo Gariletti were consulted and identified these as Orthotrichum shawii (Garilleti et al. 2006) and O. acuminatum (Vigalondo et al. 2016). I annotated all the specimens I studied. Unless they had gemmae, sterile specimens were not annotated, as they could not be identified following the methods I used.

Microscopic photos were captured through an Olympus CX41 compound microscope (40×, 100×, 400×; Center Valley, PA) and an Olympus SZX9 dissecting microscope (6.3–57×; Center Valley, PA) using Infinity Capture v. 6.5.2 software

(Lumenera Ottawa, Ontario). Macro photos were captured with a Nikon (Melville, NY) digital single-lens reflex camera through a 105 mm lens. For species that I was unable to

photograph, I obtained permission to reuse photos from the journals The Bryologist and

Taxon.

I illustrated one (but not all) of the distinctions for each couplet. My photos are meant to discriminate between characters used in two leads that divide species into small groups rather than to define characters as applied to all mosses. I added labels, arrows and circles to explicate the terminology of the couplet. My images are of specimens that were not cleared or stained. I only cropped and adjusted lighting. My images have the virtue of appearing similar to what the casual microscopist will see.

My key and the California Moss eFlora takes advantage of links. It was written in

HTML (edited in Sublime Text v. 2.0.2, Sydney, New South Wales). The leads in the key are interactive allowing users to click to go to the next couplet or to a species page in the eFlora, which in turn is linked to a great deal of other information (Legget & Kirchoff

2011). Another advantage of producing a web-based key is the ease with which the key can be updated as new discoveries are made.

A taxonomist may present a synoptic key or an artificial key. A synoptic key emphasizes characters that are relevant to classification, whereas an artificial key relies on characters that are easy to distinguish but are not necessarily relevant to classification

(Bell 1967). Synoptic keys highlight “natural” taxonomic relationships (Scharf 2009), which can impede identification. First, the synoptic key uses characters that may be difficult to observe. Second, species that are similar because of convergence are typically far from one another in the key. The keys in the California Moss eFlora are artificial.

Keys may have one character per couplet or provide the user all the differences known about a separation (Sokal 1985). The former is simple and makes the key a tidy outline. However, the latter enables identification of intermediate, ambiguous, or incomplete specimens. When possible, I noted differences in multiple characters, but I only provided images for one character per couplet. Within a couplet, I placed first the character that I considered to be the most readily observed, most categorically different, or unique to a species or larger group. For example:

A. Axillary hairs hyaline, distinctively long and protruding from the foliose shoots; capsule strongly constricted when dry; leaves unistratose or distal portion rarely with scattered bistratose ridges ..... Orthotrichum pilosissimum

A' Axillary hairs, not long and not protruding from foliose shoots; capsule slightly or not constricted when dry; leaves unistratose ...... Orthotrichum norrisii

The order in which the characters are listed reflects their priority at facilitating identification.

I also made an effort to arrange the couplets so those occurring early in the key use only characters that do not vary seasonally (e.g., gametophytic characters such as leaf morphology). The first lead in the key divides species according to whether leaves are curled when dry or straight. As a last resort and generally near terminal couplets, I used characters of the peristome, which is present only seasonally.

Characters that are easily observed were used in early couplets even if they did not split the sets into equally numerous species. For instance, the red exostome of

Orthotrichum pulchellum easily separates it from O. columbicum, O. persimile, O. consimile, and O. confusum. Then, the 16 endostome segments of Orthotrichum columbicum separate it from the rest. Orthotrichum persimile is separated based on leaf

morphology, leaving O. consimile and O. confusum to be separated based on the hairs of the calyptra. It was easier to segregate off one species at a time than to start by separating the group into two equally sized subgroups.

The sequencing of couplets is often affected by intermediate species. I eliminated intermediate species in an early couplet based on characters that set it apart from the rest.

A. Capsule exserted, mostly with 8 long ribs alternating with 8 short ones ..... Orthotrichum anomalum A' Capsule emergent, variously with 8 or 16 ribs ..... B

B. Capsule with 16 ribs ..... Orthotrichum cupulatum B' Capsule with 8 ribs ..... Orthotrichum pellucidum

The in-between state of O. anomalum made me place the ribbing distinction in the latter couplet, after elimination of O. anomalum; reversing the order of the couplets would have not worked as well.

Sometimes it is helpful to write a bypass route into a key. A bypass allows the user to identify species in a distinctive phenological state. I did this for specimens with gemmae. Specimens with gemmae are keyed through a different section, and specimens with sporophytes are keyed through the main part of the key. Specimens with gemmae require the user to make fewer decisions than using sporophytic characters. Like my key, the Swedish flora includes a gemma bypass in their Orthotrichum key (Hedenäs et al.

2014).

Another question that arises is the decision to make a bracketed key as opposed to an indented key. The California Moss eFlora allows users to toggle between bracketed and indented formats in KeyBase. Indentation is good when the whole key fits on one screen. It allows users to see the structure of the key (Porter 1959). The indented display

might well be best for a synoptic key to a small number of genera in a family. But when an artificial key has more species than can fit on a screen, indented keys become worse than bracketed keys. Indented keys have some drawbacks. First, it is difficult to compare two leads in a couplet that are far apart (Porter 1959). Next, it may be difficult to locate the second lead or return to the first lead (Davis & Heywood 1963; Stace 1989). Because my key was long, I used a bracketed format in which the leads of a couplet are directly juxtaposed (Porter 1959). The format of a bracketed key is improved with the addition of images, and these images were placed side-by-side directly under the couplet (Legget &

Kirchoff 2011).

Orthotrichaceae is a difficult group in terms of identification, and more generally the terminology describing moss morphology is unfamiliar to many people. It is impossible to avoid bryological terminology when identifying species in Orthotrichum.

Closely related species may only differ in their axillary hairs and subtle dimensions. I assume that the user of my key has some knowledge of the terminology. Individuals who are just starting will find a use for the bryological glossary by Malcolm & Malcolm

(2006).

Identifying Orthotrichum reported and expected from California

Examine characters in the following order.

1. Leaves. Dry leaves are either contorted or ± erect (for photos, see Couplet A). Only a

few species of Orthotrichum have contorted leaves, so much of the family can be

eliminated based on this one obvious character.

2. Height of capsule relative to leaves. Determine the degree to which the capsule is

immersed, emergent, or exserted (Couplet X). An immersed capsule has leaves

extending beyond the mouth. An emergent capsule has leaves covering the lower part

of the capsule. An exserted capsule is on a longer seta that is surrounded by leaves

whose tips do not extend to the base of the capsule.

3. Capsule ribbing. Capsules, when dry, vary from smooth (unribbed), to weakly ribbed,

to strongly ribbed (Couplet AF). A weakly ribbed capsule is ribbed longitudinally

from just below the mouth to about the middle of the urn. A strongly ribbed capsule is

ribbed to near the base of the urn. The number of ribs on the capsule also varies: eight

ribs, 16 ribs, or eight strong ribs alternating with eight weak ribs. Species vary in the

degree to which capsules are constricted below the mouth when dry, and in general,

the more constricted the capsule, the stronger the ribbing.

4. Peristome. In many Orthotrichaceae the peristome is double. If so, the capsule mouth

will have exostome teeth and endostome segments (visible through a good dissecting

microscope; Couplet AX). The number of exostome teeth can be zero, eight, or 16.

Species with eight teeth will have teeth that are four cells wide. Species with 16 teeth

will have teeth that are two cells wide. In intermediate cases, eight teeth may be

partially split longitudinally. The number of endostome segments can also be zero,

eight, or 16. Endostome segments are usually a single cell wide in Orthotrichum.

Next, dissect the specimen. Take a shoot, and moisten it on a microscope slide.

Pull numerous leaves off of the stem from their bases. Make cross-sections of multiple leaves, one at a time, at two-thirds distally. Select a sporophyte that has an intact peristome. If only unopened capsules are available, pluck the most mature one, make a wet mount, and try to boil off the operculum by holding the slide over a flame. If only old capsules are available, try to find one with some peristome parts. Regardless, slice the capsule longitudinally, and splay out the halves with the exterior up. Add a coverslip, and if necessary, boil away air bubbles.

5. Gemmae. Gemmae in Orthotrichum, when present, grow on the leaf surface. They are

often rust colored. At maturity, they abscise and function as asexual propagules.

Orthotrichum lyellii has very prominent gemmae, which can be observed with the

naked eye. Gemmae in other species may be much smaller, and observable only with a

microscope (100×; Couplet J).

6. Microscopic leaf characters. Many useful characters pertain to the shape of leaves,

their apex, margin, and thickness. The apex of the leaf can end acutely without

differentiated cells, with a mucron (short accentuated tip), or with a hyaline awn (a

pigmentless extension of the costa beyond the lamina). The number of cell layers is

best examined in cross-sections in the distal two-thirds of the leaf, as this is where

species differ. Species vary from being unistratose, to having bistratose streaks, to

being fully bistratose beyond approximately the midleaf. In some species with a

unistratose lamina, the leaf margins may be bistratose, or even tristratose (Couplet Q).

Leaf margins are usually recurved, but a few species distinctively have plane margins.

Lastly, leaf papillae can be single or forked, and their location (over the lumens or

over the cell walls) can vary, and where they are prominent on the leaf (on the distal

third) can be diagnostic.

7. Stomata. Stomata are observed on a longitudinal section of the capsule. Stomata are

either phaneroporous or cryptoporous (Couplet S). When stomata are phaneroporous,

exothecial cells abut guard cells in the same plane. When stomata are cryptoporous,

guard cells are recessed and may be partly covered or almost completely covered by

specialized subsidiary cells.

8. Ornamentation of exostome teeth. The outer surface of exostome teeth may be

nearly smooth, or have coarse papillae, or have fine papillae, and sometimes papillae

swirl into finger-print-like striations (Couplet AN).

9. Hairs of calyptra and of vaginula. Calyptrae may be hairless or hairy, with hairs

straight or wavy. Distinctive hairs may also occur on the vaginula, which is the collar

that surrounds the base of the developing sporophyte. Microscopically (400×), hairs

may be smooth or papillose (Couplet I).

10. Axillary hairs. Axillary hairs are organs adaxial to a leaf where it joins a stem

(Couplet AU). They are best seen (400×) when still attached after tearing most of the

leaf off of its shoot. Axillary hairs are unusually long in a few species of

Orthotrichum, whereas most species of Orthotrichum have axillary hairs that are about

the same length as in other mosses.

Key

A. Leaves weakly crispate to contorted when dry ..... B

A' Leaves ± erect when dry ..... J

Orthotrichum pulchellum crispate leaves (Norris Orthotrichum flowersii erect leaves (Coleman 1350) 74989, UC 1673879)

B. Apex of leaf usually filiform, uniseriate, 2–5 cells long ..... C

B' Apex of leaf not uniseriate ..... D

Zygodon rupestris filiform apex (Wilson 5050, SFV Orthotrichum norrisii leaf apex blunt (Shevock 21569) 32529, UC 1933675)

C. Plant stoloniferous with prostrate axes possessing abundant red-brown rhizoids; gemmae absent; erect branches short, not hamate when dry; leaf ovate with a long acumen, mid-laminal cells pluripapillose with papillae more than half the thickness of cells in cross-section ..... Ulota megalospora

C' Plant erect and tufted; abundant gemmae in leaf axils; stem apices of dry plant strongly hamate; leaf lanceolate, mid-laminal cells pluripapillose with 2–4 papillae per cell, papillae much less than half the thickness of cells in cross-section ..... Zygodon rupestris

Ulota megalospora prostrate axes with abundant Zygodon rupestris erect and tufted, lacking red- red-brown rhizoids (Wilson 206, SFV 20008) brown rhizoids (Wilson 5050, SFV21569)

D. Margins serrulate; apical cell longer than cells of lamina; basal marginal cells quadrate to very short rectangular, abruptly differentiated into several rows from more elongate interior basal cells; capsules exserted with a long and well-demarcated neck; stomates phaneroporous ..... E

D' Margins entire; apical cell same length-to-width ratio as cells of distal lamina; basal marginal cells not so abruptly differentiated; capsules and stomates various ..... F

Ulota obtusicula serrulate leaf margins (Norris Orthotrichum pulchellum leaf apex (Norris 75064, 78463, SFV 20907) UC 1650214)

E. Apex of leaves toward shoot tip with a reddish-brown cluster of septate gemmae; occurring within ~1 km of ocean ..... Ulota phyllantha

E' Apex of leaves not gemmiparous; moss more widely distributed ..... Ulota obtusicula

Ulota phyllantha leaf apices with gemmae cluster Ulota obtusicula leaf apices lacking gemmae (Wilson 537, SFV 20009) (Norris 78463, SFV 20907)

F. Exostome ± red; exothecial bands mostly 2 cells wide ..... Orthotrichum pulchellum

F' Exostome not red; exothecial bands mostly 4–6 cells wide ..... G

Orthotrichum pulchellum highly recurved ± red Orthotrichum consimile exostome teeth not red exostome teeth (Norris 75064, UC 1650214) (Norris 50401, UC 1722823)

G. Endostome in 16 segments (sometimes 8 of them reduced alternating with 8 that are more prominent), inner peristome layer with conspicuous longitudinal striae; operculum with a red basal rim; seta commonly 2.5–5.5 mm long; calyptra naked ..... Orthotrichum columbicum

G' Endostome in 8 segments, inner peristome layer smooth or weakly papillose; operculum with a thin orange basal rim; seta commonly 1.0–2.0 mm; calyptra naked or with ± abundant hairs ..... H

Orthotrichum columbicum 16 endostome segments Orthotrichum consimile 8 endostome segments (Shevock 41858, CAS 1187203) (Dillingham 2006, UC 1932571)

H. Leaf margins plane (sometimes one leaf margin recurved); leaf apex various, not filiform; rhizoids rough, not noticeably ascending along stem; vaginula hairy; calyptra ± hairy; stomata in proximal half of urn and neck ..... Orthotrichum persimile

H' Leaf margins recurved; leaf apex commonly filiform; rhizoids smooth, ascending along stems; vaginula naked or very sparsely hairy; calyptra naked or hairy; stomata usually in middle of urn (rarely in proximal half, never on neck) ..... I

Orthotrichum persimile plane leaf margins (Medina Orthotrichum consimile recurved leaf margins et al. 2012. Taxon 61: 1180–1198) (Medina et al. 2012. Taxon 61: 1180–1198)

I. Calyptra with abundant to scarce papillose hairs; vaginula (0.25)0.4–0.8 mm long, naked or with sparse papillose hairs; leaf apex merely following the curve of the leaf .....

Orthotrichum consimile

I' Calyptra naked, rarely with very few inconspicuous smooth hairs; vaginula very long,

(0.6)0.8–1.1(1.3) mm, naked; leaf apex distinctly unguiculate ..... Orthotrichum confusum

Orthotrichum consimile papillose calyptra hairs Orthotrichum confusum smooth calyptra hairs (Dillingham 2006, UC 1932571) (Medina et al. 2012. Taxon 61: 1180–1198)

J. Gemmae present (appearing as brown multicellular warts on leaves) ..... AZ—bypass for mosses with gemmae, does not rely on sporophytes

J' Gemmae absent ..... K—species with gemmae can be keyed via this lead, though with greater difficulty

Orthotrichum obtusifolium gemmae present on leaf Orthotrichum cupulatum, gemmae absent from leaf (Norris 68813, UC 1643011) (Norris 87675, UC 1759893)

K. Cells above the base of the leaf and inward from the margin with high, pointed papillae inserted not on lumen but on lateral walls; dioicous; leaf apices either very long and somewhat flexuose when dry or with apex bluntly eroded by gemma production .....

K' Papillar arrangement not as above (proximal cells smooth or papillose over the lumen); monoicous; leaf apices usually not long and flexuose; gemma production various

..... M

Orthotrichum papillosum high pointed papillae on Orthotrichum rupestre papillae on lumens of walls of proximal cells (Norris 83851, UC 1774498) proximal cells (Coleman 1651)

L. Gametophytes usually densely covered with brown wart-like multicellular gemmae; leaves not long acuminate ..... 2Orthotrichum lyellii sensu stricto

L' Gametophytes without gemmae; leaves usually having a long, slightly serrulate acumen ..... Orthotrichum papillosum

Orthotrichum lyellii with gemmae covering leaves Orthotrichum papillosum without gemmae (Wilson (Coleman 2) 5358)

M. Distal leaf lamina bistratose or at least with some bistratose streaks ..... N

M' Leaf lamina unistratose ..... R

Orthotrichum bolanderi distal portion of leaf lamina Orthotrichum rupestre unistratose leaf lamina bistratose (Norris 81945, UC 1774444) (Coleman 1651)

N. Stomata phaeneroporous; gametophytes in loose wefts with shoots often greater than 1 cm long ..... Orthotrichum bolanderi

N' Stomata cryptoporous; ascending shoots less than 1 cm long ..... O

Orthotrichum bolanderi gametophytes in loose Orthotrichum cupulatum gametophytes in short wefts (© John Game) tufts (© Scott Loring) O. Gametophytes consisting of prostrate stems giving rise to differentiated vegetative and fertile uprights (unlike other Orthotrichum species); vegetative leaves nearly obtuse; on sandstone .....2Orthotrichum kellmanii

O' All stems of the same sort, in tufts (like other Orthotrichum species); on limestone or granite ..... P

Orthotrichum kellmanii fertile leaves (left) and Orthotrichum hallii undifferentiated stems and sterile vegetative leaves (right) (Shevock 29890, UC leaves (Norris 18242) 1872681)

P. Mature capsules with 16 ribs; distal part of leaf mainly unistratose with bistratose streaks (margins not thicker) ..... 2Orthotrichum cupulatum

P' Mature capsule with 8 prominent ribs or with 8 faint ribs between 8 prominent ribs; distal part of leaf uniformly bistratose its full width or margins bi- to tristratose ..... Q

Orthotrichum cupulatum caspules with 16 ribs and Orthotrichum shevockii immersed capsule, strongly 16 erect exostome teeth (Norris 84832, UC 8 ribbed (Norris 87128, UC 1759935) 1759913)

Q. Leaf margins not thicker than leaf lamina; capsule immersed to emergent; exostome teeth 8 or irregularly split into 16, erect or spreading; leaf papillae small and conical; gametophyte ± green in clumps of short shoots ..... Orthotrichum hallii

Q' Leaf margin thicker (more cells) than leaf lamina, sometimes lamina also with bistratose streaks; capsule immersed; exostome 8; leaf papillae high, simple to forked .....

2Orthotrichum shevockii

Orthotrichum hallii nearly uniformely bistratose Orthotrichum shevockii leaf margins bi- to (Norris 80330, UC 1757602) tristratose (Norris 87128, UC 1769935)

R. Margins incurved to plane throughout; gametophyte green, often with a ± red-brown cast; lamina unistratose; epiphytic, in mountains, often near running water .....

2Orthotrichum obtusifolium

R' Margins recurved to revolute at least at base and usually along most of leaf; plant color various; lamina unistratose or not; habitat various ..... S

Orthotrichum obtusifolium plane leaf margins Orthotrichum pylaisii recurved leaf margins (Norris 68813,UC 1643011) (Coleman 1010)

S. Stomata phaneroporous as seen in longitudinal section of capsule; usually basal cells elongate, somewhat porose or with sinuose walls ..... T

S' Stomata cryptoporous; usually basal cells short-rectangular with evenly thickened walls ..... AH

Orthotrichum holzingeri phaneroporous stomata Orthotrichum norrisii cryptoporous stomata (Norris 70467, UC 1649719) (Shevock 32529, UC 1933675)

T. Exostome reduced to few basal plates or to narrow endostome segments ..... U

T' Exostome fully developed ..... V

Orthotrichum spjutii peristome reduced (Shevock Orthotrichum laevigatum fully developed peristome 14898, UC 1754439) (Wilson 4486, SFV 21283)

U. Gametophyte glaucous (due to very high multifid papillae); endostome absent; adaxial face of costa covered by laminal cells in at least distal third of leaf; spores to 11 µm in diameter ..... Orthotrichum spjutii

U' Gametophyte not glaucous; endostome in 8 segments; adaxial face of costa with elongate cells throughout; spores >11 µm ..... Orthotrichum acuminatum

Orthotrichum spjutii leaf cross-section, single and Orthotrichum acuminatum leaf cross-section, low multifid papillae (Shevock 14898, UC 1754439) papillae (Shevock 32571, CAS 1146075)

V. Leaves smooth distally ..... W

V' Leaves papillose distally ..... X

Orthotrichum holzingeri smooth leaves (Norris Orthotrichum laevigatum papillose leaves (Wilson 70467, UC 1649719) 4486, SFV 21283)

W. Calyptra hairless; peristome smooth; on volcanic rock ..... Orthotrichum holzingeri— endemic of volcanic plateaus of northeastern California and neighboring states to the northeast.

W' Calyptra hairy; peristome papillose; usually epiphytic ..... 2Orthotrichum shawii

Orthotrichum holzingeri smooth exostome teeth Orthotrichum shawii papillose exostome teeth (Norris 70467, UC 1649719) (Shevock 24632, CAS 1113091)

X. Capsule fully exserted with base of capsule beyond parichaetial leaves (side branch may extend in parallel) ..... Y

X' Capsule emergent to immersed ..... AB

Orthotrichum pylaisii exserted capsules (Coleman Orthotrichum rupestre weakly emergent capsule 1010) (Coleman 1651)

Y. Capsule weakly 8-ribbed, short-oblong gradually expanding from seta (less than 3:1 although difficult to judge); exostome teeth recurved to reflexed with age when dry; leaves broad distally (not acuminate, at two-thirds their length greater than half of their width) ..... Z

Y' Capsule weakly ribbed to not ribbed, cylindric (>3:1); exostome teeth typically erect to spreading with age when dry; leaves lanceolate (at least some somewhat acuminate)

..... AA

Orthotrichum pylaisii weakly ribbed capsule, Orthotrichum laevigatum capsules ± smooth exostome recurved when dry with age (Coleman (Wilson 4486, SFV 21283) 1010)

Z. Endostome segments one cell wide, smooth or finely papillose; leaf ovate-lanceolate to lanceolate ..... Orthotrichum pylaisii

Z' Endostome segments with remnants of a second cell width, coarsely and strongly papillose; leaf lanceolate to narrowly lanceolate ..... 2Orthotrichum speciosum

Orthotrichum pylaisii endostome segment one cell Orthotrichum speciosum endostome with remnants wide (Coleman 1010) of a second cell width (Norris 68877, UC 1723786)

AA. Leaves bluntly acute to obtuse; mid-laminal cells mostly rather low papillose; adaxial surface of costa with elongate to linear, thick-walled cells above its middle .....

Orthotrichum laevigatum—compared to Orthotrichum pylaisii, the gametophyte is smaller.

AA' Leaves long-acuminate to narrowly acute; mid-laminal cells with papillar salients at least as high as one-half of cell thickness; adaxial surface of costa covered with cells like those of adjacent lamina above its middle ..... Orthotrichum macounii—not recognized as distinct from Orthotrichum laevigatum by Vitt (2014).

Orthotrichum laevigatum adaxial surface; costa Orthotrichum macounii adaxial surface; costa with with elongate, thick– walled cells (Wilson 4486, cells like those of adjacent lamina (MacFadden SFV 21283) 19054, CAS 441460)

AB. Exostome generally separated into 16 teeth; capsules immersed to emergent ..... AC

AB' Exostome generally of 8 teeth with pairs not separated; capsules emergent ..... AF

Orthotrichum shawii exostome separated into 16 Orthotrichum affine exostome of 8 teeth (Norris teeth (Shevock 24632, CAS 1113091) 68516, SFV 20954)

AC. Capsule smooth; exostome papillose ..... Orthotrichum striatum—exostome 16, endostome 16.

AC' Capsule ribbed at least in distal half; exostome papillose to nearly smooth ..... AD

Orthotrichum striatum smooth capsule; exostome Orthotrichum rupestre nearly smooth exostome 16, endostome 16 (© Helge G. Gundersen) (Coleman 1651)

AD. Capsule emergent, weakly ribbed with metallic glints; exostome teeth tending to recurve with age; usually epiphytic ..... 2Orthotrichum shawii—exostome 16, endostome

8 short segments.

AD' Capsule immersed, ribbed entire length of capsule, without metallic glints; exostome teeth erect; usually epilithic ..... AE

Orthotrichum shawii emergent capsule with Orthotrichum rupestre immersed capsule without metallic-copper sheen (Shevock 24632, CAS copper sheen (Coleman 1651) 1113091)

AE. Gametophytes growing as relatively large, loose mats; exostome low papillose to almost smooth ..... Orthotrichum rupestre—exostome 16, endostome, if present, 8.

AE' Gametophytes growing in small, closely aggregated tufts; exostome densely papillose ..... Orthotrichum texanum—not recognized as distinct from Orthotrichum rupestre by Vitt (2014); exostome 16, endostome, if present, 8.

Orthotrichum rupestre nearly smooth exostome Orthotrichum texanum densely papillose exostome (Wilson 5359) (Hermann 20411, CAS 469527)

AF. Capsule ribbed only in distal half ..... 2Orthotrichum speciosum—endostome 8, exostome 8.

AF' Capsule ribbed more than half its length ..... AG

Orthotrichum speciosum capsule weakly ribbed Orthotrichum franciscanum capsule ribbed more (Norris 68877, UC 1723786) than half its length (Shevock 20202, CAS 1193149)

AG. Capsule long-cylindric to cylindric (widest proximally when old); exostome 8, endostome segments often 8, rarely 16; leaf apices acute, not generally bleached; generally living on bark .....Orthotrichum affine

AG' Capsule ovate-oblong to oblong-cylindric (widest distally when old); exostome teeth

8, endostome segments 16; leaf apices acuminate, cuspidate, often bleached .....

Orthotrichum praemorsum—compared to O. pylaisii with its emergent to exserted capsules, O. praemorsum has less emergent capsules that are more strongly ribbed.

Orthotrichum affine cylindric capsule, 8 endostome Orthotrichum praemorsum oblong capsule, 16 segments (Norris 68516, SFV 20954) endostome segments (Shevock 22039, CAS 1040112)

AH. Leaves terminating in hyaline awn; terminal cell of leaf elongate (more than three times as long as mid-laminal cells) ..... AI

AH' Leaves without hyaline awn ..... AJ

Orthotrichum diaphanum leaf with hyaline awn Orthotrichum euryphyllum leaf lacking hyaline awn (Shevock 19240, UC 1739285) (Shevock 32504, UC 1933798)

AI. Leaves terminating in a hyaline awn of one or a few cells; exostome teeth 8; endostome segments 8; gemmae absent ..... Orthotrichum franciscanum

AI' Leaves terminating in an awn of many hyalline cells; exostome teeth 16; endostome segments 16; gemmae often present ..... 2Orthotrichum diaphanum

Orthotrichum franciscanum hyaline awn of one cell Orthotrichum diphanum many hyaline cells (Shevock 20202, CAS 1193149) (Cardenas 5637, UC 1779402)

AJ. Exostome absent (endostome present); capsules puckered at mouth by 8 short ribs; calyptra hairy ..... AK

AJ' Exostome present; capsules various (many species not particularly constricted at mouth); calyptra hairy or hairless ..... AL

Orthotrichum mazimpakanum exostome absent, Orthotrichum underwoodii exostome teeth present endostome present with mouth puckered by 8 short (Toren 9511, CAS 1048109) ribs (Garilleti et al. 2011. The Bryologist 114: 346- 355)

AK. Endostome segments 16, very apparent when dry, ± white, strongly papillose, widely covering mouth; capsule mouth star-shaped when dry; vaginula naked or with scattered thin and smooth hairs; operculum convex, mucronate, without differentiated basal rim; leaf margin and lamina unistratose ..... Orthotrichum anodon

AK' Endostome segments usually 8, inconspicuous when dry, hyaline, smooth, sparse around mouth; capsule mouth round when dry; vaginula hairy, with thick papillose hairs; operculum slightly convex and short rostrate, with bright orange rim; leaf margins bistratose, distal leaf lamina partially to almost completely bistratose ..... Orthotrichum mazimpakanum

Orthotrichum anodon endostome segments ± white, Orthotrichum mazimpakanum exostome segments covering mouth (Garilleti et al. 2006. The inconspicuous, sparse around mouth (Garilleti et al. Bryologist 109: 188-196) 2011. The Bryologist 114: 346-355)

AL. Leaf apex rounded-obtuse, often somewhat toothed; leaves smooth to inconspicuously papillose; costa sinuose; mosses seasonally inundated ..... AM

AL' Leaf apex acute to acuminate or mucronate, usually entire; leaves smooth to densely papillose; costa straight; mosses not typically inundated ..... AN

Orthotrichum rivulare rounded-obtuse leaf apex, Orthotrichum shevockii acute leaf apex, not toothed toothed, sinuose costa (Norris 100722, UC (Norris 87128, UC 1759935) 1759034)

AM. Gametophytes dark green to nearly black, sometimes dark-red brown, in compact tufts; apex entire to irregularly serrulate; distal laminal cells approaching 20 µm; epilithic, in headwaters .....Orthotrichum euryphyllum

AM' Gametophytes rich dark green, sometimes black on older portions, in loose patches; apex irregularly dentate; distal laminal cells mostly to 12 µm; epiphytic or epilithic, in frequently inundated flood-plains of large streams ..... Orthotrichum rivulare

Orthotrichum euryphyllum slightly serrulate apex Orthotrichum rivulare irregularly dentate apex (Shevock 32504, CAS 1111476) (Lenz 630, UC 1920649)

AN. Exostome erect or spreading, with striate ridges on dorsal face resembling a fingerprint pattern; endostome absent; leaves often with bistratose streaks; epilithic .....

AN' Exostome reflexed to recurved, papillose to papillose-striate but never with a fingerprint-like pattern; endostome present; leaves unistratose or bistratose margins distal two-thirds of leaf; most species epiphytic ..... AQ

Orthotrichum cupulatum striate ridges on exostome Orthotrichum norrisii papillose exostome (Shevock teeth (Norris 84832, UC 1759913) 32529, UC 1933675)

AO. Capsule exserted and cylindrical, mostly with 8 long ribs alternating with 8 short ones ..... Orthotrichum anomalum—not yet verified from CA but reported from NV and

OR.

AO' Capsule immersed to emergent; with 8 or 16 ribs ..... AP

Othotrichum anomolum exserted capsules with 8 Orthotrichum cupulatum capsules immersed to ribs and 8 alternating short ribs (Allen 29190. CAS emergent (Norris 84832, UC 1759913) 1135847)

AP. Capsule with 16 ribs; leaves very weakly keeled, acute; gametophyte not glaucous

..... 2Orthotrichum cupulatum

AP' Capsule with 8 ribs; leaves distally keeled, obtuse to bluntly acute; gametophyte glaucous ..... Orthotrichum pellucidum

Orthotrichum cupulatum capsule emergent with 16 Orthotrichum pellucidum capsule with 8 ribs, plant ribs (Norris 84832, UC 1759913) glaucous (Shevock 32482, UC 1933690)

AQ. Leaf margins bistratose in distal two-thirds of leaf; capsules immersed with seta less than 0.5 mm long ..... 2Orthotrichum shevockii

AQ' Leaf margins unistratose; length of seta various ..... AR

Orthotrichum shevockii bistratose leaf margins Orthotrichum cucullatum capsules emergent (Norris (Norris 87128, UC 1759935) 106515, UC 1768584)

AR. Capsules fully exserted with seta visible ..... 2Orthotrichum coulteri

AR' Capsules immersed to emergent ..... AS

Orthotrichum coulteri exserted capsules (Norris Orthotrichum flowersii emergent capsule (Coleman 109508, UC 1917203) 1350)

AS. Capsule more than four times as long as wide when wet ..... AT

AS' Capsule much less than four times as long as wide when wet ..... AV

Orthotrichum norrisii urn longer than wide Orthotrichum alpestre urn wide (Shevock 14804, (Shevock 32529, UC 1933675) UC 1712492)

AT. Leaf apices obtuse or rounded and frequently cucullate ..... 2Orthotrichum cucullatum

AT' Leaf apices acute and flat to channeled ..... AU

Orthotrichum cucullatum obtuse leaf apex (Norris Orthotrichum norrisii acute leaf apex (Norris 80668, UC 1768584) 55099A, UC 1757638)

AU. Distal leaf lamina unistratose with rarely scattered bistratose ridges; basal marginal cells often quadrate, thick walled; axillary hairs 700–1200 µm long, multiseriate and branched, in dense fascicles, forming tomentum on stems; leaves distinctly narrowed with strongly revolute margins in distal half of leaf; apex acuminate to subulate .....

2Orthotrichum pilosissimum—not yet verified from CA, reported from NV.

AU' Leaf lamina unistratose entire length; basal marginal cells thin walled, occasionally thick walled and nodose; axillary hairs 75–250 µm long, filiform or subclaviform, uniseriate, not branched; leaves broader .....2Orthotrichum norrisii

Orthotrichum pilosissimum axillary hairs long, Orthotrichum norrisii axillary hairs short, branched, multiseriate (Shevock 22397, CAS unbranched, and uniseriate (Shevock 32529, UC 1040002) 1933675)

AV. Tip of leaf nearly obtuse except for one-cell mucron; distal one-fourth of leaf tongue-shaped (after narrowing from broader basal half); calyptrae mostly hairless .....

2Orthotrichum flowersii

AV' Tip of leaf acute; distal one-fourth of leaf triangular; calyptrae hairy to nearly hairless ..... AW

Orthotrichum flowersii apex with one-celled Orthotrichum alpestre acute leaf tip (Shevock mucron (Coleman 1350) 14804, UC 1712492)

AW. Subsidiary cells with strongly thickened radial walls that obscure outline of guard cells below; leaf papillae high, sometimes branched; exostome teeth striate-papillose distally ..... AX

AW' Subsidiary cells around stomates without thickened radial walls, thickened only at tips, thus guard cells only partially covered by subsidiary cells; leaf papillae low; exostome teeth uniformly papillose, not striated distally..... AY

Orthotrichum alpestre with strongly thickened Orthotrichum pumilum subsidiary cells thickened radiall walls, obscuring outline of guard cells only at tips, guard cells somewhat covered (Norris (Shevock 14804, UC1712492) 77959, UC1675390)

AX. Capsules ribbed throughout, short-cylindric; exostome teeth 16, erect when mature, reflexed when old; endostome teeth 16, filiform ..... Orthotrichum underwoodii— endemic to Santa Clara mountains, below 2000 m elevation.

AX' Capsules lightly ribbed primarily toward strangulate mouth, mostly oblong in shape; exostome teeth 8, reflexed; endostome with 8 segments ..... Orthotrichum alpestre— widespread above 2000 m elevation.

Orthotrichum underwoodii peristome with 16 Orthotrichum alpestre peristome with 8 endostome endostome and 16 exostome teeth (Toren 9511, and 8 exostome teeth (Shevock 14804, UC CAS 1048109) 1712492)

AY. Leaves parallel-sided from half to three-fourths distally; endostome with 16 segments; calyptra plicate; capsule gradually tapering to seta; distal laminal cells less than 14 µm ..... Orthotrichum pallens

AY' Leaves gradually tapering from base to three-fourths of total length; endostome with

8 segments; calyptra smooth; capsule abruptly narrowing into seta; distal laminal cells more than 13 µm; gemmae sometimes present ..... 2Orthotrichum pumilum

Orthotrichum pallens leaves parallel over three- Orthotrichum pumilum leaf broadly lanceolate, fourths their length, calyptra plicate (© Michael tapering from a broad base (Norris 77959, UC Luth, US Department of Agriculture) 1675390)

AZ. Multicellular gemmae densely covering leaves; pre-basal marginal cells with high pointed papillae on cell wall ..... 2Orthotrichum lyellii

AZ' Multicellular gemmae not densely covering leaves ..... BA

Orthotrichum lyellii gemmae densely covering Orthotrichum obtusifolium gemmae not densely leaves (Coleman 2) covering leaves (Wilson 607, SFV 19764)

BA. Leaves terminating in a hyaline awn ..... 2Orthotrichum diaphanum

BA' Leaf tip at most irregularly bleached, not hyaline ..... BB

Orthotrichum diaphanum leaf with hyaline awn Orthotrichum obtusifolium leaf tip lacking hyaline (Shevock 19240, UC 1739285) awn (Norris 68813, UC 1643011)

BB. Leaf lamina uniformly bistratose; gametophytes with prostrate stems consisting of differentiated vegetative and fertile uprights (unlike other Orthotrichum species); vegetative leaves nearly obtuse .....2Orthotrichum kelmanii—stomata cryptoporous.

BB' Leaf lamina unistratose or occasionally scattered bistratose streaks ..... BC

Orthotrichum kelmanii differentiated fertile (left) Orthotrichum coulteri leaf lamina with bostratose and vegetative (right) stems (Shevock 29890, UC streaks (Norris 109508, UC 1917203) 1872681)

BC. Each leaf cell with a single papilla over lumen on abaxial and adaxial surfaces .....

2Orthotrichum obtusifolium

BC' Leaves smooth to having 1-3 small conic papillae per cell ..... BD

Orthotrichum obtusifolium single papilla on lumen Orthotrichum pumilum leaves smooth (Norris on both abaxial and adaxial surfaces (Wilson 607, 77959, UC 1675390) SFV 19764)

BD. Capsules fully exserted; subsidiary cells with strongly thickened radial walls that obscure outline of guard cells below; leaves not decurrent ..... 2Orthotrichum coulteri

BD' Capsules immersed to emergent ..... BE

Orthotrichum coulteri exserted capsules (Norris Orthotrichum flowersii emergent capsules (Coleman 109508, UC1917203) 1350)

BE. Leaf apices channeled ..... BF

BE' Leaf apices more or less flat, acute to acuminate, never channeled or cucullate .....

Orthotrichum norrisii leaf apex channeled (Norris Orthotrichum pumilum leaf apex flat (Norris 77959, 55099A, UC 1757638) UC 1675390)

BF. Axillary hairs hyaline, distinctively long, protruding from foliose shoots; capsule cylindric, strongly constricted when dry; leaves unistratose or distal portion rarely with scattered bistratose streaks .....2Orthotrichum pilosissimum— not yet verified from CA, reported from NV.

BF' Axillary hairs not long, not protruding from foliose shoots; capsule cylindric, weakly or not constricted when dry; leaves unistratose ..... 2Orthotrichum norrisii

Orthotrichum pilosissimum axillary hairs Orthotrichum norrisii axillary hair (Shevock 32529, distinctively long, hyaline (Shevock 22397, CAS UC 1933675) 1040002)

BG. Tip of leaf nearly obtuse except for one-celled mucron; distal one-fourth of leaf tongue-shaped (after narrowing from broader basal half); calyptrae mostly hairless .....

2Orthotrichum flowersii

BG' Tip of leaf acute; distal quarter of leaf triangular; calyptrae sparsely hairy .....

2Orthotrichum pumilum

Orthotrichum flowersii leaf tip with one celled- Orthotrichum pumilum triangular leaf tip (Norris mucron (Coleman 1350) 77959, UC 1675390)

Discussion

In addition to revising the Orthotrichum key, I added images of characters that are useful for species identification, thereby reducing the need to examine herbarium specimens. My photographic key is an important contribution to the endeavor of identifying Orthotrichum in California. First, it serves as a model for future keys in the

California Moss eFlora. Second, reliable identification is necessary for protecting rare species and extraordinary areas where sensitive species live. Third, identification is a prerequisite to investigating the origins of the flora—how species distributions correlate with climate, endemism, and phylogeny.

Orthotrichum might have the highest proportion of endemics of any comparably sized clade of mosses in California (or perhaps it will be the second highest after new species in the Bryaceae have been described: Carter et al. 2016). My key could be used to identify the 3,447 specimens of Orthotrichaceae already in herbaria (2016 query of

Consortium of North American Bryophyte Herbaria). Identifying a large number of those specimens would enable robust analyses of the distributions of the various species. Once georeferenced, species distributions could be plotted on dot maps (Blockeel et al. 2014) and subject to species distribution modeling. Comparing species distribution models, in turn, might shed light on diversity gradients and modes of diversification within the group. As a challenge for future workers, I discuss below how such models could be used to study diversification in the group by comparing the distributions of endemic, disjunct, and widespread species. I refer to geographic patterns that species in Orhotrichum appear

to exhibit, although data to support these trends have yet to be assembled. I end with predictions on the results of such a study.

Georeferenced localities are used in species distribution models (SDMS). SDMS summarize the relationship between known occurrences of a species and climatic variables, and are also referred to as ecological niche models. SDMs are used to geographically delineate the habitat of species (Irfan-Ullan et al. 2007; Rotenberry et al.

2006), to predict potential ranges of species (Dettmers & Bart 1999; Peterson 2001;

Anderson et al. 2002), and to assess factors that allow for the persistence of species

(Swenson 2006). Inputs into SDMs include climate layers and distribution data in a GIS framework. One SDM program that allows for a presence-only analysis is MaxEnt (Elith et al. 2006; Pearson et al. 2007). MaxEnt analysis of various Orthotrichum species would find the environmental variables that most account for the species’ distributions. Wiens &

Stralburg (2009) described this method as a correlative approach using environmental variables that characterize where a species occurs and that find the variables that might determine the distribution of the species.

SDMs have been used to study areas of endemism (Escalante et al. 2007). An

SDM could be used to test the assertion that Orthotrichum inhabits regions with a

Mediterranean climate, i.e., areas with hot dry summers and moist cool winters. The

SDMs of various species could be compared. For example, species could be separated into those that are endemic versus those that are more widespread. Then, for the non- endemics, a spatially explicit analysis of the climate variables could test for a relationship

with species richness. This would test the impression that Orthotrichum species occur predominantly in areas where the climate is the most Mediterranean.

The degree to which the climate of California is in fact Mediterranean varies. In northern California and beyond into Oregon, winters are wetter, whereas to the south, many fewer days per year are wet (Sawyer 2006). The higher elevations are somewhat less Mediterranean because snow accumulates on many of the surfaces where species of

Orthotrichum might otherwise grow, and thunderstorms occur in summer over the high mountains (Abatzoglou et al. 2009). Deserts occur in the rain shadow of the Sierra,

Transverse, and Peninsular Ranges (Barbour et al. 2007). The redwood forests of

California’s cool and foggy north coast and extending inland along river bottoms are famous for retaining moisture replenished by fog (Sawyer 2006).

Assuming one confirmed the pattern that Orthotrichum species richness is highest in the archetypical Mediterranean climate, one would then ask, how do the endemics differ with respect to climatic tendencies compared to the non-endemics? Brown (1957) envisioned what he called “a centrifugal mode of speciation” whereby species form through a series of geographic expansions into favorable habitat and contractions eroding ranges in unfavorable habitats. If Orthotrichum diversified in a centrifugal mode, then endemic species should occur in climates that are typical of the center of richness, so that their center of distribution is in the Mediterranean-type climate, e.g. the part of our region with truly hot dry summers yet moist snow-free winters. In contrast, Mayr (1954) envisioned “a peripheral mode of speciation” whereby species form from populations that are isolated in somewhat different habitats. If Orthotrichum diversified in a peripheral

fashion, then endemic species would be expected to occur not in the core Mediterranean- climate areas but in the high mountains, the deserts, and the redwood forests, at the extremes of Orthotrichum’s climatic center.

For vascular plants, California’s Central Coast ranges, with their archetypical

Mediterranean climate, present the most prominent centers of neoendemism (Stebbins &

Major 1965; Thorne 2009; Kraft et al. 2010; Baldwin 2014). In contrast, the Klamath

Region of northwest California and southwest Oregon has a concentration of paleoendemics (Sawyer 2006). Paleoendemics are species that once occupied a larger geographic range and have become more restricted geographically over geological time as a result of environmental changes, whereas neoendemics are species that arose in the small area(s) that they continue to occupy. Phylogenetically, neoendemics are known by how they are nested closely among related species, whereas paleoendemics are on long branches attached near the base of their phylogeny (Mishler et al. 2014). What might be predicted of California’s endemic Orthotrichum species? Certain species, such as O. coulteri, O. cucullatum, O. franciscanum, and O. norrisii appear to be neoendemics of a highly Mediterranean climate, growing in the lower elevation mountains in

Mediterranean-type chaparral and woodlands (Medina et al. 2008). Other endemics, such as the rare O. kellmanii, which grows only in a small area overlooking the Pacific Ocean, are in a milder region of California, and perhaps such species are paleoendemics (Norris et al. 2004).

If the Mediterranean-climate is confirmed to be the center of richness, then the question arises, how much of the phylogeny is involved? Do the species that thrive in

Mediterranean climates constitute a subclade within the genus, the whole genus, or a group of genera? The genera of Orthotrichaceae other than Orthotrichum are not particularly associated with Mediterranean climates. Species of Ulota and Zygodon occur in California, but either tend to be more coastal or to have ranges more in the Pacific

Northwest (Goffinet et al. 1998). It also seems unlikely that Orthotrichum as a whole was simply subtended by the evolution of characters that favor a Mediterranean climate.

Apparently, the crytoporous clade of Orthotrichum (sometimes treated as a subgenus) arose from within the phaneroporous lineages, and the cryptoporous species appear to live in more arid climates (Lewinsky-Haapasaari & Hedenäs 1998). Considering only

Californian species, phaneroporous Orthotrichum may be less Mediterranean in their habitats than cryptoporous Orthotrichum species. It has been suggested that the crytopores radiated in relatively arid habitats compared to the phaneropores (Vitt 1971,

1973). However, this generalization is simplistic given that cryptoporous and phaneroporous species of Orthotrichum are found in similar habitats and that western

North America is home to many phaneroporous endemics (Paton & Pearce 1957; Sawicki et al. 2009; Medina et al. 2012).

Depending on climate, various Orthotrichaceae are epiphytic, epilithic, or both.

Consider three transitions away from core Mediterranean-type climate. (1) Comparing

Mediterranean-climate areas to more arid areas, mosses cease to inhabit bark, presumably because too few days of the year are moist (personal impression of experience collectors

D. Norris & J. Shevock). (2) Comparing Mediterranean-climate foothills to the upper

Sierran slope, epiphytes again drop out, although this may be partially the result of

angiosperm trees giving way to an exclusively coniferous forest (Coleman 2014). (3)

Comparing Mediterranean-climate areas to the humid coastal forests, bryophytes not in the Orthotrichaceae become abundant on trees—even on conifers—thus, perhaps

Orthotrichaceae are or were out-competed on the large trunks and they mainly live toward the branch tips.

In Orthotrichum, many geographic disjunctions are apparent. Orthotrichum columbicum has a trans-Atlantic distribution. This species is related to three species that are endemic to western North America: O. consimile, O. persimile, and O. confusum

(Medina et al. 2012). These species all live on the branches of trees and shrubs in humid forests. Orthotrichum columbicum is scattered in western Europe and the Mediterranean

Basin as well as in western North America (Medina et al. 2012). Another trans-Atlantic distribution involves O. tenellum and O. norrisii. They presumably diverged allopatrically after long-distance dispersal given that they are endemic to Mediterranean climate areas in the Old World and western North America, respectively (Medina et al.

2012). A special case of disjunction occurs with O. acuminatum. This species is found in western North America, the Mediterranean Basin, and eastern Africa. It has differentiated genetically among the three areas but not morphologically (Vigalondo et al. 2016).

Are the geographic patterns of Orthotrichum similar to or different from the patterns of vascular plants in California that have stimulated so much evolutionary theory? As a moss, Orthotrichum is desiccation tolerant, enabling species in this group to inhabit sun-exposed meso-habitats (albeit usually in shady microhabitats). By thriving in areas with summer drought, the Orthotrichum clade was apparently predisposed to

speciate in California. While some rules that shape vascular plant distributions may apply to mosses (Harrison et al. 1971; Woodward & Williams 1987; Kelly & Goulden 2008;

Baldwin 2014), we cannot ignore the physiology of mosses (Goffinet & Shaw 2008), their great dispersal ability (Miles & Longton 1992; Stoneburner et al. 1992; Marshall &

Convey 1997), or their ability to inhabit tree trunks and rocks. These characteristics play a greater role in moss biogeography than in the biogeography of vascular plants (Mishler

1988; Oliver et al. 2000; Kroemer et al. 2004; Jenks & Hasegawa 2005). For vascular plants, the Mediterranean-type climate of California favors neoendemism because of the close interdigitation of habitats (Stebbins & Major 1965). Close interdigitation sets the stage for centers of endemism to evolve in organisms with limited dispersal abilities.

Such organisms under those circumstances tend to differentiate into varying ecotypes and then into varying species (Stebbins 1974; Wiens 2011). Since the biology of mosses differs, it would be interesting to test whether these generalizations fail to apply to biogeographic patterns in Orthotrichum.

I conclude with hypotheses concerning the expansion and radiation of

Orthotrichum species in western North America. I hypothesize that speciation in

Orthotrichum occurred in a centrifugal fashion and that neoendemic species will be located at the center of Orthotrichum’s climatic center. Furthermore, I predict that paleoendemics will occur in more peripheral areas that resemble the widespread climate of the Tertiary (i.e., wetter areas such as the humid foggy coasts of the Pacific

Northwest). Next, I predict that there is no single clade of the phylogeny associated with the adaptation for Mediterranean climates. Rather, I would favor the notion that such

adaptations predate occupancy of this type of climate and that multiple species and clades of Orthotrichum evolved in parallel in the Mediterranean-climate regions of the Northern

Hemisphere. I suspect that California was colonized long ago both by Orthotrichum from moister adjacent areas and subsequently by long-distance dispersal from the

Mediterranean Basin. Furthermore, the pattern of some endemics suggests that they many have their closest relatives nearby, whereas others diverged allopatrically following disjunctions.

These suggested patterns and hypotheses can only be tested if specimens are identified by today’s standards. In addition, some of the patterns ought to be tested with a well-supported phylogeny. With the verification of specimens in Orthotrichum, using the new photographic key, a comparison of SDMs will shed light into their diversification and how this relates to the Mediterranean-type climate of California. This will also reveal the general climate niche of Orthotrichum and perhaps suggest what environments and characters are involved in the adaptive evolution of various Orthotrichum species.

References

Abatzoglou, J. T., K. T. Redmond & L. M. Edwards. 2009. Classification of regional climate variability in the state of California. Journal of Applied Meteorology and Climatology 48: 1527–1541.

Anderson, R. P., M. Laverde & A. T. Peterson. 2002a. Geographical distributions of spiny pocket mice in South America: Insights from predictive models. Global Ecology and Biogeography 11: 131–141.

Anderson, R. P., M. Laverde & A. T. Peterson. 2002b. Using niche-based GIS modeling to test geographic predictions of competitive exclusion and competitive release in South American pocket mice. Oikos 93: 3–16.

Baldwin, B. G. 2014. Origins of plant diversity in the California Floristic Province. Annual Review of Ecology, Evolution, and Systematics 45: 347–369.

Barbour, M., T. Keeler-Wolf & A. A. Schoenherr. 2007. Terrestrial vegetation of California, 3rd edition. University of California Press, Berkeley, CA.

Bell, C. R. 1967. Plant Variation and Classification. Wadsworth Publishing, Belmont, CA.

Blockeel, T. L., S. D. S. Bosanquet, M. O. Hill & C. D. Preston. 2014. Atlas of British and Irish Bryophytes. Pisces Publications, Newbury, England.

Brown, W. J. 1957. Centrifugal speciation. Quarterly Review of Biology 32: 247–277.

Carter, B. E., B. Shaw & A. J. Shaw. 2016. Endemism in the moss flora of North America. American Journal of Botany 103:769–779.

Coleman, L. A. 2014. Bryophyte diversity and niche relations along a 3000 M elevational gradient in Sequoia National Park. California State University, Northridge.

Davis, P. H. & V. H. Heywood. 1963. Principles of Angiosperm Taxonomy. D. Van Nostrand, Princeton, NJ.

Dettmers, R. & J. Bart. 1999. A GIS modeling method applied to predicting forest songbird habitat. Ecological Applications 9: 152–163.

Elith, J., C. H. Graham, R. P. Aderson, M. Dudík, S. Ferrier, A. Guisan, R. J. Hijmans, F. Huettmann, J. R. Leathwick, A. Lehmann, J. Li, L. Lohmann, B. A. Loiselle, G. Manion, C. Moritz, M. Nakamura, Y. Nakazawa, J. McC. M. Overton, A.

Townsend Peterson, S. J. Phillips, K. Richardson, R. Scachetti-Pereira, R. E. Schapire, J. Soberón, S. Williams, M. S. Wisz & N. E. Zimmerman. 2006. Novel methods improve prediction of species’ distributions from occurrence data. Ecography 29: 129–151.

Escalante, T., V. Sanchez-Cordero, J. J. Morrone & M. Linaje. 2007. Areas of endemism of Mexican terrestrial mammals: a case study using species’ ecological niche modeling, parsimony analysis of endemicity and Goloboff fit. Interciencia 32: 151–159.

Faubert, J. 2014. Flore des bryophytes du Québec–Labrador. Volume 3: Mousses, second partie. 455 pp. Société Québécoise de Bryologie, Saint-Valérien, Québec.

Fischer, M. A. 2010. Do plant identification keys enable identification? Phytologia Balcanica 16: 175–185.

Flora of North America Editorial Committee [R. H. Zander & P. M. Eckel, eds]. 2014. Flora of North America North of Mexico, Volume 28: Bryophyta, part 2. 702 pp. Oxford University Press, New York, NY.

Fransworth, E. J., M. Chu, W. J. Kress, A. K. Neill, J. H. Best, J. Pickering, R. D. Stevenson, G. W. Courtney, J. K. VanDyk & A. M. Ellison. 2013. Next- generation field guides. BioScience 63: 891–899.

Garilleti, R. & F. L. Mazimpaka. 2006. On the presence of Orthotrichum shawii (Orthotrichaceae) in California. The Bryologist 109: 510–515.

Garilleti, R., F. Lara, V. Mazimpaka, D. Bot, F. De. Farmacia & D. Valencia. 2006. Orthotrichum anodon (Orthotrichaceae), a new species from California, and its relationships to other Orthotricha with puckered capsule mouths. The Bryologist 109: 188–196.

Goffinet, B., R. J. Bayer & D. H. Vitt. 1998. Circumscription and phylogeny of the Orthotrichales (Bryopsida) inferred from rbcL sequence analyses. American Journal of Botany 85: 1324–1337.

Goffinet, B. & A. J. Shaw. 2008. Bryophyte Biology. Cambridge University Press, New York, NY.

Griffing, L. R. 2011. Who invented the dichotomous key? Richard Waller’s watercolors of the herbs of Britain. American Journal of Botany 98: 1911–1923.

Guerra, J., M. J. Cano & M. Brugués (eds). 2014. Flora Briofítica Ibérica, Volumen V. 264 pp. Sociedad Espanola de Briologia, Murcia, Murcia.

Harrison, A. T., E. Small & H. A. Mooney. 1971. Drought relationships and distribution of two Mediterranean-climate California plant communities. Ecology 52: 868– 875.

Hedenäs, L. 2001. Environmental factors potentially affecting character states in pleurocarpous mosses. The Bryologist 104: 72–91.

Hedenäs, L., C. Reisborg & T. Hallingbäck. 2014. Nationalnyckeln till Sveriges Flora och Fauna: Bladmossor, Kompaktmossor-kapmossor. 366 pp. ArtDatabanken (Swedish Threatened Species Unit), Uppsala, Sweden.

Irfan-Ullah, M., G. Amaranth, M. S. R. Murthy & A. T. Peterson. 2007. Mapping the geographic distribution on Aglaia bourdillonii Gamble (Meliaceae), an endemic and threatened plant, using ecological niche modeling. Biodiversity and Conservation 16: 1917–1925.

Jarvie, J. K. & P. F. Stevens. 1998. Interactive keys, inventory, and conservation. Conservation Biology 12: 222–224.

Jenks, M. A. & P. M. Hasegawa. 2005. Plant Abiotic Stress. Blackwell Publications, Oxford, England.

Karttunen, K. 1984. Zygodon gracilis, Z. rupestris and Z. dentatus, comb. nova (Musci, Orthotrichaceae): History and nomenclature. Annales Botanici Fennici, 21: 343- 348.

Kelly, A. E. & M. L. Goulden. 2008. Rapid shifts in plant distribution with recent climate change. Proceedings of the National Academy of Sciences, USA 105: 11823– 11826.

Kraft, N. J. B., B. G. Baldwin & D. D. Ackerly. 2010. Range size, taxon age and hotspots of neoendemism in the California flora. Diversity and Distributions 16: 403–413.

Kroemer, K., R. Reski & W. Frank. 2004. Abiotic stress response in the moss Physcomitrella patens: evidence for an evolutionary alteration in signaling pathways. Plant Cell Reports 22: 864–870.

Lamarck, J. P. 1779. Flore Francoise. Ed. 1. Imprimerie Royale, Paris, France.

Leggett, R. & B. K. Kirchoff. 2011. Image use in field guides and identification keys: review and recommendations. AoB Plants 11: 1–37.

Lewinsky, J. 1993. A synopsis of the genus Orthotrichum Hedw. (Musci, Orthotrichaceae). Bryobrothera 2: 1–59.

Lewinsky-Haapasaari, J. & L. Hedenäs. 1998. A cladistics analysis of the moss genus Orthotrichum. The Bryologist 101: 519–555.

Malcolm, B. & N. Malcolm. 2006. Mosses and Other Bryophytes, an Illustrated Glossary, 2nd Ed. Micro-Optics Press, Nelson, New Zealand.

Malcolm, B., N. Malcolm, D. Norris & J. Shevock. 2009. California Mosses. Micro- Optics Press, Nelson, New Zealand.

Marshall, W. A. & P. Convey. 1997. Dispersal of the moss propagules on Signy Island, maritime Antarctic. Polar Biology 18: 376–383.

Mayr, E. 1954. Change of genetic environment and evolution. Pp. 157–180 in J. Huxley, A. C. Hardy & E. B. Ford (eds). Evolution as a Process. Allen and Unwin, London, England.

Medina, R., F. Lara, V. Mazimpaka & R. Garilleti. 2008. Orthotrichum norrisii (Orthotrichaceae), a new epiphytic Californian moss. The Bryologist 111: 670– 675.

Medina, R., F. Lara, B. Goffinet, R. Garilleti & V. Mazimpaka. 2012. Integrative taxonomy successfully resolves the pseudo-cryptic complex of the disjunct epiphytic moss Orthotrichum consimile s.l. (Orthotrichaceae). Taxon 61: 1180– 1198.

Medina, R., F. Lara, B. Goffinet, R. Garilleti & V. Mazimpaka. 2013. Unnoticed diversity within the disjunct moss Orthotrichum tenellum s.l. validated by morphological and molecular approaches. Taxon 62: 1133–1152.

Méndez, N. 2015. New volumes in multivolume moss floras. The Bryologist 118: 116– 117.

Miles, C. J. & R. E. Longton. 1992. Deposition of moss spores in relation to distance from parent gametophytes. Journal of Bryology 17: 355–368

Mishler, B. D. 1988. Reproductive ecology of bryophytes. Pp. 285–306 in J. Lovett- Doust & L. Lovett-Doust (eds). Plant Reproductive Ecology: Patterns and Strategies. Oxford University Press, New York, NY.

Mishler, B., K. Nunzio, C. E. González-Orozco, A. H. Thornhill, S. W. Laffan & J. T. Miller. 2014. Phylogenetic measures of biodiversity and neo- and paleo- endemism in Australia Acacia. Nature Communications 5: 1–10.

Norris, D. H. & J. R. Shevock. 2004. Contributions toward a bryoflora of California: II A key to the mosses. Madroño 51: 133–269. 86

Norris, D. H., J. R. Shevock & B. Goffinet. 2004. Orthotrichum kellmanii (Bryopsida, Orthotrichaceae), a remarkable new species from the central coast of California. The Bryologist 107: 209–214.

Oliver, M. J., J. Velten & A. J. Wood. 2000. Bryophytes as experimental models for the study of environmental stress tolerance: Tortula ruralis and desiccation-tolerance in mosses. Plant Ecology 51: 73–84

Paton, J. A. & J. V. Pearce. 1957. The occurrence, structure and functions of stomata in British bryophytes. Transactions of the British Bryological Society 3: 228–259.

Pearson, R. G., C. J. Raxworthy, M. Nakamura & A. T. Peterson. 2007. Predicting species distributions from small numbers of occurrence records: a test case using cryptic geckos in Madagascar. Journal of Biogeography 34: 102–117.

Peterson, A. T. (2001) Predicting species’ geographic distributions based on ecological niche modeling. Condor 103: 599–605.

Porter, C. L. 1959. Taxonomy of Flowering Plants. W. H. Freeman, San Francisco, CA.

Rotenberry, J. T., K. L. Preston & S. T. Knick. 2006. GIS-based niche modeling for mapping species’ habitat. Ecology 87: 1458–1464

Sawicki, J., P. Vítězslav & M. Szczecińska. 2009. Preliminary studies on the phylogeny of Orthotrichum (Bryophyta) inferred from nuclear ITS sequence. Annales of Botanici Fennici 46: 507–515.

Sawyer, J. O. 2006. Northwest California: A Natural History. University of California Press, Berkeley, CA.

Scharf, S. T. 2009. Identification keys, the “natural method,” and the development of plant identification manuals. Journal of the History of Biology 42: 73–117.

Sharp, A. J., H. Crum & P. M. Eckel. 1994. The Moss Flora of Mexico. Memoirs of the New York Botanical Garden 69, New York, NY.

Sokal, R. R. 1985. The continuing search for order. American Naturalist 126: 729–749.

Stace, C. A. 1989. Plant Taxonomy and Biosystematics. 2nd ed. Cambridge University Press, Edward Arnold, London.

Stebbins, G. L. 1974. Flowering plants: evolution above the species level. Harvard University Press, Cambridge, MA [chapter 8].

Stebbins, G. L. & J. Major. 1965. Endemism and speciation in the California flora. Ecological Monographs 35: 1–35.

Stonebruner, A., D. M. Lane & L. E. Anderson. 1992. Spore liberation distances in the moss Atrichum angustatum (Polytrichaceae). The Bryologist 95: 324–328.

Swenson, N. G. 2006. GIS-based niche models reveal unifying climatic mechanisms that maintain the location of avian hybrid zones in a North American suture zone. Journal of Evolutionary Biology 19: 717–725.

Thorne, J. H., J. H. Viers & J. Price. 2009. Spatial patterns of endemic plants in California. Natural Areas Journal 29: 344–366.

Vigalondo, B. F. Lara, I. Draper, V. Valcarcel, R. Garilleti & V. Mazimpaka. 2016. Is it really you, Orthotrichum acuminatum? Ascertaining a new case of intercontinental disjunction in mosses. Botanical Journal of the Linnean Society 180: 30–49.

Vitt, D. H. 1971. The infrageneric evolution, phylogeny, and taxonomy of the genus Orthotrichum (Musci) in North America. Nova Hedwigia 21: 683–711.

Vitt, D. H. 1973. A revision of the genus Orthotrichum in North America, north of Mexico. Bryophytorum Bibliotheca 1: 1–208.

Waller, R. 1686. A catalogue of simple and mixt colours, with a specimen of each colour prefixt to its proper name. Philosophical Transactions of the Royal Society B 16: 24–32.

Walter, D. E. & S. Winterton. 2007. Keys and the crisis in taxonomy: extinction or reinvention? Annual Review of Entomology 52: 193–208.

Wiens, J. A., D. Stralburg, D. Jongsomjit, C. A. Howell & M. A. Snyder. 2009. Niches, models, and climate change: assessing the assumptions and uncertainties. Proceedings of the National Academy of Sciences, USA 106: 19729–19736.

Wiens, J. J. 2011. The niche, biogeography and species interactions. Philosophical Transactions of the Royal Society of London B 366: 2336–2350.

Woodward, F. I. & B. G. Williams. 1987. Climate and plant distribution at global and local scales. Advances in Vegetation Science 8: 189–197.

Appendix: Useful Specimens Examined

Orthotrichum acuminatum U.S.A. California: Riverside Co., 33° 50' 13.92"N -116° 48' 32.0394"W. Quercus forest with scattered conifers, elevation 4940 ft. Trunk of Quercus wislizenii in filtered light. 16.XI.2008, Shevock 32571 (CAS 1146075)

Orthotrichum affine U.S.A. California: Modoc Co., 41° 26'N -121° 03'W. Modoc National Forest: Cottonwood Flat Campground, in Pseudotsuga forest on moist diffusely lit bark of Populus, elevation 4921 ft., 21.IV.1983, 21.IV.1983, D. H. Norris 68516 (SFV 20954) California: Lassen Co., 40° 19’N -121°01’W. Lassen National Forest: in Pinus ponderosa forest, 22.XI.1971, D. H. Norris 19236 (SFV 21208)

Orthotrichum alpestre U.S.A. California: Tulare Co., 36° 2' 59.9994"N -118° 7' 44.7594"W, 16.II.1997, Shevock 14804 (UC 1712492)

Orthotrichum anomolum U.S.A. Maine: Knox Co., 44° 11' 56.76"N -69° 6' 30.9594"W, Town of Rockport. Simonton Corners Quarry Preserve., X.VI.2009, Allen 29190 (CAS 1135847) Utah: Iron Co., 37° 37' 7.1”N -125° 0' 11.34"W, Cedar City, on private land, elevation 7850 ft. 20.IV.2014, Weiss 20157 (CAS 1220879)

Orthotrichum bolanderi U.S.A. California: Placer Co., 38°51' 0"N -121°12'59.7594"W, 18.I.1994, D. H. Norris 81945 (UC 1774444) California: Tulare Co., 36° 20' 49.2"N -118° 45' 28.7994"W, Sequoia National Forest, elevation 3772 ft., 28.I.1998, D. H. Norris 92748 (UC 1774447) California: Tulare Co., 36° 9' 43.1994"N -118° 41' 27.6"W, Southern Sierra Nevada, Sequoia National Forest, elevation 3200 ft., 17.III.1996, J. R Shevock 13100 (UC 1730763) California: Riverside Co., 33° 30' 14.3994" N -117.283 W, Peninsular Ranges in oak savanna and vernal pool community, Santa Ana mountains, elevation 2500 ft., 28.IV.1975, J. R Shevock 4273 (UC 1710769) California: Los Angeles Co., 34°5'24"N -118° 43' 11.9994"W, Malibu Creek, Santa Monica Mountains, elevation 760 ft., 23.V.2004, T. Sagar 372 (SFV 20461) California: Placer Co., 38° 50' 56.3994"N -121° 12' 57.5994"W, Quercus agrofolia forest, elevation 1968 ft., 18.I.1994, J. R Shevock 81945 (UC 1774444) California: Ventura Co., 34°7'37.2"N - 118° 55' 55.2"W. Santa Monica Mountain: Circle X Ranch, Carlisle Canyon, on Quercus agrifolia woodland, elevation 2329 ft., 27.XII.2003, T. Sagar 211 (SFV 20462)

Orthotrichum columbianum U.S.A. California: Del Norte Co., 41° 54' 12.9594"N -123° 51' 16.9194"W, 27.II.2013, Shevock 41858 (CAS 1187203)

Orthotrichum consimile U.S.A. California: Plumas Co., 39° 36' 46.44"N -121° 3' 14.76"W, 5.VI.2005, Dillingham 2006 (UC 1932571); Humboldt Co., 40° 46' 59.8794"N -123° 46' 59.8794"W, 29.X.1977, Norris 50401 (UC 1722823) California: Humboldt Co., 40° 42’N -123° 56’W. On moist diffusely lit twigs of Acer macrophyllum in riverine forest with Pseudotsuga, Thuja, and Acer along small stream, elevation 1312 ft., 10.II.1983, D. H. Norris 68267 (SFV 21220) Oregon: Benton Co., 44° 38' 23.6394"N -123° 17' 25.7994"W. Mc Donald/Dunn Research Forest: on fallen twig of grand fir with Orthotrichum papillosum, elevation 600 ft. 24.III.2008, P. Wilson 5005 (SFV 21630) California: Humboldt Co. Humboldt State University: On maple between Founders hall and the Art building, elevation 200 ft., 14.II.1984, P. Wilson 95 (SFV 19754) California: Humboldt Co., in second growth redwood forest, on Acer macrophyllum, elevation 500 ft., 28.XII.1983, P. Wilson 83 (SFV 19755)

Orthotrichum coulteri U.S.A California: Contra Costa Co., 37° 53' 42" N -122° 14' 42" W, 29.XI.2006, Norris 109508 (UC 1917203) California: Alameda Co., Tilden Regional Park: George Trail toward lake Anza, 29.XI.2006, Norris 109508 (UC 1917203)

Orthotrichum cucullatum U.S.A. California: Contra Costa Co. On trail, south shore of Briones Reservoir, 1.XI.2003, Norris 106515 (UC 1768584) California: Los Angeles Co., 34°23’N -118°27’W. San Gabriel Mountains: Placerita Canyon Park, Walker Ranch, on Fraxinus in partial shade, elevation 1500 ft., 9.III.2002, P. Wilson 3764 (SFV 23079)

Orthotrichum cupulatum U.S.A California: Tulare Co., 36° 7' 59.8794"N -118° 36' 59.76"W, 27.III.1996, Norris 87675 (UC 1759893) California: Shasta Co., 40° 52' 59.8794"N -122° 13' 59.8794"W, 5.II.1995, Norris 84832 (UC 1759913) California: Shasta Co., 40°47’N -122°00’W. 1 mi. north of Mineral School at intersection of road to Oak Run, on moist, diffusely lit limestone outcrop in mixed conifer forest, elevation 1312 ft., 22.IV.1983, D. H. Norris 68614 (SFV 20955) California: Trinity Co., 40° 46’N -123° 20’W. Ironside Mountain Quad: on limestone outcrop, elevation 110 ft., 20.III.1987, P. Wilson 1246 (SFV 19756) Oregon: Mutnomah Co., 45° 32' 13.1994"N -122° 13' 11.9994"W. Columbia River Gorge, Latourelle Falls, on vertical surface of concrete bridge, elevation 100 ft., 25.III.2008, P. Wilson 5049 (SFV 21570)

Orthotrichum diaphanum Mexico, Hidalgo: 19° 52' 59.8794"N -98° 25' 59.88"W, 25.IX.1997, UC1779402 U.S.A California: San Francisco Co., 37° 47' 27.5994"N -122° 25' 44.4"W, San Francisco Parks and Recreation areas: Lafayette Park, elevation 350 ft., 16.IV.2000, J. R. Shevock 19240 (UC 1739285)

California: San Francisco Co., 37° 46' 19.1994"N -122° 26' 27.6"W, San Francisco Parks and Recreation areas: Panhandle section of Park, elevation 230 ft., 14.II.2000, J. R. Shevock 18931 (UC 1739286)

Orthotrichum euryphyllum U.S.A. California: Lassen Co., 41° 8' 55.68"N -120° 30' 28.08"W, 30.X.2008, Shevock 32504 (UC 1933798) California: Lassen Co., 41°8’55.8” N -120° 30’ 28.4” W, Juniperus occidentalis woodland, elevation 5095 ft. 30.X.2008, J. R. Shevock 3205 (CAS1111476) California: Modoc Co., T41N R10E. Juniper/sagebrush, on volcanic rock, ft. 3.V.2005. C. Beyer 5039 (CAS 1182878)

Orthotrichum flowersi U.S.A. Nevada: NYE Co., 36° 58' 3"N -117° 5' 18.2394"W, 18.I.2003, Shevock 23691 (UC 1783158) California: Tulare Co., 36° 32' 0.5994"N -118° 46' 21.3594"W. Sequoia National Park: amongst Quercus wislizeni and Quercus douglasii, on soil slope shaded by trees and on Quercus douglasii trunk, elevation 3627 ft., 1.IV.2012, L. A. Coleman 1350 (SFV 22746) California: Tulare Co., 36°20'50.0"N 118°45'30.0"W. Sequoia National Park: in savannah on temporarily moist, diffusely lit bark of Aesculus in savannah with Quercus chrysolepis, Quercus wislizenii, Aesculus californica, Cercocarpus and Umbellularia in area of metamorphic rocks on south-facing slopes, elev. 3772 ft., 28.I.1998, D. H. Norris 92730 (UC1767850) California: Los Angeles Co., 34° 5' 52.7994"N - 118° 48' 46.8"W. Santa Monica Mountains: NPS Rocky Oaks Park, elevation 1706 ft., 8.III.2004, T. Sagar 241 (SFV20517)

Orthotrichum franciscanum U.S.A. California: San Francisco Co., 37° 44' 39.8394"N -122° 23' 15"W, 22.X.2000, Shevock 20202 (CAS 1193149)

Orthotrichum hallii U.S.A. California: Fresno Co., 36° 48' 14.4"N -118° 48' 57.6"W, Southern Sierra Nevada, Sequoia National Forest, elevation 5475 ft., 27.I.1996, J. R Shevock 13634 (UC 1730772) California: Shasta Co., 40° 45' 59.7594"N -121° 58' 59.8794"W, in Quercus kellogii and Pseudotsuga forest, elevation 410 ft., 30.I.1993, D. H. Norris 80330 (UC 1757602) Colorado: Conejos Co., 37° 8' 9.6"N -106° 22' 12"W. In Pseudotdsuga, Pinus ponderosa and Picea forest, elevation 9186 ft., 7.IX.1970, D. H. Norris 18242 (UC 1649714) Utah: Garfield Co., 37° 37' 2.9994"N -129° 9' 9"W. Bryce Canyon National Park: Abies concolor and Picea glauca forest, on diffusely lit rocks, elevation 7874 ft., 30.IX.1984, D. H. Norris 71713 (SFV 21236) California: Los Angeles Co., 34° 6' 35.9994"N -118° 46' 48"W, Santa Monica Mountains National Park, elevation 932 ft., 18.I.2005, T. Sagar 708 (SFV 20675) California: Trinity Co., 40° 24’00.0”N 123°01’00.0”W, Shasta –Trinity National forest: in forest with Pseudotsuga and Quercus chrysolepis in deep canyon over limestone at Hall City Caves, elevation 3937 ft., 15.IX.1984, D. H. Norris 71663 (UC 1669109)

Orthotrichum holzingeri U.S.A. California: Modoc Co., 41° 30' 59.7594"N -121° 2' 59.9994"W, 22.IV.1984, Norris 70467 (UC 1649719) Oregon: Grant Co., 44° 21' 0"N -119° 36' 59.76"W, 4.IX.1992, Norris 79089 (UC1649721) California: Modoc Co., 41°31'00.0"N 121°03'00.0"W. Modoc National Forest: On moist, diffusely lit rocks at headwaters of rivulet in flat and poorly drained area with Pinus ponderosa, Juniperus and Artemisia, elevation 5085 ft., 22.IV.1984, D. H. Norris 70467 (UC1649719) Oregon: Grant Co., 44°21'00.0"N 119°37'00.0"W. Ochoco National Forest: On moist, diffusely lit basaltic boulder in Pinus ponderosa forest, elevation 2952 ft., 24.IX.1992, D. H. Norris 79089 (UC1649721)

Orthotrichum kellmanii U.S.A. California: Monterey Co., 36° 13' 14.8794"N -121° 30' 52.92"W, 27.V.2007, Shevock 29890 (UC 1872681)

Orthotrichum laevigatum U.S.A. California: Madera Co., 37° 38' 24"N -119° 5' 24"W. Sierra Nevada Mountains: NW of Devil’s Postpile National Monument, on vertical face of granite in partial shade, elevation 7349 ft., 3.VIII.2006, P. Wilson 4486 (SFV 21283) California: Tulare Co., 36° 33' 34.452"N -118° 47' 0.924"W. Sequoia National Park: Crystal Cave Road, in mixed conifer forest with Abies concolor, Cornus nuttallii, and Calocedrus decurrens, on fallen log near stream base of slope, elevation 5464 ft., 13.VI.2011, L. A. Coleman 803 (SFV 22753) California: Tulare Co., 36° 33' 37.44"N -118° 46' 43.2834"W. Sequoia National Park: Slope above junction to Crystal Cave, in Abies concolor, Pinus jeffreyi, and Quercus kelloggii, on rock (side), elevation 5980 ft., 12.VI.2012, L. A. Coleman 1564 (SFV 22751) California: Fresno Co., 37°15’N -118°59’W. Sierra National Forest: 1-2 miles west of Florence at Dutch Lake, amid lodgepole pine forest, on granite outcrop, elevation 9186 ft., 9.IX.1984, P. Wilson 458 (SFV 23081) California: Siskyo Co. Klamath National Forest: in mixed conifer forest, on oak, elevation 5600 ft., 12.VI.1984, P. Wilson 243 (SFV 23084)

Orthotrichum lyellii U.S.A. California: Tulare Co., 36° 31' 12.324"N -118° 46' 42.348"W, 23.III.2009, L. A. Coleman 2 California: Madera Co., 37° 5' 45.348"N -119° 44' 24.1434"W, 8.IV.2014, Wilson 5356 California: Humboldt Co., 40° 38' 59.9994"N -123° 35' 56.4"W, Six Rivers National Forest: Abies concolor and Pseudotsuga young forest, elevation 1968 ft., D. H. Norris 83851 (UC 1774498) California: Tulare Co., 36° 20' 49.2"N -118° 45' 28.7994"W, Sequoia National forest: in savannah with Quercus wislizenii, Aesculus californica, Cercocarpus and Umbellularia, elevation 3772 ft., 28.I.1998, D. H. Norris 92731 (UC 1774518) California: Tulare Co., 36° 8' 27.5994"N -118° 35' 59.9994"W, Sequoia National Forest: Giant Sequoia-mixed conifer forest, elevation 5000 ft., 17.III.1996, J. R. Shevock 13124 (UC 1754019) California: Glenn Co., 39° 33' 57.6" N -122° 39' 28.8" W, Mendocino National Forest: black oak- foothill woodland, elevation 1800 ft., 2.VII.1997, J. R. Shevock 15803 (UC 1730605)

California: Mariposa Co., 37° 43' 30" N -119° 43' 4.8" W, Yosemite National Park: on bark of canyon live oak, Douglas fir and ponderosa pine, elevation 4425 ft., 30.IX.2000, J. R. Shevock 20177 (UC 1741882) California: San Benito Co., 36° 28' 37.1994" N -121° 11' 6" W, Pinnacles National Monument: volcanic boulders above stream channel with Quercus agrifolia, Adenostoma, and scattered Pinus sabiana, elevation 1400 ft., 2.I.2005, J. R. Shevock 26238 (UC 1798868) California: Santa Barbara Co., 34° 32' 23.9994" N -120° 2' 59.9994" W, Santa Ynez mountains, elevation 3900 ft., 11.II.2006, P. Wilson 4284 (SFV 20449) California: Santa Clara Co., 37° 11' 23.9994" N -121° 32' 24" W, Henry W. Coe State Park, elevation 2650 ft., 1.II.2006, T. Sagar s.n. (SFV 20565) California: Humboldt Co., coordinates unavailable, Lanphere-Christensen Dunes Preserve: Pinus/Picea forest over stabilized dunes with a dense Ericaceous/Garrya understory, elevation not recorded, 5.II.1985, P. Wilson 536 (SFV 197659) California: Mendocino Co., coordinates unavailable, Paperwood Springs Mountains: Mixed evergreen forest, elevation not recorded, 23.XI.1983, P. Wilson 50 (SFV 19760) California: Humboldt Co., coordinates unavailable, Sonoma Peninsula, elevation not recorded, 25.V.1984, P. Wilson 165 (SFV 19761) California: Humboldt Co., coordinates unavailable, Arcata, elevation not recorded, 13.V.1984, P. Wilson 147 (SFV 19763) California: Madera Co., 37° 5' 30.3714"N -119° 42' 55.944"W. San Joaquin experimental range: in area with scattered shrubs and trees, on limb of live oak, elevation 1131 ft., 21.XII.2008, P. Wilson 5189 (SFV 21503) California: Tulare Co., 36° 30' 44.6394"N -118° 47' 56.4"W. Sequoia National Park: near footbridge across middle fork of Kaweah River, on partly shaded tree branch amid oak forest, elevation 2000 ft., 11.I.2009, P. Wilson 5236 (SFV 21509)

Orthotrichum macounii U.S.A. Oregon: Wallowa Co., along road to Grizzly Ridge, ca. 2 miles E., 18.VII.1963, Hermann 18940 (CAS 444093) Idaho: Elmore Co., Boise National Forest., 1.VII.1942, MacFadden 19054 (CAS 441460)

Orthotrichum norrisii U.S.A. California: Humboldt Co., 40° 51' 59.7594"N -124° 3' 59.7594"W, -.-.1979, Norris 55099A (UC 1757638) Riverside Co., 33° 36' 38.1594"N -117° 26' 15"W, 14.XI.2008, J. R. Shevock 32529 (UC 1933675) California: Riverside Co., 33°26’N -117°12’W. Santa Margarita Reserve: on Sambucus neomexicana, elevation 600 ft., 26.III.2001, P. Wilson 3690 (SFV 23080) California: Los Angeles Co., 34° 4' 37.1994"N -118° 48' 57.6"W. Santa Monica Mountains: in Newton Canyon, on Salix sp., elevation 1394 ft., 12.VI.2004, P\T. Sagar 504 (SFV 23133)

Orthotrichum obtusifolium U.S.A. California: Trinity Co., 40° 51' 59.7594"N -122° 54' 59.76"W, 14.V.1983, D. H. Norris 68813 (UC 1643011) Michigan: Cheboygan Co., 45° 32' 53.1594"N -84° 40' 37.2"W, 20.VI.1985, P. Wilson 607 (SFV 19764)

California: Tehema Co., 40° 08’00”N -122° 50-52’00”W. In open Quercus forest, on dry sunny bark of Quercus in Pinus ponderosa and Abies concolor forest, II.XII.1980, D. H. Norris 56896 (UC 1650154) California: Modoc Co., 41° 26’00.0”N -121° 03’00.0”W. Modoc National Forest: in Pseudotsuga forest on moist diffusely lit bark of Populus, elevation 4921 ft., 21.IV.1983, D. H. Norris 68517 (UC 1650153) California: Modoc Co., 41° 26’00.0”N -121° 03’00.0”W. Modoc National Forest: in Pseudotsuga forest on moist diffusely lit bark of Populus, elevation 4921 ft., 21.IV.1983, D. H. Norris 68512 (UC 1650152) Michigan: Cheboygan Co. On Acer trunk, off Hogsback Rd., in a young-growth woodland, P. Wilson 607 (SFV 19764)

Orthotrichum papillosum U.S.A. California: Monterey Co., 35° 57' 3.6" N -121° 25' 15.6" W, Los Padres National Forest: open woodland of Quercus chrysolepis, Q. lobata, Arctostaphylos, Arbutus, and Pinus coulteri, elevation 3275 ft., 22.X.2005, J. R. Shevock 27717 (UC 1798628) California: Monterey Co., 36° 22' 40.8" N -121° 34' 4.7994" W, U.C. Hastings Natural History Reservation: woodland of California live oak and madrone, elevation 1830 ft., 21.III.2004, J. R. Shevock 24683 (UC 1782503) California: San Francisco Co., 37° 45' 21.6"N -122° 27' 7.1994"W, San Francisco Parks and Recreation areas: Eucalyptus and cypress forest, elevation 750 ft., 29.IV.2000, J. R. Shevock 19300 (UC 1739292) California: Kern Co., 35° 29' 13.2"N -118° 32' 27.5994"W, Sequoia National Park, canyon live oak woodland, elevation 4100 ft., 1.III.1997, J. R. Shevock 14875 (UC 1754029) California: Tulare Co., 36° 33' 28.7994"N -118° 32' 27.5994"W, Sequoia National Park: canyon live oak woodland, elevation 4800 ft., 26.IV.1997, J. R. Shevock 15225 (UC 1783172) California: Shasta Co., 41° 10' 1.2" N -122° 19' 26.3994" W, Shasta National Forest: Castle Craig Wilderness in open mixed conifer forest with Quercus chrysolepis, elevation 3800 ft., 23.IV.2006, J. R. Shevock 27938 (UC 1798373) California: Glenn Co., 39° 33' 57.6" N -122° 39' 28.8" W, Mendocino National Forest: black oak- foothill woodland, elevation 1800 ft., 2.VII.1997, J. R. Shevock 15803 (UC 1730605) California Madera Co., 37° 17' 42"N -119° 21' 53.9994"W, Sierra National Forest: mixed conifer-hardwood woodland elevation 4593 ft., 29.VII.2000, D. H. Norris 100036 (UC 1759949) California: Fresno Co., 36° 44' 27.5994"N -119° 2' 27.5994"W, Sequoia National Forest: canyon live oak-transition forest, elevation 4600 ft., 29.V.1997, J. R. Shevock 15612 (UC 1730807) California: San Benito Co., 36° 29' 52.0794"N -121° 12' 19.4394"W, Pinnacles National Monument: volcanic rocks along intermittent streamlet with Adenostoma fasciculatum and Juniperus, elevation 1550 ft., 12.II.2005, Shevock 26313 (UC 1798845) California: Alameda Co., 83.37° 52'19"N -122°14'17"W. Strawberry Creek Ecological Area, U.C. Botanic Garden, common on trunks of Quercus agrifolia, elevation 400 ft., 16.III.2008, J. R. Shevock 31566 (CAS 482680) California: Humboldt Co., 40° 38' 59.9994"N -123° 35' 59.9994"W, 22.XI.1994, D. H. Norris 83851 (UC 1774498) California: Lake Co., 39° 27' 45"N -122° 44' 27"W. Mixed conifer forest, on trunk and branches of Quercus garryana breweri, elevation 5100 ft, 3.VII.1997, J. R. Shevock 15858 (UC 1730827)

California: Mariposa Co., 37° 42'56"N -119° 39'50:W. Mixed conifer forest of Pseudotsuga menziesii, Abies concolor, Pinus ponderosa with Cornus & Acer, elevation 3854 ft. 7.X.2006, J. R. Shevock 29092 (CAS 482690) California: Tulare Co., 36° 27'45"N -118 44'15"W. Mixed conifer forest, elevation 5680 ft., 25.III.1996, J. R. Shevock 13180 (CAS 482699) California: Trinity Co., 40° 21'21"N -123° 9'33"W. Mixed conifer forest, On bark of Garry oak trunk 2-3 m above the ground elevation 5249 ft., 16.VI.2001, L. P. Janeway 7346 (UC 1932873) California: Napa Co., 38° 46'49" N -122° 22'33"W. Serpentine chaparral, on bark of toyon and scrub oak, elevation 1850 ft. 12.V.2002, L. P. Janeway 7545 (UC 1932923) California: Tehama Co., 40° 18' 57.6" N -122° 52' 58.7994" W, Yolla Bolla Mountains, conifer forest with canyon live oak and black oaks, elevation 3000 ft., 26.V.2000, J. R. Shevock 19375 (UC 1739811) California: Tulare Co., 36° 8' 59.9994"N -118° 30' 57.6"W, Sequoia National Forest: Pinus jeffreyi, Quercus kelloggii, Calocedrus and Pseudotsuga forest, elevation 5249 ft., 7.I.1991, D. H. Norris 76218 (UC 1723875) California: Lake Co., 38° 59' 41.9994" N -122° 45' 54" W, North Coast Ranges, Clear Lake Region: in shaded transition forest dominated by Douglas fir, bay laurel, and madrone, elevation 2400 ft., 1.IV.2001, J. R. Shevock 20601 (UC 1741428) California: San Benito Co., 36° 47' 56.3994"N -121° 29' 52.8"W, San Juan Canyon: Quercus wislizeni forest, elevation 656 ft., 20.II.2002, D. H. Norris 103529 (UC 1767845) California: Tulare Co., 36° 8' 27.5994"N -118° 35' 59.9994"W, Sequoia National Forest: Giant Sequoia-mixed conifer forest, elevation 5000 ft., 17.III.1996, J. R. Shevock 13124 (UC 1754019) California: Humboldt Co. In mixed conifer forest on leafless Salix sp., 12.V.1983, C. J. Thomas 37 (SFV 19758) California: Shasta Co., 40° 53’N -121° 44’W. Along Hwy 299 on North side of Haynes flat, on moist diffusely lit bark of Quercus in Pinus ponderosa and Quercus forest, elevation 3280 ft. , 22.IV.1983, D. H. Norris 68539 (SFV 20956) California: Tulare Co., 36° 30' 44.6394"N -119° 47' 56.4"W. Sequoia National Park: south of Potwisha camp near footbridge across Middle Fork of Kaweah River, amid oak forest on live oak, elevation 2000 ft., 11.I.2009, P. Wilson 5232 (SFV 21504) California: Tulare Co., 36° 34' 33.204" N -118° 47' 5.9994" W. Sequoia National Park: Crystal Cave Road trail bridge, in mixed conifer forest with mountain Dogwood, Incense cedar, Jeffrey Pine, Black Oak, and White Fir, on bark of Black Oak, elevation 5235 ft., 23.V.2009, L. A. Coleman 810 (SFV 22756) California: Tulare Co., 36° 33' 33.7674"N -118° 47' 2.724"W. Sequoia National Park: at beginning of Crystal Cave Road, in mixed conifer forest with Abies concolor, Cornus nuttallii, and Calocedrus decurrens, on streambank with fallen log, elevation 5411 ft., 13.VI.2011, L. A. Coleman 161 (SFV 22694)

Orthotrichum pellucidum U.S.A. California: Tulare Co., 36°08'00.0"N 118°37'00.0"W. Sequoia National Forest: On moist, diffusely lit limestone boulder in Calocedrus decurrens, Quercus chrysolepis and Pinus ponderosa woodland, elevation 4757 ft., 27.III.1996, D. H. Norris 87688 (UC1774546)

California: Inyo Co., 37°18’47.7”N -118°10’23.1”W. White mountains: ~1 mi. walk near Piñon picnic area, under an overhanging ledge of slate, elevation 7211 ft., 20.VI.2004, P. Wilson 3972 (SFV 20585)

Orthotrichum pilosissimum U.S.A. Nevada: NYE Co., 38° 47’36”N -116° 51’29”W. Toquima Range, Toiyabe National Forest: Pine creek Campground, in riparian woodland with quaking aspen and black cottonwood within a pinyon-juniper woodland, elevation 7700 ft. 31.V.2002, J. R. Shevock 22397 (CAS 1040002)

Orthotrichum praemorsum U.S.A. Nevada: Carson City Co., 39° 09’48”N -119° 55’50”W. Northern Sierra Nevada, Lake Tahoe Basin: in open Jeffrey pine, incense cedar and white fir forest, elevation 6385 ft. 20.IV.2002, J. R. Shevock 22039 (CAS1040112) Nevada: Carson City Co., 39° 06’52”N -119° 48’36”W. Carson Range: Clear Creek Canyon, on hillside of granitic rock outcrops in a Jeffrey pine forest with Eriogonum, Huecher and Purshia, on rock, elevation 5200 ft., 5.IV.2002, J. R. Shevock 21963 (CAS1040076)

Orthotrichum pulchellum U.S.A. California: San Luis Obispo Co., 35° 16’N -120° 53’W. Montana de Oro State Park: in Eucalyptus forest on moist diffusely lit twigs of Baccharis, elevation 164 ft., 25.XII.1982, D. H. Norris (UC California: Humboldt Co., 41°05’N -124°09’W. In Alnus, Pseudotsuga and picea sitchensis forest, on bark of Sambucus, elevation 0-164 ft., 9.V.1976, Norris 68200 (SFV 21218) California: Humboldt Co., 41°08'00.0"N 124°09'00.0"W. Patrick Points state Park: On moist, diffusely lit bark of Sambucus in second-growth Alnus forest on sea cliff, elevation n/a, 23.IX.1989, D. H. Norris 74989 (UC1673879) California: Humboldt Co., 41°05’N -124°09’W. College Cove: ½ mile north of Trinidad, on moist diffusely lit bark of Sambucus in Alnus, Pseudotsuga, and Picea sitchensis forest, elevation 164 ft., 9.V.1976, D. H. Norris 47747 (SFV21201) California: Humboldt Co. Soma Peninsula, near Coast Guard Station near north spit, on Salix on sand dunes, elevation 50 ft., 25.V.1984, P. Wilson 166 (SFV 197659) California: Del Norte Co., 41°34’N -124°06’W. Redwood National Park: along coast trail between Falsa Klamath Cove and Requa, elevation 328 ft., 13.IV.1984, D. H. Norris 70366 (SFV 20957)

Orthotrichum pylaisii U.S.A. California: Tulare Co., 36° 39' 1.1154"N -118° 49' 50.124"W. Sequoia National Park: Dorst Creek Trail, in mixed conifer forest with Giant Sequoia, White fir, and Chinquapin, elevation 6596 ft., 28.VI.2010, L. A. Coleman 593b (SFV 22817) California: Tulare Co., 36° 34' 2.2434"N -118° 46' 38.4594"W. Sequoia National Park: Rock Trail at Beetle Rock, on large shaded boulder in mixed conifer forest with Abies concolor, Pinus lambertiana, Pinus jeffreyii, Manzanita, Linanthus and Mimulus, elevation 6331 ft., 14.VI.2011, L. A. Coleman 888 (SFV 22816) California: Tulare Co., 36° 39' 3.0594" N -118° 49' 41.6994"W. Sequoia National Park: Dorst Creek Trail, on large rock outcrop in mixed conifer forest with Giant Sequoia and Chinquapin, elevation 6589 ft., 28.VI.2010, L. A. Coleman 585 (SFV 22815)

California: Tulare Co., 36° 36' 32.256"N -118° 44' 2.0754"W. Sequoia National Park: Cahoon Meadow Trail, on granite outcrop, on slope, amid conifer forest with Red Fir, White Fir, and Jeffrey Pine, elevation 7207 ft., 15.VIII.2009, L. A. Coleman 486 (SFV 23086)

Orthotrichum rivulare U.S.A. California: Del Norte Co., 1°48'00.0"N 124°05'00.0"W. Jedediah smith State Park: On moist, diffusely lit bark of Alnus in old growth Sequoia sempervirens forest and on riverine corridor of Umbellularia and Acer, elevation 164 ft., 13.VI.1983, D. H. Norris 68926 (UC1650328) California: Mendocino Co., 39°05'00.0"N 123°28'60.0"W. Hendy Woods State Park: On moist, diffusely lit bark of Acer macrophyllum in riverine forest with Sequoia and Quercus, elevation 328 ft., 27.VI.1979, D. H. Norris 53107 (UC1650324) California: Lake CO., 39°02’N 123°49’W. Corinthian Bay County Park: on moist diffusely lit bark of Salix in shore corridor of Salix and Populus on south shore of Clear Lake, elevation 1312 ft., 21.XII.1984, D. H. Norris 71776 (SFV 21237) California: Mendocino Co. on rock along Hwy 101 between Laytonville and Willitts, just upstream from bridge 10-98, 3.III.1984, P. Wilson 96 (SFV 19768) California: Shasta Co., 40°37’N -122°35’W. Whiskeytown National Recreation Area: At Brandy Creek Picnic Area, on moist diffusely lit boulder in stream corridor of very large Quercus chrysolepis, Pinus ponderosa, and Pseudotsuga menziesii along stream in area of old chaparral, elevation ~1148 ft., 28.II.1988, D. H. Norris 73623 (SFV 20960)

Orthotrichum rupestre U.S.A. California: Tulare Co., 36° 39' 31.968"N -118° 49' 42.276"W. Sequoia National Park: on Alder trunk at a river on small trail past Crystal Cave, in Quercus chrysolepis, Abies concolor, Athyrium, Acer macrophyllum, Taxus, and Alder forest, elevation 4413 ft., 14.VI.2012, L. A. Coleman 1651 (SFV 22745) California: Tulare Co., 36° 34' 31.368"N -118° 47' 5.9994"W. Sequoia National Park: on large granodiorite wall with seasonal seepage now dry, in mixed conifer forest, along crystal Cave Road, right before bridge, elevation 5243 ft., 23.V.2009, L. A. Coleman 133 (SFV 22752) California: Tulare Co., 36° 34' 31.368" N -118° 47' 5.9994" W. Sequoia National Park: on large granodiorite wall with seasonal seepage now dry, in mixed conifer forest, along crystal Cave Road, right before bridge, elevation 5243 ft., 23.V.2009, L. A. Coleman 132 (SFV 22812) California: Kern Co., 35° 3' 36"N -118° 29' 23.9994"W. Tehachapi Mountain Park: on hard sandstone boulder on north facing slope south of campground, elevation 6725 ft., 15.III.2014, N. Sarkissian 21 (SFV 23108)

Orthotrichum shawii U.S.A. California: Lake Co., 39° 27’46” N -122° 45’18” W. 4850 feet. On metavolcanic rock outcrop. 10.VII.2012, D. Toren 9891 (CAS1181013); Contra Costa Co., 37°52’55” N -121 54’49” W. Oak woodland, elevation 3800 ft. On large metamorphic boulders and rock outcrops. 28.II.2004, J. R. Shevock 24632 (CAS1113091)

Orthotrichum speciosum U.S.A. California: Del Norte Co., 41° 46' 58.8"N -123° 43' 58.8"W, Bear Basin Creek, in Abies and Pseudotsuga forest, elevation 4101 ft., 12.VI.1983, D. H. Norris 68877 (UC 1723786)

Orthotrichum texanum U.S.A. Montana: Flathead Co., Shepard Memorial Spring, Bad Rock Canyon: On shaded argillite bluff, elevation n/a. 24.VII.1966, F. J. Herman 20411 (CAS 49527)

Orthotrichum underwoodii Information unavailable due to protected/threatened status: K. Kellman 1183 (CAS 1073366) Information unavailable due to protected/threatened status: D. Toren 9511 (CAS 1048109)