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Transcript
1
SPORE-BEARING VASCULAR PLANTS:
IDENTIFICATION KEYS FOR USE IN THE INTERMOUNTAIN REGION
OF WESTERN NORTH AMERICA.
Mary E. Barkworth
Intermountain Herbarium, Department of Biology
Utah State University
With support from the Utah Agricultural Experiment Station
2
PREFACE
This treatment is a derivative work. I have prepared it for use in teaching plant taxonomy at the Logan
Campus of Utah State University. It makes available a treatment of the spore-bearing plants of the
Intermountain Region that reflects current taxonomic opinion and direct users to works that provide
detailed descriptions and illustrations of the species.
Geographic scope: For the purposes of this treatment, the Intermountain Region is interpreted to
include the region treated by Cronquist et al. (1973) plus the southern tip of Nevada. Decisions as to
which species to include are based on three primary sources, volume 1 of the Intermountain Flora
(Cronquist et al. 1972), volume 2 of the Flora of North America north of Mexico (Flora of North
America Editorial Committee 1993), and edition 4 of A Utah Flora (Welsh et al. 2008). In addition, I
have included several species that are grown in the greenhouse of the Logan Campus of Utah State
University so as to make it more useful in teaching plant taxonomy.
Information sources: The family treatments were initially based on Smith et al. (2006) but they have
been modified to reflect the treatment in Christenhulz et al. (2011) which reflects subsequent
research. The generic treatments are based primarily on those in the Flora of North America.
References following species names are to the relevant pages in volume 1 of the Intermountain Flora
(IMF), volume 2 of the Flora of North America (FNA), or A Utah Flora (UF). The Intermountain Flora
provides descriptions and line drawings for all the species recognized; FNA provides descriptions for all
species and line drawings for representative species, some of which are from IMR; UF provides
descriptions. In addition, I have consulted Wikipedia, particularly for information about the greenhouse
species and various other floristic treatments for information about species that are grown in the
greenhouse. The families are treated in the sequence recommended by Christenhulz et al. (2011). The
genera and species are treated alphabetically within each family.
Links: There are three sets of links. Those for the global distribution of a taxon are to the Global
Biodiversity Information Facility. To see the map, it is necessary first to scroll down the page that
comes up and accept the terms, then to scroll down the next page that comes up because the map is
some distance down. Links to the Encyclopedia of Life should take you directly to the appropriate page
on that web site. Links for the Intermountain Biota pages will take you directly to the relevant pages in
the Intermountain Biota site. For further information about the genera, there are links to Wikipedia. I
have found it useful. All these projects are works in progress. Some of the linked pages will have almost
no information; others will be a rich resource. With time, more will be rich resources. Of course,
increased resources would increase the rate at which herbaria and taxonomists are able to share their
knowledge.
Still to be completed: There are still items on the to-do list. One is completion of a more complete list
of works consulted. For now, I need to create keys to other plant groups.
Preparation of this treatment would have been impossible were it not for the enormous amount of
detailed field, herbarium, and laboratory study that has been conducted by others as reflected in the
preliminary listing of references plus the information that people have contributed to Wikipedia.
Having said which, I am responsible for all the errors and misinterpretations that are present. Please let
me know about them by emailing mary.barkworth[at]usu.edu.
Mary E. Barkworth
February 20, 2011
3
SPORE-BEARING VASCULAR PLANTS
There are two divisions currently recognized among the extant spore-bearing plants, the
Lycopodiopsida or lycophytes, a group that used to dominate the vegetation, back in the Devonian, and
and the Equisetopsida or monilophytes, a recently recognized group that encompasses all today’s ferns
and the horsetails. The formal name of the monilophytes is Equisetopsida because that is the first
name published at the rank of class that includes one of its members, but it has been differently
interpreted over the past century. The informal name “monilophytes” has come into use very recently
and is less likely to be misinterpreted. Ranks are relative, not a measure of size.
1a. Leaves 1-veined, green; sporangia not lobed, solitary on the adaxial or inner side of leaf-like
sporophylls; plants up to 20 cm tall ............................................................................................ Lycopodiopsida
1b. Leaves usually green and many-veined, someimtes absent or 1-veined and black at least distally;
sporangia usually unlobed and several to many on leaves or specialized sporophores, 3-lobed if
solitary; plants usually more than 20 cm tall ................................................................................. Equisetopsida
LYCOPODIOPSIDA BARTL
Species homosporous or heterosporous. Plants perennial, herbaceous. Gametophytic and sporophytic
phases free-living, sporophyte dominant. Sporophyte: Stems unbranched or with dichotomous or
monopodial branching; leaves microphyllous, with only one vein; sporophylls sometimes aggregated
into distinct strobili, sometimes morphologically distinct from leaves. Sporangia adaxial on or axillary to
the sporophylls, reniform, dehiscing transversely.
The Lycopodiophyta appeared shortly after the first tracheophytes. They reached their peak in the
Carboniferous, around 350–300 MYA, at which time they included many arborescent species. Today
they include only three families and 1200 species, most of which are tropical and either creeping or
pendant from other vegetation. .
1a. Leaves and sporophylls 2–100 cm; stems erect, short, not evident, unbranched; sporangia
embedded in the leaves...................................................................................................................... Isoëtaceae
1b. Leaves and sporophylls 0.2–2 cm long; stems trailing, elongate, trailing, branched; sporangia not
embedded in the leaves..................................................................................................................................... 2
2a. Plants usually 5–20 cm tall, homosporous; strobili not differentiated or dense and cylindrical.........................
..................................................................................................................................................... Lycopodiaceae
2b. Plants usually less than 3 cm tall, heterosporous; strobili often weakly differentiated, 4-sided or
open and cylindrical .................................................................................................................... Selaginellaceae
Lycopodiaceae P. Beauv. ex Mirb. [Wikipedia]
Plants 5–20 cm tall, homosporous. Stems elongate, horizontal, anchored by rhizophores, branching
dichotomously or monopodially. Leaves 1.5–20 mm, flat, thin, not ligulate, margins spinulose.
Sporophylls like vegetative leaves or non-photosynthetic and smaller, sometimes forming distinct,
cylindrical, terminal strobili. Sporangia solitary, adaxial or axillary.
There are 350-400 species of Lycopodiaceae in the world. They are most common in tropical montane
and alpine habitats.
Lycopodium L. [Wikipedia]
There is only one species that may be present in the Intermountain Region.
4
Lycopodium annotinum L. [Bristly clubmoss; FNA 2:25, 26] [Global distribution; Encyclopedia of Life;
Intermountain Biota]
In swampy or moist coniferous woods and exposed grassy or rocky sites. A northern hemisphere
species. Presence in IMR needs verification. FNA (2:26) shows it as being present in southwestern
Idaho and northeastern Utah but it was not included by either Cronquist et al. (1973) or Welsh et al.
(2008). Following FNA, no varieties are recognized.
Isoëtaceae Dumort.
Stems condensed, 2-lobed. Leaves 2–50 cm long, more or less cylindrical, narrowing distally, bases
thick. Sporangia adaxial, embedded in the base of the leaves, partly to wholly covered from above by a
thin membrane, the velum; inner leaves with megasporangia, outer leaves with microsporangia.
Isoëtaceae tend to grow at the edges of seasonal fresh water pools such as Dry Lake in Cache Valley,
Utah and the vernal pools of California.
Isoëtes L. [Wikipedia]
1a. Megaspore surface spiny ............................................................................................................... I. echinospora
1b. Megaspores surface cristate, reticulate, rugulate, tuberculate, or smooth ...................................................... 2
2a. Plants terrestrial or becoming so, growing in seasonally saturated soils such as temporary
streams and vernal pools; leaf bases pale to brown or lustrous black. ............................................................. 3
3a. Velum covering less than 3/4 of the sporangium; megaspore girdle obscure; massed microspores
gray ............................................................................................................................................... I. melanopoda
3b. Velum covering less than 1/2 of the sporangium; megaspore girdle smooth; massed microspores
brown .................................................................................................................................................... I. howellii
2b. Plants submerged or emergent aquatics or persistent lakes or pools; leaf bases pale to pale
brown ................................................................................................................................................................. 4
4a. Megaspores averaging less than 0.5 mm diameter ........................................................................... I. bolanderi
4b. Megaspores averaging 0..5–0.75 mm in diameter ............................................................................................ 5
5a. Leaves gradually tapering to the tip; sporangium wall not pigmented; massed microspores brown
....................................................................................................................................................... I. occidentalis
5b. Leaves abruptly tapering to the tip; sporangium walls re or less brown-streaked; massed
microspores gray ................................................................................................................................. I. lacustris
Isoëtes bolanderi Engelm. [Bolander’s quillwort; IMF 1:182–183; FNA 2: 71, 73; UF 1]. [Global
distribution, Encyclopedia of Life, Intermountain Biota]
Subalpine to alpine lakes and pools.
Isoëtes echinospora Durieu Spiny-spored quillwort; [IMF 1:183, as I. echinospora; FNA 2: 65, 68; UF 1].
[Global distribution; Encyclopedia of Life; Intermountain Biota]
In oligotrophic, non-calcareous lakes and ponds. Primarily a northern species; known from the Uinta
Mountains and, possibly the northwestern IMR. North American plants belong to I. echinospora subsp.
muricata (Durieu) Á. Löve & D. Löve. They differ from European plants, which belong to I echinospora
subsp. echinospora, in having stomates.
Isoëtes howellii Engelm. Western quillwort [IMF 1:182–183; FNA 2: 71, 74; UF 1]. [Global distribution;
Encyclopedia of Life; Intermountain Biota]
Wet depressions and lake margins in the mountains. IMF gives reasons for excluding Utah plants from
I. melanopoda.
5
Isoëtes lacustris L. [FNA 1:184; UF 1]. [Global distribution; Encyclopedia of Life; Intermountain Biota]
In lakes at middle elevations in the mountains, often submerged in deep water. Very similar to, and
sometimes included in, I. occidentalis..
Isoëtes melanopoda Gay & Durieu [Black-footed quillwort; IMF includes in I. howelli; FNA 2:73; UF 1
suggests the FNA report may be based on a misidentification]. [Global distribution; Encyclopedia of
Life; Intermountain Biota]
In non-calcareous soil, primarily east of the Rocky Mountains but, according to FNA, known from the
Uinta Mountains. See comments under I. howellii.
Isoëtes occidentalis L.F. Henders. [Western quillwort; FNA 2:74, 71, IMF 6: 24, as I. lacustris L.; UF 2
suggests that it should be included in I. lacutris] [Global distribution; Encyclopedia of Life;
Intermountain Biota]
In lakes at middle elevations in the mountains, often submerged in deep water. Very similar to I.
lacustris and included in it by IMF
Selaginellaceae Willk.
Plants usually shorter than 2 cm, heterosporous. Stems elongate, horizontal, often anchored by
rhizophores, branching dichotomously or monopodially. Leaves 2–10 mm long, flat, thin, ligulate,
margins dentate to spinulose, at least in part. Sporophylls often more densely packed than the leaves
and forming distinct strobili; strobili usually terminal and 4-sided, if cylindrical the strobili open;
sporangia adaxial or axillary, either or both megasporangia and microsporangia present within a
strobilus, if both, the megasporangia proximal and the microsporangia distal within each strobilus.
Selaginellaceae tend to grow in wet places. Those of the Intermountain Region are associated with
rocky habitats.
Selaginella P. Beauv. [Wikipedia]
There is only one genus¸ Selaginella, in the family. For the description, see the family description.
1a. Leaves on aerial stems dimorphic, arranged in 4 ranks; axillary leaves present at branching points;
rhizophores present; not native in IMR ........................................................................................ S. lepidophylla
1b. Leaves on aerial stems monomorphic, not in distinct ranksm without axillary leaves at branching
points; rhizophores absent ................................................................................................................................ 2
2a. Sporophylls loosely packed, strobili cylindrical; rhizophores not present; leaf margins shortly
spiny ..............................................................................................................................................S. selaginoides
2b. Sporophylls densely packed; strobili 4-sided; rhizophores developed; leaf margins dentate,
serrate, or ciliate, not spiny ............................................................................................................................... 2
3a. Leaves on main stem differing in color from stem, bases usually rounded, sometimes slightly
decurrent and cuneate ......................................................................................................................... S. mutica
3b. Leaves on main stem similar in color to stem, bases cuneate or oblique, decurrent ....................................... 3
4a. Leaves on upper and lower sides of stem unequal in size, their bases decurrent and oblique ........................ 4
5a. Leaf tips with a bristle 1–2 mm long; sporophyll margins ciliate throughout ....................................... S. densa
5b. Leaf tips with a bristle 0.4–1.3 mm long; sporophyll margins with cilia 0.02–0.15 mm long or
denticulate on lower half, smooth distally .................................................................................. S. underwoodii
4b. Leaves usually all similar in size, if slightly unequal the leaf bases decurrent cuneate ..................................... 5
6a. Lateral branches 1–3-divided; dry stems not readily fragmenting ..................................................... S. watsonii
6b. Lateral branches 1-divided; dry stems readily fragmenting .............................................................................. 6
7a. Leaf tips with a bristle 0.2–0.6 mm long; leaves not in well-defined pseudowhorls ................. S. leucobryoides
6
7b. Leaf tips without a bristle or with bristle to 0.4 mm long; leaves in defined, alternate
pseudowhorls ................................................................................................................................... S. utahensis
Selaginella densa Rydb. [Prairie club-moss, Rocky Mountains spike-moss; IMF 2:178–179, FNA 2: 56;
UF 2] [Global distribution; Encyclopedia of Life; Intermountain Biota]
Rocky, exposed sites, at high elevations, often in the alpine zone.
Selaginella lepidophylla (Hook. & Grev.) Spring [Resurrection plant] [Global distribution; Encyclopedia
of Life; Intermountain Biota]
Plants terrestrial or on rock, forming rosettes. Rhizophores borne on upperside of stems, restricted to
basal part of the rosettes. Leaves thick and stiff.
Selaginella leucobryoides Maxon [Mojave spike-moss; not in IMF; FNA 2:55; not in UF] [Global
distribution; Encyclopedia of Life; Intermountain Biota]
Rock crevices or exposed rocky sites. Often confused with S. utahensis which has an overlapping range.
Selaginella mutica D.C. Eaton ex Underw. [Awnless spike-moss; IMF 1:180, 181; FNA 2: 51; UF 2]
[Global distribution; Encyclopedia of Life; Intermountain Biota]
Crevices and ledges on igneous, limestone, or sandstone rocks, usually below the forest zone. IMR
plants belong to Selaginella mutica D.C. Eaton ex L. Underw. var. mutica.
Selaginella selaginoides (L.) Link [Northern spike-moss, Prickly mountain moss; IMF 1:178, 179; FNA 2:
39, 41; not in UF] [Global distribution; Encyclopedia of Life; Intermountain Biota]
Wet places, often on mossy stream banks.
Selaginella underwoodii Hieron. [Underwood’s spike-moss; IMF 1:180,181; FNA 2: 53; UF 2] [Global
distribution; Encyclopedia of Life; Intermountain Biota]
Moist or shaded cliffs, rocky slopes, and crevices on granite, sandstone, or limestone.
Selaginella utahensis Flowers [Utah spike-moss; IMF 180, 181; FNA 2:55; UF2] [Global distribution;
Encyclopedia of Life; Intermountain Biota]
Dry sandstone crevices or sandy or clay soils. Often confused with S. leucobryoides and having an
overlapping range.
Selaginella watsonii Underw. [Watson’s spike-moss; IMF1: 179–180, 181; FNA 2: 54] [Global
distribution; Encyclopedia of Life; Intermountain Biota]
Exposed or shaded rocky sites or in gravel or sandy soil, sometimes in alpine regions.
EQUISETOPSIDA C. AGARDH [MONILOPHYTES]
Plants herbaceous or arborescent, usually homosporous, aquatic genera often heterosporous. Stems
often subterranean and horizontal, sometimes surficial, sometimes vertical. Leaves usually compound,
large, and with many veins, sometimes small and lacking veins or with 1 vein. Sporangia usually on the
abaxial surface of leaves, sometimes terminal on specialized branches or modified leaves, if axillary to
a reduced leaf, sporangia 3-lobed. Sporangia eusporangiate (with walls 2 or more cell layers thick) or
leptosporangiate (with walls only 1 cell layer thick).
7
Monilophytes encompass, in addition to the plants traditionally considered ferns, the Psilotaceae and
Equisetaceae, two families that used to be called “fern allies”. There are about 300 genera and 9,000
species of monilophytes.
Plants floating aquatics; shorter than 10 mm long ......................................................................... Salviniaceae
Plants rooted in soil or growing on trees ........................................................................................................... 2
Stems vertical, less than 1.5 cm in diameter; leaves scale-like, unlobed, 1-veined or without veins ............... 3
Leaves whorled, connivent; sporangia unlobed, on sporangiophores that are clustered together
into a terminal strobilus; plants native, growing outside ............................................................... Equisetaceae
3b. Leaves alternate, not connivent; sporangia 3-lobed, axillary to short branches, not in strobili; plants
not native, grown in greenhouses ..................................................................................................... Psilotaceae
2b. Stems if present, usually horizontal and subterranean, sometimes horizontal and above ground,
vertical and more than 10 cm in diameter in tree ferns, sometimes essentially non-existent;
leaves conspicuous, with multiple veins, usually lobed or compound .............................................................. 4
4a. Plants tree-like; stems vertical, more than 10 cm in diameter ......................................................... Cibotiaceae
4b. Plants herbaceous; stems, of present, horizontal, usually below ground, usually less than 1 cm in
diameter ............................................................................................................................................................ 5
5a. Plants emergent aquatics, heterosporous; sporangia formed in specialized sporocarps ...................................
........................................................................................................................................................Marsileaceae
5b. Plants terrestrial, homosporous; sporangia borne on leaves, not in sporocarps .............................................. 6
6a. Sporangia on modified leaf blades, about 1 mm in diameter, thick-walled, without an annulus;
leaves not circinate ................................................................................................................... Ophioglossaceae
6b. Sporangia on the lower surface of the leaf blades, less than 0.5 mm in diameter, thin-walled,
usually with an annulus; leaves circinate in bud................................................................................................ 7
7a. Rhizomes and leaves with hairs but not scales; sori marginal, protected by the reflexed leaf
margins; petioles with several vascular bundles .................................................................... Dennstaedtiaceae
7b. Rhizomes and often the petioles with scales, often also with hair; sori variously located,
sometimes protected by the reflexed leaf margins; petioles often with only 1 or 2 vascular
bundles .............................................................................................................................................................. 8
8a. Rhizomes above ground, conspicuously scaly; indusium cup-shaped, opening towards the margin .................
.........................................................................................................................................................Davalliaceae
8b. Rhizomes below ground, not conspicuously scaly; indusial variously shaped, usually round,
reniform, or elongate, not cup-shaped.............................................................................................................. 9
9a. Petioles with a single vascular bundle, always slender, often dark and wiry; indusium formed by
the reflexed leaf margins or indusium absent ................................................................................. .Pteridaceae
9b. Petioles with 2 or more vascular bundles, at least towards the base, sometimes slender and dark;
indusium, if present, not formed by the reflexed leaf margin ........................................................................ 10
10a. Sori elongate, parallel to the midvein of the pinnae; veinlets anastomosing, forming closed ovals
on each side of the pinnae midveins ............................................................................................... Blechnaceae
10b. Sori variously shaped, not parallel to the midvein; all veinlets free ................................................................ 11
11a. Leaves evergreen, merely pinnatifid, pinnae broadly attached to the rachises; indusium absent
...................................................................................................................................................... Polypodiaceae
11b. Leaves annual, 1–4 times pinnate, pinnae narrowly attached to the rachises; indusium present ................. 12
12a. Sori elongate, parallel to primary veins of pinnae; indusial flap-like attached along the vein that
bears the sorus ............................................................................................................................... Aspleniaceae
12b. Sori round; indusia radially symmetric ............................................................................................................ 13
13a. Petiole with three or more veins at the base ........................................................................... Dryopteridaceae
13b. Petiole with two veins at the base ................................................................................................................... 14
14a. Indusia attached at the center of the sori, radially symmetric, composed of a central circular
portion with radiating filaments or ribbons ...................................................................................Woodsiaceae
14b. Indusia absent or attached on the sides of the sori, forming a half-cup or composed of ribbons
that cover the sorus from the sides ................................................................................................................. 15
1a.
1b.
2a.
3a.
8
15a. Indusia absent or small, paralleling the veins, and soon deciduous ................................................. Athyriaceae
15b. Indusia present, cup-shaped, lying across the veins, persistent but sometimes inconspicuous .........................
................................................................................................................................................. Cystopteridaceae
EQUISETIDAE WARM
Equisetaceae Michx. ex DC. Horsetails
Plants perennial, rhizomatous. Aerial stems jointed, with distinct nodes, annual or perennial, with
hollow centers; internodes ridged, ridges continuing into the fused leaves and terminating in the leaf
tips; branches, when present, borne at the nodes and breaking through the base of the subtending
leaves. Sporangia borne on peltate sporangiophores; sporangiophores clustered into distinct, terminal
strobili.
There is only one extant genus of the Equisetaceae, Equisetum.
Equisetum L. Horsetail [Wikipedia]
For the generic description, see the family description.
Of the approximately 15 species in the world, four species and one hybrid grow in the IMR. They grow
in moist to wet places.
1a. Aerial stems persisting one year or less, usually with regular whorls of branches; stomates on
surface, scattered or in bands; strobili with rounded apices............................................................... E. arvense
1b. Aerial stems usually persisting more than a year, usually unbranched; stomates sunk, in single
lines; strobili usually with pointed apices ......................................................................................................... 2.
2a. Spores white, misshapen ....................................................................................................................... E. × ferrisii
2b. Spores green, spherical. ....................................................................................................................................... 3
3a. Leaves at most nodes with a dark band across their base; leaf tips 14 or more, usually deciduous,
articulation line visible ........................................................................................................................E. hyemale
3b. Leaves green or those at the base of the stems with inconspicuous bands; leaf tips 3–32, usually
persistent, sometimes deciduous, articulation line not visible ......................................................................... 4
4a. Leaf tips usually deciduous; strobili with rounded to apiculate but blunt tips; stem ridges flattened
or ± convex .................................................................................................................................... E. laevigatum
4b. Leaf tips usually persistent; strobili with sharply apiculate tips; stem ridges minutely grooved
....................................................................................................................................................... E. variegatum
Equisetum arvense L. Common horsetail [IMF 1: 188; FNA 2: 78, 81; UF3] [Global distribution;
Encyclopedia of Life; Intermountain Biota].
Strobili mature in early spring. In disturbed areas, river banks, fields, and tundra. The most widespread
species of Equisetum in North America.
Equisetum ferrissii Clute [IMF 1: 186; FNA 2: 83; UF 3] [Global distribution; Encyclopedia of Life;
Intermountain Biota]
Strobili maturing in late spring to early summer but spores not shed. Equisetum × ferrissii is a hybrid
between E. hyemale and E. laevigatum that grows in a wide range of open, moist areas such as lake
shores, riverbanks, and roadsides. It is discussed, but not described or illustrated, in IMF.
9
Equisetum hyemale L.. [Scouring rush; [IMF 185–187; FNA 2: 78, 82; UF 3] [Global distribution;
Encyclopedia of Life; Intermountain Biota]
Strobili maturing in summer, old stems sometimes developing branches with strobili in the fall. Moist
roadsides, riverbanks, lake shores, and woodlands. Plants in IMR belong to Equisteum hyemale subsp.
affine (Engelm.) Calder & R.L. Taylor according to FNA; IMF agrees, but with considerable doubt
expressed as to whether or not subsp. affine really is distinct from subsp. hyemale which is supposedly
restricted to Eurasia.
Equisetum laevigatum A. Br. [Smooth scouring rush; IMF 1: 186–187; FNA 2: 82; UF 3] [Global
distribution; Encyclopedia of Life; Intermountain Biota]
Strobili maturing in spring to early summer. Grows in moist prairies, riverbanks, and road banks.
Equisetum variegatum Schleich. ex F. Weber & D. Mohr [Variegated scouring rush; IMF 1: 186–188;
FNA 2: 83; UF 3] [Global distribution; Encyclopedia of Life; Intermountain Biota]
Strobili maturing in late summer or overwintering and shedding spores in spring. Grows on lake shores,
river banks, and in ditches and woods. Plants from the IMR belong to E. variegatum Schleich. ex F.
Weber & D. Mohr subsp. variegatum.
OPHIOGLOSSIDAE KLINGE
Ophioglossaceae Martinov [Wikipedia]
Vernation nodding, not circinate. Rhizomes and petioles fleshy; fertile leaves each with a single
sporophore arising along or at the base of the trophospore (photosynthetic portion of leaf ) stalk or at
the base of the trophophore blade. Sporangia large, with walls 2 cells thick, lacking an annulus.
There are four genera in the Ophioglossaceae. Only one, Botrychium, is present in the IMR.
Botrychium Sw. [Wikipedia]
Roots 0.5–2 mm thick. Stems upright, forming caudices up to 5 mm thick. Trophophores usually
pinnately or ternately-pinnately compound or dissected, sometimes simple; veins forked and free.
Sporophores usually 1 per leaf, 1–3-pinnate, long-stalked. Sporangia borne in 2 rows on pinnate
sporophore branches.
The distribution of Botrychium is rather poorly known because the plants are inconspicuous and grow
early in the season. There is disagreement between the FNA account of the genus and that in IMF and
UF as to how many species are present in the IMR. This may reflect, at least in part, the extensive
searches for Botrychium in the IMR that took place in the late 1990s. This treatment follows the
Wagners’ FNA treatment but indicates where the disagreements lie in order to encourage further
research.
1a. Leaf blades deltate, usually 5–25 cm, usually sterile, sporophores absent or misshapen; plants
usually over 12 cm tall ....................................................................................................................................... 2
2a. Trophophore blade thin; leaf sheaths open; sporophores, when present, arising from the base of
the trophophore blade high on the common stalk; leaves absent during winter ........................ B. virginianum
2b. Trophophore blade thick to leathery; leaf sheaths closed; sporophores, whenpresent arising near
the ground from the basal portion of the common stalk .............................................................. B. multifidum
10
1b. Leaf blades oblong to linear, usually 2–4 cm long, all fertile, sporophores always present; plants
to 15 cm tall, usually shorter than 10 cm .......................................................................................................... 3
3a. Trophophores present or replaced by sporophore, if present, venation of basal pinnae or
segments pinnate; midrib present; basal pinnae or segments oblanceolate to linear, lanceolate,
or ovate .............................................................................................................................................................. 4
4a. Leaves with 2 sporophores and no trophophore ..........................................................................B. paradoxum
4b. Leaves with 1 sporophore and 1 trophophore .................................................................................................. 5
5a. Trophophore blades deltate; sporophores divided proximally into several equally long branches ...................
..................................................................................................................................................... B. lanceolatum
5b. Tropophore blades oblong-linear or nearly deltate; sporophores with single midrib or 1 dominant
midrib and 2 smaller ribs. .................................................................................................................................. 6
6a. Large trophophore blades nearly deltate, basal pinna pair elongate; pinnae distal to basal pair
close together or overlapping; segments and lobes with rounded tips ........................................ B. hesperium
6b. Large trophophore blades oblong-deltate; basal pinnae not elongate; apices truncate to
somewhat acute ..................................................................................................................................................
..........................................................................................................................................................B. pinnatum
3b. Trophophores present; basal pinnae or segments with venation like ribs of fan; midrib absent;
basal pinnae fan-shaped to spatulate................................................................................................................ 7
7a. Trophophore blades ovate to deltate .................................................................................................. B. simplex
7b. Trophophore blades oblong to oblong-lanceolate ............................................................................................ 8
8a. Basal pinnae broadly fan-shaped ....................................................................................................................... 9
9a. Plants herbaceous; trophophores usually smaller than 4 x 1.5 cm; pinnae 2–5 pairs, well
separated; margins usually crenate to dentate; sporophores 1.3–3 times length of the
trophophores .................................................................................................................................B. crenulatum
9b. Plants fleshy; trophophores on most plants larger than 5 x 2 cm; pinnae 4–9 pairs, close together,
sometimes overlapping; margins usually entire to undulate, rarely dentate; sporophores 0.9–2
times length of trophophore ................................................................................................................ B. lunaria
8b. Basal pinnae narrowly fan-shaped, cuneate, lanceolate, or linear ................................................................. 10
10a. Pinnae strongly ascending; margins conspicuously dentate-lacerate ............................................. B. ascendens
10b. Pinnae spreading or only moderately ascending; outer margins usually entire to crenate, rarely
dentate......................................................................................................................................... B. minganense
Botrychium ascendens W.H. Wagner [Upswept Moonwort; not in IMF; FNA 2: 95; not in UF] [Global
distribution; Encyclopedia of Life; Intermountain Biota]
Leaves appearing late spring to midsummer; grassy fields. According to FNA (2: 95), grows in southern
Nevada, an area not included in IMF or UF.
Botrychium crenulatum W.H. Wagner [Dainty Moonwort; not in IMF; FNA 2: 95; UF includes in B.
lunaria] [Global distribution; Encyclopedia of Life; Intermountain Biota]
Leaves appearing in mid to late spring, dying in late summer; local in marshy and springy areas. FNA 2:
95 shows the species as widespread in the IMR.
Botrychium hesperium (Maxon & R.T. Clausen) W.H. Wagner & Lellinger [Western Moonwort; FNA 2:
96; not in UF] [Global distribution; Encyclopedia of Life; Intermountain Biota]
Leaves appear in mid-spring, die in late fall; grassy mountain slopes, snow fields, ditches with willows,
and sand dunes. IMF does not mention the species as such but states that B. matricariifolium (Döll) A.
Braun ex W.D.J. Koch, in which B. hesperium is sometimes included as a subspecies, is reputedly in the
IMR. Neither B. hesperium nor B. matricariifolium is mentioned in UF.
11
Botrychium lanceolatum (S.G. Gmel.) Ångström [Triangle Moonwort; IMF 1: 191–192; FNA 2: 96–97;
UF 4] [Global distribution; Encyclopedia of Life; Intermountain Biota]
Leaves usually drying in midsummer; open fields. Plants from the IMR belong to Botrychium
lanceolatum (S.G. Gmel.) Ångström subsp. lanceolatum.
Botrychium lunaria (L.) Sw. [Common Moonwort; IMF 1: 190, includes B. minganense; FNA 2: 97; UF
includes B. mingananese and D. crenulatum] [Global distribution; Encyclopedia of Life; Intermountain
Biota]
Leaves appearing in spring, dying in late summer. Moist to wet places at middle to high elevations in
the mountains.
Botrychium minganense Vict. Mingan [Mingan Moonwor; FNA 2: 98; IMF and UF include in B. lunaria]
[Global distribution; Encyclopedia of Life; Intermountain Biota]
Leaves appearing in spring through summer; widely scattered.
Botrychium multifidum (S.G. Gmel.) Rupr. [Leather Grape-fern; IMF 1:189 says not in IMR; FNA 2: 92,
93; UF 5] [Global distribution; Encyclopedia of Life; Intermountain Biota]
Leaves remain green overwinter; mainly in fields. FNA 2: 93 shows it as widespread in the IMR.
Botrychium paradoxum W.H. Wagner [Paradox Moonwort; FNA 2: 99; UF includes in B. lanceolatum]
[Global distribution; Encyclopedia of Life; Intermountain Biota]
Plants usually under other vegetation, in snowfields and secondary growth pastures. Not mentioned in
IMF; one record from south-central Utah in FNA 2: 99.
Botrychium pinnatum H. St. John [Northwestern Moonwort; IMF 1: 191–192 includes in B. boreale;
FNA 2: 100; not mentioned in UF] [Global distribution; Encyclopedia of Life; Intermountain Biota]
Leaves appearing June to August. Grassy slopes, stream banks, and woods.
Botrychium simplex E. Hitchc. [Least Moonwort; IMF 1: 189–190; FNA 2: 101; UF 5] [Global
distribution; Encyclopedia of Life; Intermountain Biota]
Leaves appearing mid spring to late fall; meadows at middle elevations, not above timberline.
Botrychium virginianum (L.) Sw. [Rattlesnake Fern, Common Grape fern; FNA 2: 86, 91; UF 5 states
that it is not possible to say with certainty that Utah plants belong to this species rather than B.
multifidum] [Global distribution; Encyclopedia of Life; Intermountain Biota]
An early spring species that dies in late summer; shady forests and shrubby second growth. IMF (1:
189) states that it does not grow in the region; FNA 2: 91 shows it as being widespread within the IMR.
Psilotaceae J.W. Griff. & Henfr. [Wikipedia]
Plants perennial, terrestrial, with branched, rhizoid-bearing, subterranean axes. Stems erect, simple or
dichotomously branched; appendages leaf-like or bract-like, alternate. Sporangia fused into synangia
of 2–3 homosporous eusporangia, solitary in axils of appendages.
There are two genera in the Psilotaceae. The one treated here, Psilotum, is native in Florida and often
grown in greenhouses.
Psilotum Sw. [Wikipedia]
12
Stems branched distally, ridged. Appendages 0.7–2.5 mm long; sterile appendages subulate, those
subtending synangia 2-lobed. Synangia 3-lobed, globose.
Psilotum nudum (L.) Beauv. [Global distribution; Encyclopedia of Life; Intermountain Biota]
Native habitat is low to mesic woods, thickets, swamps, hammocks, and rocky slopes. It is often grown
in greenhouses for teaching purposes.
POLYPODIIDAE CRONQUIST, TAHKT. & ZIMMERM.
Marsileaceae Mirb. [Wikipedia]
Plants emergent aquatics, rhizomatous, heterosporous. Stems growing on soil surface or subterranean,
main stems horizontal; roots arising at nodes and along internodes. Leaves with circinate vernation,
distichous, with 0, 2, or 4 terminal pinnae. Sori bisexual, enclosed in a nutlike sporocarp.
Two of the three genera in the family grow in the IMR.
1a. Leaves composed of a petiole and a blade divided into four pinnae ................................................... Marsilea
1b. Leaves about 0.5 mm wide, without a blade ........................................................................................ Pilularia
Marsilea L. [Four-leaf clover fern] [Wikipedia]
Leaves deciduous in temperate regions heteromorphic, floating leaves usually larger than land leaves;
petioles filiform, stiffly erector procumbent on land leaves, lax on floating leaves; blades divided into 4
wedge-shaped pinnae. Sporocarps ovoid, borne on branched or unbranched stalks at the base of the
petioles, usually above ground.
There are 45 species in the world, with two being known from the IMR. Sporocarps are essential for
accurate identification. The English name reflects the appearance of the genus.
1a. Plants producing 1-3 roots between the nodes as well as at the nodes; pinnae glabrous to
sparsely pubescent; stalks of sporocarps often branched; sporocarps usually not toothed; not
native in IMR ................................................................................................................................. M. quadrifolia
1b. Plants not producing roots between the nodes; pinnae; pilose, pubescent or glabrous; sporocarp
stalks unbranched; sporocarps toothed where joined to stalk; native in IMR .................................................. 2
2a. Each sporocarp with a broad, blunt distal tooth up to 0.4 mm long or with no distal tooth ........ M. oligospora
2b. Each sporocarps with an acute distal tooth 0.4–1.2 mm long ............................................................. M. vestita
Marsilea oligospora Goodd. [FNA 2: 333–234; UF says not in Utah] [Global distribution; Encyclopedia
of Life; Intermountain Biota]
Sporocarps produced June to October. Around ponds and marshes and in wet depressions in
sagebrush communities, sometimes on river margins. In IMF, this species is included in M. vestita. As
described in FNA, its distribution and range of variation in all but the characters used in the key fall
within that of M. vestita.
Marsilea quadrifolia L. [Global distribution; Encyclopedia of Life; Intermountain Biota]
Sporocarps produced summer to fall. In mud and shallow water. Native to Eurasia, established in
northeastern North America but probably introduced as a curiosity.
13
Marsilea vestita Hook. & Grev. [Pepperwort; Water-clover; IMF 1: 220–221; FNA 2:333–334; UF 4]
[Global distribution; Encyclopedia of Life; Intermountain Biota].
Sporocarps produced April-October. In ponds and wet depressions along the shores of lakes, streams,
and vernal ponds.
Pilularia L. Pillwort [Wikipedia]
Plants emergent aquatics. Leaves filiform, with a single midvein, without a blade. Sporocarps globose,
subterranean. There is only one species in North America.
Pilularia americana A. Braun [American pillwort; IMF 1:220-221] [Global distribution; Encyclopedia of
Life; Intermountain Biota]
Sporocarps produced spring-fall. In shallow water around ponds, reservoirs, and temporary pools.
Might be present at low elevations in southern Nevada, based on map in FNA.
Salviniaceae Martinov
Plants floating aquatics, heterosporous. Stems filiform. Leaves sessile or nearly so, simple and to 1.5
cnd about 2mm long. Sporocarps containing one unisexual sorus, soft-walled; microsporocarps about
half the size of megasporocarps.
Only one genus, Azolla, is present in the IMR. The other, Salvinia, may be grown in greenhouses.
1a. Leaves to 2 mm long, unequally bilobed, one lobe submersed and forming a float, the other
smaller, emersed, and turning red in fall .................................................................................................. Azolla
1b. Leaves 3-15 mm long, simple, not turning red in fall .............................................................................. Salvinia
Azolla Lam. [Wikipedia]
Plants moss-like, heterosporous. Leaves bilobed, one lobe submersed and functioining as a float, other
lobe emersed, containing a symbiotic blue-green bacterium, Anabaena azollae. Sporocarps, when
present, borne in pairs on the first leaf of a branch.
There is only one species in the IMR.
Azolla mexicana Presl [Waterfern, Mosquitofern; IMF 1: 222; FNA 2: 340–341; UF 13] [Global
distribution; Encyclopedia of Life; Intermountain Biota]
Stagnant or slow flowing ponds and backwaters, often with Lemna from which it is differs in turning
red in the fall.
Salvinia Ség. [Wikipedia]
Plants floating aquatics, heterosporous. Leaves simple, in whorls of , 2 of the leaves entire, floating, 1
descending, rootlike. Sporocarps unisexual, borne in bisexual clusters, sometimes submersed.
Salvinia includes about 10 species, none of which is native to the U.S. Salvinia minima used to be
considered native (see FNA 2:337) but Jacono et al. 1 (2001) argued that was probably introduced
around 1920-1930. The genus is included here because it is frequently grown in college greenhouses
as a demonstration plant. The following key treats three species that have invasive properties, some of
1
Jacono, .C.C., T.R. Davern, and T. D. Center. 2001. The Adventive Status of Salvinia minima and S. molesta in the
Southern United States and the Relateed Distribution of the Weevil Cyrtobagous salvinae. Castanea 66(3):214226. Also:http://fl.biology.usgs.gov/posters/Nonindigenous/Status_of_Salvinia/status_of_salvinia.html
14
which are sold as aquarium plants. The information has been taken from Flora Europaea, Flora
Mesoamericana, FNA, and Wikipedia.
1a. Leaf papillae 2-3 mm long, dividing into 4 branches near the tip, the parts diverging and then
coming together forming a cage-like structure ...................................................................................S. molesta
1b. Leaf papillae 0.2-0.9 mm long, dividing into 3 or 4 branches but the branches remaining separate,
not forming a cage-like structure at their tip .................................................................................................... 2
2a. Floating leaves almost round to elliptic, 5-16 mm long, 4-14 mm wide.............................................. S. minima
2b. Floating leaves elliptic, 10-14 mm long, 6-9 mm wide ......................................................................... S. natans
Salvinia minima Baker [Global distribution; Encyclopedia of Life; Intermountain Biota]
This species is present in the southeastern U.S. where it was thought to be native. It is now considered
to be a South American species.
Salvinia molesta D.S. Mitch. [Global distribution; Encyclopedia of Life; Intermountain Biota]
This species was introduced to the southern U.S. rather recently. It is a serious concern in several
states because ot its ability to blanket suitable water bodies. It has been found in Arizona, in locations
including near the Nevada border. It can be controlled with a weevil, Cyrtobagous salviniae.
Salvinia natans All. [Global distribution; Encyclopedia of Life; Intermountain Biota]
This species is advertised as being a desirable aquarium plant.
Cyatheales A.B. Frank
This is the order of the tree ferns. It includes eight families. The Logan Campus of USU has one species
growing in its greenhouse.
Cibotiaceae Korall
There is only one genus in this family.
Cibotium Kaulf [Wikipedia]
There are about 11 species of Cibotium. They are native to southeast Asia, Hawai‛I and Central to
Tropical America. It is likely that the species in the USU greenhouse is Cibotium glaucum (Sm.) Hook. &
Arn., this being the one most commonly encountered in the horticultural trade. Cibotium forests in
Hawai‛I are suffering from invasion by a similar-appearing Australian species, Cyathea cooperi (F.
Muell.) Domin.
Dennstaedtiaceae Lotsy [Wikipedia]
Plants terrestrial, colonial. Rhizomes bearing jointed hairs, without scales; petioles often with
epipetiolar buds, with several vascular bundles; blades often large, usually 2–3 or more times pinnate
or ternately-pinnate, veins free, forked, or pinnate, rarely anastomosing and then without included
veinlets; sori marginal or submarginal continuous or discrete; indusia linear, cuplike at blades margins
or sori protected by the rolled leaf margin on one side and a delicate, concealed indusium on the other
side.
There are several genera in the family. One, Pteridium, is native in the IMR; Microlepia can be grown
indoors.
15
1a. Sori circular to oval, just inside the pinna margins and terminal on the veins; indusia half-cup
shaped................................................................................................................................................. Microlepia
1b. Sori continuous along the margin, on a marginal connecting vein; reflexed pinna margin forming
false indusium ...................................................................................................................................... Pteridium
Microlepia C. Presl
Rhizomes hairy. Blades pinnate to pinnatifid, ultimate segments usually oblique at the base. Sori near
the margins, terminal on veins. Indusi thin, half-cup shaprd. Annulus of 14-21 cells.
About 50 species, all native to tropics and subtropics of the Eastern Hemisphere..
Microlepia platyphylla (Don) J. Sm. [Global distribution; Encyclopedia of Life; Intermountain Biota]
Rhizome with dark brown hairs. Petioles 70-100 cm long, straw-colored, glabrous, shiny; blade 100-140
cm long, bipinnate. Native to southeast asia.
Pteridium Gled. ex Scop. [Wikipedia]
Plants terrestrial, often covering large areas. Rhizomes horizontal, long-creeping, without scales.
Leaves widely spaced, 50–450 cm long; petioles glabrous or hairy, with numerous vascular bundles;
blades 2–4-pinnate; pinnae pinnately divided. Sori more or less continuous, covered by recurved false
indusium formed by the leaf margin; true indusium inconspicuous, concealed by false indusium.
Pteridium is treated here as in FNA, as having only one species.
Pteridium aquilinum (L.) Kuhn Bracken [IMF 1: 195–196; FNA 2: 202–203; UF 12] [Global distribution;
Encyclopedia of Life; Intermountain Biota]
Dry to moist woods and open areas, in partial to full sun. Pteridium aquilinum is found in temperate
regions throughout the world, often forming extensive colonies. Plants in the IMR belong to P.
aquilinum var. pubescens L. They are distinguished from other varieties in North America by the
abundant, spreading hairs on the underside of the blades.
Pteridaceae E.D.M. Kirchn. [Wikipedia]
Plants terrestrial, epipetric, or epiphytic. Rhizomes long or short, ascending to erect, usually bearing
scales, sometimes only hairs. Leaves monomorphic to dimorphic, usually pinnate, sometimes pedate;
veins free and forming or anastomosing and forming a reticulate pattern without included veinlets; sori
marginal or submarginal and without a true indusium although often protected by the reflexed blade
margin or sporangia along the veins; sporangia with an interrupted vertical annulus.
The Pteridaceae constitute one of the largest fern families, having around 50 genera and 950 species
worldwide. In the IMR, the family is represented by six genera and 23 species.
1a. Leaves with distinct, clearly separate ultimate segments; veins of ultimate segments prominent,
essentially parallel distally; sporangia located at the tips of the reflexed marginal lobes of the
ultimate segments; veins ..................................................................................................................... Adiantum
1b. Leaves with unlobed or the ultimate segments not distinct; veins of ultimate blade segments
indistinct or pinnate; sporangia on leaf surface or marginal ............................................................................. 2
2a. Petioles longitudinally ridged and with 2-3 grooves; sporangia marginal to submarginal, often
covered by the reflexed leaf margin ........................................................................................................... Pteris
2b. Petioles rounded, flattened or with a single longitudinal groove; sporangia scattered along the
veins or submarginal .......................................................................................................................................... 3
16
3a. Leaves strongly dimorphic, the sterile leaves well developed but obviously shorter than the fertile
leaves and with shorter and relatively wider ultimate segments; petioles greenish to yellowish at
least distally, essentially glabrous; mature spores yellowish ..................................................... Cryptogramma
3b. Leaves monomorphic or only weakly dimorphic; petioles usually brown to black or, if paler,
sparsely to densely pubescent; mature spores usually brown to black, rarely yellowish ................................. 4
4a. Blades with white or yellow farina on the abaxial surface; stem scales bicolored, with a dark
central stripe and tan margins; sporangia scattered along the veins; indusia absent ................. Pentagramma
4b. Blades usually without farina on the abaxial surface, if a white farina present, sporangia marginal
and covered by the reflexed margin; stem scales usually uniformly yellow or brown to black,
sometimes bicolored; sporangia marginal or along the veins; indusium or false indusium (reflexed
leaf margin) present .......................................................................................................................................... 5
5a. Ultimate segments of leaves linear to lanceolate; blades glabrous, adaxial surface lustrous; false
indusia strongly differentiated from segment margin .......................................................................... Aspidotis
5b. Ultimate segments of leaves usually narrowly elliptic to rounded, occasionally linear; blades
glabrous, farinose, or sparsely pubescent, adaxial surface usually dull; false indusia, when
present, poorly differentiated from the leaf margin ......................................................................................... 6
6a. Stem scales strongly bicolored or, if uniform in color, ultimate segments of the leaves more than
4 mm wide ............................................................................................................................................... Pellaea
6b. Stems scales uniformly colored or only weakly bicolored; ultimate segments of the leaves usually
less than 4 mm wide .......................................................................................................................................... 7
7a. Blades with whitish farina on abaxial surface or, if glabrous, petioles lustrous and dark; ultimate
leaf segments somewhat cordate at the base .............................................................................. Argyrochosma
7b. Blades sparsely pubescent or glabrous, never fainose; petioles dull, greenish; ultimate segments
usually truncate to cuneate at the base ........................................................................................... Cheilanthes
Adiantum L. Maidenhair fern [Wikipedia]
Plants medium-sized to small, mesophytic. Rhizomes covered with yellow to dark reddish brown to
blackish scales. Leaves deciduous (IMR taxa); petioles slender, dark, shiny, with a single vascular
bundle; blades usually glabrous, 1-several times pinnate or pinnate and dichotomous; ultimate
segments petiolulate, usually wide and often more or less flabellate; veins free; sori on the reflexed
margins; true indusium not present, sori protected by the reflexed tips of the ultimate segments.
There are 150–200 species of Adiantum; only two are known from the IMR. The generic name comes
from Greek, meaning "not wetting", referring to the leave's ability to shed water without becoming
wet [Information from Wikipedia].
1a.
1b.
2a
2b.
Leaf blades 2–4 times as long as wide, longest pinnae with 15-35 pairs of pinnules; basal pinnae A. aleuticum
Leaf blades 1-2 times as wide as long; longest pinnae with about 6 pairs of pinnules .................................... 2
Veins of the ultimate sterile segments terminating in the teeth ........................................... A. capillus-veneris
Veins of the ultimate sterile segments terminating in the sinuses between the teeth .............. A. raddianaum
Adiantum aleuticum (Rupr.) Paris [Western maidenhair fern; IMF 1:127, as A. pedatum Paris; FNA
2:129; not in UF] [Global distribution; Encyclopedia of Life; Intermountain Biota]
Sporulating in summer-fall. Wooded ravines and shaded banks and cliffs.
IMF includes A. aleuticum in A. pedatum Paris. FNA (2: 126) states that the two are reproductively
isolated and morphologically distinct, A. aleuticum having segments at the middle of the penultimate
blade divisions that are being longer in proportion to their width, have more sharply denticulate,
angular lobes and shorter petiolules.
17
Adiantum capillus-veneris L. [Southern maidenhair fern; IMF 1: 196; FNA 2: 125, 127; UF 6]. [Global
distribution; Encyclopedia of Life; Intermountain Biota]
Sporulating spring-summer. Dripping cliffs and seeps, especially on calcareous or basic rocks. Native to
North America and Europe, often cultivated.
Adiantum raddianum C. Presl [Global distribution; Encyclopedia of Life; Intermountain Biota]
An ornamental species, native to Central and South America. It is likely to be moved to another genus
in the near future.
Argyrochosma (J. Sm.) Windham [Wikipedia]
Plants usually on rock. Rhizomes short, usually unbranched; covered with tan to brown, rarely black,
scales. Leaves monomorphic, 3–30 cm tall; petioles brown or black, glabrous apart from a few scales
near the base, with a single vascular bundle; blades 2–6-pinnate, leathery to herbaceous, lower surface
glabrous, sometimes covered with waxy scales; ultimate segments stalked or subsessile, usually less
than 4 mm wide, margins flat or recurved and forming false indusia. Sporangia scattered along veins,
often submarginal, usually intermixed with glands.
Argyrochosma is restricted to the Americas. It includes about 20 species. The IMF includes it in
Notholaena.
1a. Leaves 2–3-pinnate toward the base, not conspicuously waxy............................................................. A. jonesii
1b. Leaves 4–5-pinnate towards the base, conspicuously waxy below .................................................. A. limitanea
Argyrochosma jonesii (Maxon) Windham [Jones’ cloakfern; IMF 1:206–207 as Notholaena jonesii
Maxon; FNA 2: 170, 174; in UF as Pellaea jonesii (Maxon) Morton] [Global distribution; Encyclopedia of
Life; Intermountain Biota]
Rocky crevices in mountains within sagebrush and pinyon-juniper zones.
Argyrochosma limitanea (Maxon) Windham [Waxy cloakfern; IMF 1:207, as Notholaena limitanea
Maxon; FNA 1:173–174; UF as Pellaea limitanea (Maxon) Morton] [Global distribution; Encyclopedia of
Life; Intermountain Biota]
Rock ledges and crevices from valleys to middle elevations. Plants from the IMR belong A. limitanea
(Maxon) Windham subsp. limitanea.
Aspidotis Copeland [Wikipedia]
Plants terrestrial, sometimes on rock. Rhizomes short, densely covered with dark brown scales, these
sometimes with lighter margins. Leaves 8–35 cm monomorphic to somewhat dimorphic; petioles
usually dark reddish brown, grooved adaxially, with a single vascular bundle; blades 3–4(5) pinnate,
glabrous abaxially and adaxially; ultimate segments short-stalked or with narrow decurrent bases,
veins obscure free, unbranched. Sporangia in marginal, submarginal or continuous sori.
Terrestrial, often at the bases of boulders or in rock crevices, Aspidotis has four species. Three grow in
the U.S. and one in Mexico. Only one species grows in the IMR.
Aspidotis densa (Brackenb.) Lellinger [Indian’s dream, Dense lace-fern; IMF 1:199–200; FNA 2:173; UF
6] [Global distribution; Encyclopedia of Life; Intermountain Biota]
Rocky crevices and moist rocky slopes, at upper elevations to timberline.
18
Cheilanthes Sw. [Wikipedia]
Plants usually on rock. Rhizomes short to long, with linear-subulate to ovate- lanceolate, brown to
black or bicolored scales; Leaves 4–60 cm, monomorphic; petioles brown to black or straw-colored,
rounded, flat, or with a groove, hairy, scaly or glabrous, with a single vascular bundle; blades pinnatepinnatifid to 4-pinnate, leathery or somewhat herbaceous, lower surface hairy and/or scaly, rarely
glabrous, margins usually recurved to form false indusia, continuous or only on lobes, veins usually
free. Sporangia submarginal on vein tops or scattered along veins near margins, n intermixed with
farinose-producing glands.
Cheilanthes is the largest xeric-adapted fern genus in the world.
1a. Lower surface of blades without multicellular scales, sometimes hairy, sometimes glabrous;
vernation usually circinate ................................................................................................................................. 2
2a. Ultimate segments of fertile leaves 1–3 mm long, nearly round, beadlike; blades 3-pinnate near
the base; blades not densely airy on the upper surface ............................................................................ C. feei
2a. Ultimate segments of fertile leaves 3–5 mm; blades 2-pinate-pinnatifid near the base; blades
densely hairy on the supper surface ....................................................................................................... C. parryi
1b. Lower surface of blades with multicellular scales over the veins, intermixed with hairs in some
species; vernation not circinate, expanding leaves hooked but not coiled at the tips...................................... 3
3a. Largest scales on lower surface of blades 0.1–0.4 mm wide, linear, inconspicuous ...................... C. gracillima
3b. Largest scales on lower surface of blades 0.4–1.5 mm wide ............................................................................. 4
4a. Scale margins entire, erose, or denticulate, occasionally with 1 or 2 cilia .......................................... C. eatonii
4b. Scale margins ciliate, usually more so near the base......................................................................................... 5
5a. Blade scales lanceolate with truncate or subcordate bases, basal lobes, if present, not
overlapping, margins often ciliate throughout; rhizomes usually long, leaves scattered ................. C. wootonii
5b. Blade scales ovate-lanceolate with deeply cordate bases, basal lobes usually overlapping, distal
margins usually not ciliate; rhizomes short, leaves in dense clusters. .............................................................. 6
6a. Blade scales ciliate on the basal lobes only; ultimate segments glabrous or with a few entire to
weakly ciliate scales, without branched hairs; stem scales usually dark brown or black
throughout, rarely with light margins .................................................................................................. C. covillei
6b. Blade scales conspicuously ciliate over most of the lower 1/2; ultimate segments of with
branched hairs on the lower surface and ciliate scales on the lower surface; stem scales usually
bicolored .......................................................................................................................................... C. intertexta
Cheilanthes covillei Maxon [Coville’s lipfern; IMF 1: 295; FNA 2:161; UF 8] [Global distribution;
Encyclopedia of Life; Intermountain Biota]
Sporulating late spring-fall. Rocky crevices in desert mountains and canyons, from the Larrea to Pinyon
pine zones, usually on igneous rock. In dry weather, the blades curled up so that only the scales are
visible.
Cheilanthes eatonii Baker [Eaton’s lipfern; IMF 1:204; FNA 2: 160 UF 8] [Global distribution;
Encyclopedia of Life; Intermountain Biota]
Sporulating late summer-fall. Rocky slopes and ledges, on a variety of substrates.
Cheilanthes feei Moore [Slender lipfern; IMF 1: 203–204; FNA 2: 168; UF 8] [Global distribution;
Encyclopedia of Life; Intermountain Biota]
Sporulating late-spring-fall. Cliff crevices, usually on limestone or sandstone, usually above the Larrea
zone.
19
Cheilanthes gracillima D.C. Eaton [Lace lipfern; lace fern; IMF 1: 204-–205; FNA 2:158–159; UF 8]
[Global distribution; Encyclopedia of Life; Intermountain Biota]
Sporulating summer-fall. Crevices, cliffs, and rocky slopes, usually on igneous substrates.
Cheilanthes intertexta (Maxon) Maxon [Coastal lipfern; IMF 2:161; not in UF] [Global distribution;
Encyclopedia of Life; Intermountain Biota]
Sporulating late spring-fall. Rocky slopes and ledges, usually on igneous substrates. Cheilanthes
intertexta has sometimes been treated as a subspecies of C. covillei. Preliminary isozyme evidence
supports the suggestion that it is an allotetraploid hybrid between C. gracillima and C. covillei
(Windham and Rabe in FNA 2:161). It is not mentioned in IMF.
Cheilanthes parryi (D.C. Eaton) Domin [Parry’s lipfern; IMF 1: 106, as Notholaena parryi D.C. Eaton;
FNA 2: 133; UF *] [Global distribution; Encyclopedia of Life; Intermountain Biota]
Sporulating late spring-fall. Cliffs, ledges, and crevices in desert mountains, usually in the Larrea zone.
Cheilanthes wootonii Maxon [Wooton’s lipfern; FNA 2: 163; Not treated in IMF which excludes
southern Nevada; not in UF] [Global distribution; Encyclopedia of Life; Intermountain Biota]
Sporulating summer-fall. Rocky slopes and ledges, usually on igneous substrates.
Cryptogramma R. Br. [Wikipedia]
Plants on rock. Rhizomes little to much branched, covered with scales; scales colorless, brownish, or
mixed, with entire margins. Leaves dimorphic, scattered or densely tufted; fertile leaves 5–25 cm;
sterile leaves 3–20 cm, shorter than the fertile leaves; petioles dark brown proximally, light brown to
green distally, grooved, with one vascular bundle; blades 2–4 -pinnate, lower surfaces glabrous, upper
surfaces glabrous to sparsely hairy; ultimate segments of sterile blades shortly stalked or sessile,
usually less than 4 mm wide, with flat, dentate to deeply flat at maturity. Sporangia scattered along
veins, often intermixed with farinose producing glands.
1a. Rhizomes (including persistent leaf bases) 10–20 mm thick, many branched; leaves densely
tufted, blades of sterile leaves somewhat leathery, overwintering; petioles dark for no more than
1/8 their length ......................................................................................................................... C. acrostichoides
1b. Rhizomes (excluding leaf bases) 1–1.5 mm thick, little branched; leaves scattered, blades of sterile
blades delicate, not overwintering; petioles dark brown up to 1/2 their length ................................. C. stelleri
Cryptogramma acrostichoides R. Br. [American parsleyfern; IMF 1:198–199 as C. crispa var.
acrostichoides (R. Br.) C.B. Clarke; FNA 2:137, 139; UF 9 as C. crispa var. acrostichoides]. [Global
distribution; Encyclopedia of Life; Intermountain Biota]
New growth starts in spring, sporulation is in summer, sterile leaves overwinter, senescing in their
second spring. Non-calcareous crevices, rocks, and talus, up to timberline.
Cryptogramma stelleri (Gmel.) Prantl [Delicate cliffbrake, Slender cliffbrake; IMF 1: 198–199; FNA 2:
138; UF 9] [Global distribution; Encyclopedia of Life; Intermountain Biota]
New growth develops in spring, withers by late summer. Sheltered crevices and ledges on calcareous
substrates, usually in coniferous zone.
Pellaea Link [Wikipedia]
Plants usually on rock. Rhizomes compact to elongate, usually branched, covered with scales; scales
brown to tan or with a dark central stripe, margins entire, erose or dentate. Leaves monomorphic to
20
somewhat dimorphic, clustered or scattered; petioles pale to dark, straw-colored to brown or gray to
blackish, mostly glabrous or hairy, usually with a few scales at the base, with one vascular bundle;
blades 1–4 pinnate, usually leathery, sometimes somewhat herbaceous, lower surfaces glabrous, hairy
or with hairlike scales along the veins, upper surfaces usually glabrous; ultimate segments usually
wider than 4 mm, usually stalked, margins reflexed, forming confluent false indusia extending around
the whole segment. Sporangia scattered along veins near the segment margins.
Pellaea includes about 40 species, most of which grow in the Western Hemisphere. There are six in the
IMR. One additional species is cultivated indoors.
1a. Petioles densely and conspicuously scaly, also with hairs; leaves 20-55 cm long; pinnae round to
oval ................................................................................................................................................. P. roundifolia
1b. Petioles almost naked to somewhat scaly, without hairs; leaves 2-55 long; pinnae often elongate ................ 2
2a. Rhizome scales bicolored, with a dark central region and lighter, brown margins ........................................... 3
3a. Ultimate segments of the blades with greenish borders; blades 3-pinnate at the base or, if 2pinnate, pinnae strongly ascending; sporangia shortly stalked ......................................................P. mucronata
3b. Ultimate segments with whitish borders; blades 2-pinnate proximally, pinnae perpendicular or
slightly ascending; sporangia long-stalked ........................................................................................................ 4
4a. Blades 4–18 cm wide; pinnae with 9-25 segments; borders of ultimate segments almost entire.... P. truncata
4b. Blades 1.5–5 cm wide; pinnae with 3-9 segments; borders of ultimate segments crenulate ........ P. wrightiana
2b. Rhizome scales uniformly reddish brown or tan ............................................................................................... 5
5a. Petioles reddish purple to nearly black, without prominent articulation lines;rachises glabrous or
nearly so......................................................................................................................................P. atropurpurea
5b. Petioles brown, sometimes with prominent articulation lines near the base; rachises glabrous or nearly so .....................6
6a. Pinnae at the base of the leaves usually deeply 2-lobed; veins of ultimate segments evident .......... P. breweri
6b. Pinnae at the base of the leaves usually with 3–7 lobes or ultimate segments; veins of ultimate
segments usually obscure ................................................................................................................... P. glabella
Pellaea atropurpurea (L.) Link [Purple cliffbrake; FNA 2: 175, 185] [Global distribution; Encyclopedia of
Life; Intermountain Biota]
Sporulating summer-fall. Calcareous, usually limestone, cliffs and rocks.
Pellaea breweri D.C. Eaton [Brewers cliffbrake; IMF 1: 200–201; FNA 2: 183; UF 10] [Global
distribution; Encyclopedia of Life; Intermountain Biota]
Sporulating summer-fall. Cliffs, ledges, and talus slopes from the foothills to treeline.
Pellaea glabella Mett. ex Kuhn [Suksdorf’s cliffbrake; IMF 1:201–202 as P. glabella subsp. simplex
(Butters) Á. Löve & D. Löve; FNA 2: 183–185; UF 10] [Global distribution; Encyclopedia of Life;
Intermountain Biota]
Sporulating summer-fall. Calcareous cliffs and ledges, usually on limestone. There are four subspecies
of P. glabella. According to the FNA account (2: 184), two of are present in the IMR:
1a. Sporangia containing 64 spores; spores averaging 35–52 µm ............................................. subsp. occidentalis
1b. Sporangia containing 32 spores; spores averaging 60–70 µm .................................................... subsp. simplex
The distributions of these two subspecies overlap in northeastern Utah. From there, Pellaea glabella
subsp. simplex (Butters) Á. Löve & D. Löve extends through southeastern Utah into northern Arizona
and Colorado, Pellaea glabella subsp. occidentalis (E.E. Nelson) Windham through eastern Idaho to
21
Manitoba and Wyoming. When the IMF account was prepared, only subsp. simplex had been found in
the IMR.
Pellaea mucronata (D.C.Eaton) Windham [IMF 1:203; FNA ] [Global distribution; Encyclopedia of Life;
Intermountain Biota]
There are two subspecies of P. mucronata. According to the maps in the account FNA (2:183), Pellaea
mucronata (D.C.Eaton) Windham subsp. mucronata grows in southern Nevada whereas Pellaea
mucronata subsp. californica (Lemmon) Windham grows in southern California., extending to the
Nevada border. They differ as follows:
1a. Blades 3-pinnate near the base; pinnae usually more or less perpendicular to the rachises, not
overlapping; usually below 1800 m ....................................................................................... subsp. mucronata
1b. Blades usually 2-pinnate near the base; pinnae ascending and overlapping, particularly distally;
usually above 1800 m .............................................................................................................. subsp. californica
Both subspecies grow on cliffs and rocky slopes. Pellaea mucronata (D.C. Eaton) Windham) subsp.
mucronata sporulates in late spring-summer and grows on acidic to mildly basic substrates. Pellaea
mucronata subsp. californica (Lemmon) Windham sporulates in summer- fall and usually grows on
granitic substrates.
Pellaea rotundifolia (Forst.) Hook. [Global distribution; Encyclopedia of Life; Intermountain Biota]
A New Zealand endemic that is apparently in cultivation.
Pellaea truncata Goodd. [Spiny cliffbrake; IMF 1:202; IMF 2:182; UF 11] [Global distribution;
Encyclopedia of Life; Intermountain Biota]
Sporulating late spring-fall. Crevices in cliffs and on open rocky slopes, from the Larrea to the oak and
juniper zones.
Pellaea wrightiana Hook. [Wright’s cliffbrake; not included in IMF; FNA 2: 175, 181–182; UF 11] [Global
distribution; Encyclopedia of Life; Intermountain Biota]
Sporulating summer-fall. Cliffs and rocky slopes, on both limestone and sandstone. At the time the IMF
account was written, P. wrightiana had not been found in the IMR.
Pentagramma Yatsk., Windham & E. Wollenw. [Wikipedia]
Plants terrestrial, sometimes on rocks. Rhizomes short, covered with scales; monomorphic, clustered;
petioles chestnut brown to black, rounded or nearly so at the base, glabrous, farinose or viscidglandular, with 1 vascular bundle; blades triangular-pentagonal, 1–2-pinnate-pinnatifid proximally,
pinnatifid distally, usually farinose on the lower surface, glabrous or glandular on the upper surface;
ultimate segments sessile, margins not recurved, veins free. Sporangia borne along the veins,
intermixed with farin-producing glands; indusium none.
Pentagramma used to be included, on the basis of superficial morphological characteristics, in
Pityrogramma Link, a genus that is now included in the Pteridaceae E.D.M. Kirchn. . Pentagramma is
restricted to western North America, including northwestern Mexico. One species grows in the IMR.
22
Pentagramma triangularis (Kaulf.) Yatsk. [Goldback fern, Silverback fern; IMF 1:194–195, as
Pityrogramma triangularis (Kaulf.) Maxon; FNA 2: 146, 150-151; UF 11 as Pityrogramma triangularis]
[Global distribution; Encyclopedia of Life; Intermountain Biota]
Sporulating ???. Rock crevices and open, rocky slopes in pine, and oak woodlands. Plants in the IMR
belong to P. triangularis (Kaulf.) Yatsk. subsp. triangularis. They have pale to bright yellow farina on
the lower surface of the blades. Some of the plants from Nevada and Utah that key here may represent
hybrids with P. triangularis subsp. maxon (Weath.) Yatsk., Windham & E. Wollenw. whose range lies to
the south of the IMR (Yatskievych and Windham FNA 2: 151).
Pteris L. [Brake fern] [Wikipedia]
Plants 1-10 m tall. Stems erect or creeping, with pale brown to black, elongate, uniformly colored
scales. Leaves monomorphic; petioles variously colored, longitudinally ridged, with 2-3 adaxial grooves,
scaly at the base, usually with 1, sometimes 2 or more, vascular bundles; blade 1-4-pinnate, glabrous,
pubescent, or scaly, abaxial surface dull; ultimate segments 1.5-8 mm wide; veins conspicuous, free.
Sori marginal, basically continuous; false indusium present. None native in the IMR. The following
species can be grown indoors.
1a. Leaves strictly 1-pinnate, pinnae not lobed or divided......................................................................... P. vittata
1b. Leaves not strictly 1-pinnate, at least the lowest pinnae lobed, forked, or divided............................. P. cretica
Pteris cretica L. [Global distribution; Encyclopedia of Life; Intermountain Biota]
Native in tropical and sub-tropical regions around the world.
Pteris vittata L. [Global distribution; Encyclopedia of Life; Intermountain Biota]
Native to Asia but escaped from cultivation in various parts of the southern U.S.A., not including the
IMR.
Cystopteridaceae Smakov [Wikipedia]
Plants small to medium sized, growing in forests or in crevices. Blades 1-3-pinnate, veins free, sori
small, rounde to cup-shaped, dorsal on veinlets; indusia,, if present, small, thin, ovate or oval, attached
at the bas, scalelike and immersed by mature sporangia. This family is one of several split out of the
Woodsiaceae by Christenhusz et al. (2011). It includes four genera, one of which is present in the IMR.
Cystopteris Bernh. [Wikipedia]
Plants terrestrial or on rock. Rhizomes short or long. Petioles 1/3–3 times the length of the blades;
blades ovate-lanceolate to deltate, gradually narrowing distally, axils or pinnae and/or lower surface
sometimes with uniseriate, multicellular hairs or unicellular glandular hairs, sometimes without hairs.
Sori in 1 row between midrib and margin, lying across the veins; indusia cuplike, attached on side of
sorus nearest pinnule midvein.
There are about 20 species of Cystopteris.
1a. Rachises, costae, indusia, and midribs or ultimate segments sparsely to densely covered with
glandular hairs; blades deltate to ovate, usually widest at or near the base .................................................... 2
2a. Rachises and costae frequently with bulblets; rachises, costae, indusia, and midribs usually
densely covered with glandular hairs; early blades lacking sori; petioles reddish when young,
becoming green or straw-colored when mature ............................................................................... C. bulbifera
23
2b. Rachises and costae frequently with bulblets; rachises, costae, indusia, and midribs usually
sparsely covered with glandular hairs; all blades with sori; petioles green to straw-colored
throughout or darker at the base ..................................................................................................... C. utahensis
1B. Rachises, costae, indusia, and midribs of ultimate segments without glandular hairs; leaf blades
elliptic to lanceolate, usually widest at or just below the middle;rachises and costae without
bulblets .............................................................................................................................................................. 3
3a. Blades 2–3-pinnate; lowest pinnae pinnate-pinnatifid to 2-pinnate, pinnules on the lower side
usually stalked; spores 33–41 µm ................................................................................................... C. reevesiana
3b. Blades 2-pinnate-pinnatifid; lowest pinnae pinnatifid to pinnate-pinnatifid, pinnules on lower side
sessile or stalked; spores 39–60 µm .................................................................................................................. 4
4a. Pinnae usually forming an acute angle with the rachis, often curvingtowards the blade tip; distal
pinnae usually ovate to narrowly elliptic, the margins usually crenate or with rounded teeth ............ C. tenuis
4b. Pinnae perpendicular to the rachises, not curving towards the blade tip;distal pinnae deltate to
ovate, the margins with sharp teeth..................................................................................................... C. fragilis
Cystopteris bulbifera (L.) Bernh. [Mountain bladder-fern; [IMF 1: 217–218, FNA 2:265–266; UF 9]
[Global distribution; Encyclopedia of Life; Intermountain Biota]
Sporulating summer-fall. Usually on moist soils at low elevations, sometimes on calcareous rocks.
Cystopteris fragilis (L.) Bernh. [Brittlefern;. IMF 1: 217–218, FNA 2: 269–270; UF 9] [Global distribution;
Encyclopedia of Life; Intermountain Biota]
Sporulating summer-fall. Usually on cliff faces or in thin soil over rocks. It has been confused with C.
reevesiana but tends to grow at higher elevations and, because its rhizomes have short internodes, its
more densely clumped leaves and petiole bases. The description in IMF may have encompassed what
are treated here as C. reevesiana and C. tenuis, with both of which C. fragilis hybridizes.
Cystopteris reevesiana Lellinger [Southwestern Brittlefern; not in IMF; FNA: 2: 266,268–269; UF 9, not
recognized as distinct] [Global distribution; Encyclopedia of Life; Intermountain Biota]
Sporulating summer-fall. Terrestrial or on rock, usually at lower elevations and further south than C.
fragilis. It was not recognized as distinct until after publication of IMF 1; in that work, specimens would
key to C. fragilis. Cystopteris reevesiana can usually be distinguished from C. fragilis by its more widely
spaced leaves but plants growing in rocks may have strongly clustered leaves.
Cystopteris tenuis (Michx.) Desv. [Mackay’s Brittlefern; FNA 2: 269] [Global distribution; Encyclopedia
of Life; Intermountain Biota]
Sporulating summer-fall. In shade, usually on rock or cliffs, sometimes on forest floor. The presence of
this species (which used to be known as C. fragilis var. mackayi G. Laws.) in western North America was
not recognized when IMF 1 was published. Specimens would key to C. fragilis in that work.
Cystopteris utahensis Windham & Laufler [Utah Brittlefern; [FNA 2: 267; UF included in C. fragilis]
[Global distribution; Encyclopedia of Life; Intermountain Biota]
Sporulating summer-fall. Cracks and ledges on sandstone, limestone and calcic cliffs. The species was
not recognized as distinct until after publication of IMF 1. In that work, specimens would probably be
identified as C. bulbifera if they have bulblets and as C. fragilis if they did not.
Aspleniaceae Newman [Wikipedia]
Plants terrestrial, often on rock. Rhizomes usually short, scaly at the apices, scales lattice-like in
appearance. Leaves monomorphic; petioles with 1 X-shaped vascular bundle; or 2 back-to-back C-
24
shaped bundles; blades simple to 4-pinnate, usually with glandular hairs, veins pinnate or forking,
usually free. Sori along the veins, lunar to linear; indusia usually present, originating on one side of
sorus. Sporangia stalked; annulus vertical.
There two genera in the family. One, Asplenium, is present in the IMR.
Asplenium L. [Wikipedia]
Stems (rhizomes) usually erect, rarely creeping in the IMR, consequently the leaves clumped. Petioles
with 1 vascular bundle, this X-shaped in cross-section, petiole bases with scales having a lattice-like
appearance; blades 1–4-pinnae, of diverse size and shape. Indusia present, originating along one side
of the sori.
There are about 700 species of Asplenium in the world. Many of them hybridize naturally and many of
its species are polyploids derived from such hybrids.
1a. Blades simple, not divided; plants requiring a greenhouse in winter .................................................... A. nidus
1b. Blades divided; plants native ............................................................................................................................. 2
2a. Blades not evidently pinnate, frequently forking into segments 1 – 2 cm long, 2 – 3 mm wide,
with a few teeth; sorus elongate, extending along the segments ............................................ A. septentrionale
2b. Blades clearly pinnately compound for all or most of their length; pinnae with several sori ........................... 3
3a. Blades 2 –4 pinnate or pinnatifid, lower pinnae evidently larger than the middle pinnae
.............................................................................................................................................A. adiantum-nigrum
3b. Blades 1-pinnate, the pinnae undivided, lower and middle pinnae similar in size............................................ 4
4a. Pinnae 6 – 11 mm long, to 5 mm wide, length of middle pinnae 3 or more times width .................. A. resiliens
4b. Pinnae 2.5 – 9 mm long, to 7 mm wide, length of middle pinnae up totwice width.
5a. Rachises dark reddish to purplish brown throughout their length.............................................. A. trichomanes
5b. Rachises mostly green, reddish brown only at the base............................................. A. trichomanes-ramosum
Asplenium adiantum-nigrum L. [Triangular spleenwort; IMF 1: 212–213, FNA 2: 240–241; UF 7] [Global
distribution; Encyclopedia of Life; Intermountain Biota]
On sandstone cliffs at 1675 – 2300 m, rare.
Asplenium nidus L. [Bird’s-nest fern] [Global distribution; Encyclopedia of Life; Intermountain Biota]
Frequently grown in greenhouses, A. nidus is a common species on trees in rain forests of the Old
World tropics.
Asplenium resiliens Kunze [Black-stemmed spleenwort; IMF 1:212–213, FNA 2:239; UF 7] [Global
distribution; Encyclopedia of Life; Intermountain Biota]
Rock crevices, usually on limestone, sometimes on sandstone at 100–1500 m.
Asplenium septentrionale (L.) Hoffm. [Forked spleenwort; IMF 1: 214; FNA 2:235-236; UF 7]. [Global
distribution; Encyclopedia of Life; Intermountain Biota]
Cliffs and rock crevices at 700–2900 m.
Asplenium trichomanes L. [Maidenhair spleenwort; IMF 212–213, FNA 2: 239; UF 7] [Global
distribution; Encyclopedia of Life; Intermountain Biota]
Usually on sandstone, granite or basalt at 0–3000 m. Plants in the IMR belong to Asplenium
trichomanes L. subsp. trichomanes They differ from those on the west and east coasts of North
America in being tetraploid rather than diploid.
25
Asplenium trichomanes-ramosum L. [Green spleenwort; IMF 1:21–213, as A. viride, FNA 2: 240]
[Global distribution; Encyclopedia of Life; Intermountain Biota]
Rocky cliffs and crevices in limestone or other basic rocks, to 4000 m.
Woodsiaceae Herter [Wikipedia]
Plants usually terrestrial. Rhizomes ascending, erect, or horizontal, apices with scales; scales usually
not lattice-like in appearance. Leaves monomorphic; petioles with 2 elongate or lunar vascular
bundles, united distally; veins pinnate or forking, usually not anastomosing. Sori round, J-shaped, or
linear; indusia absent or reniform to linear.
Christenhulz et al. (2011) adopted a narrow interpretation of the Woodsiaceae. In their treatment, the
family has three genera, only one of which, Woodsia, is in the IMR.
Woodsia R. Br. [Wikipedia]
Plants usually on rock. Rhizomes usually ascending or erect. Leaves monomorphic, usually dying back in
winter; petioles 1/5–3/4 the length of the blades; blades linear to lanceolate or ovate, 1–2-pinnatepinnatifid, glandular hairs usually present on both surfaces, sometimes absent; pinnae with entire to
dentate margins, lowest pinnae somewhat smaller than the adjacent pinnae. Sori in 1 row between
midrib and margin, round; indusia with several to many radiating filamentous or scale-like segments,
often obscure in mature sori.
1a. Both surfaces of pinnae with flattened, multicellular hairs along the midrib; mature petioles
relatively brittle and easily shattered ............................................................................................. W. scopulina
1b. Neither surface of pinnae with flattened, multicellular hairs along the midrib; mature petioles
somewhat pliable and resistant to shattering ................................................................................................... 2
2a. Rachis with scattered glandular hairs and occasional hairlike scales; indusia composed of
numerous filamentous segments that are uniseriate most of their length...................................... W. oregana
2b. Rachis with abundant glandular hairs; indusia composed of relatively few, broad segments that
are multiseriate at the base, often dividing into uniseriate segments distally ............................ W. plummerae
Woodsia oregana D.C. Eaton [Oregon cliff-fern; [IMF 1:218–219, FNA 2: 274, 278–279; UF 12] [Global
distribution; Encyclopedia of Life; Intermountain Biota]
Sporulating summer-fall. Cliffs and rocky slopes. There are two subspecies that tend to have separate,
but overlapping ranges. They differ in chromosome number as well as in the characters used in the
key. Cronquist (IMF 1: 219) did not distinguish the two and was not aware of the existence of
tetraploids in the region.
1a. Pinnae margins appearing entire; spells averaging 39–45 µm. .................................................. subsp. oregona
1b. Pinnae margins usually minute and appearing ragged; spores averaging 45–50 µm ......... subsp. cathcartiana
Woodsia oregana subsp. cathcartiana (B.L.Rob.) Windham is widely distributed in the IMR, growing on
both acidic and basic rocks. 2n = 152. The range of Woodsia oregana D.C.Eaton subsp. oregana
extends from the west coast south to northern California and Utah and east to western Wyoming. It
usually grows on granitic or volcanic substrates. 2n = 76.
Woodsia plummerae Lemmon [Plummer’s cliff-fern; FNA2: 274, 276] [Global distribution;
Encyclopedia of Life; Intermountain Biota]
Sporulating late-fall. Rocky slopes and cliff, usually on granitic or volcanic soils. It grows just south of
the region covered by the IMF and UF and was not, therefore, included in either work.
26
Woodsia scopulina D.C. Eaton [Mountain cliff-fern; IMF 1:219–219, FNA 2: 274–276; UF 12] [Global
distribution; Encyclopedia of Life; Intermountain Biota]
Sporulating summer–fall. Rocky slopes and cliffs, both basic and acidic. Since publication of IMF 1, two
subspecies have been recognized in the IMF:
1a. Scales of stems and petioles usually all of one color, sometimes with a few dark cells; spores
averaging 42–50 µm .................................................................................................................. subsp. scopulina
1b. Scales of stems and petioles with clusters of dark cells forming a narrow, discontinuous stripes;
spores averaging 50–57 µm ................................................................................................... subsp. laurentiana
The two subspecies grow in similar habitats and have similar ranges but differ in their chromosome
numbers (2n = 76 in W. scopulina D.C. Eaton subsp. scopulina, 2n = 152 in W. scopulina subsp.
laurentiana Windham) as well as the characters shown in the key. Hybrids between the two are sterile.
Blechnaceae Newman [Wikipedia]
Rhizomes short and vertical or long and horizontal, often with stolons, apices scaly, scales not latticelike. Leaves monomorphic or dimorphic; petioles with numerous round vascular bundles arranged in a
ring; blades usually pinnate or pinnatifid, sometimes simple or 2-pinnate., with scales on the surfaces,
scales not lattice- like. Sori in chains or linear, often parallel and adjacent to midribs; indusia linear,
opening towards the midveins of the pinnules sporangia stalked.
There are about nine genera in the Blechnaceae but only one, Woodwardia, is known from the IMR.
Woodwardia Sm. [Wikipedia]
Plants usually terrestrial, rarely on rock. Rhizomes horizontal, not climbing; scales brown. Leaves
monomorphic or dimorphic, clustered or separated. Petioles and veins scaly, scales not lattice-like;
blades pinnate or pinnatifid. Sori discrete, in chain-like rows along midveins of pinnules; indusia flaplike, opening towards the midveins of the pinnules.
One species of Woodwardia grows in Nevada.
Woodwardia fimbriata Sm. [Giant chainfern; IMF 208–209, FNA 224, 227; not in Utah] [Global
distribution; Encyclopedia of Life; Intermountain Biota]
In mesic to wet places along stream banks in coniferous woods. Although primarily a coastal species,
W. fimbriata also grows at disjunct locations in Nevada and Arizona.
Athyriaceae Alston [Wikipedia]
Recognition of this family is recent. The IMR has representatives of one of its six genera.
Athyrium Roth [Wikipedia]
Plants terrestrial. Rhizomes short, often ascending. Leaves monomorphic, usually not overwintering.
Petioles up to about 1/2 the length of the blades; blades lanceolate to elliptic or oblanceolate, 1–3pinnate or pinnatifid; pinnae gradually reduced distally, margins serrulate or crenate. Sori in 1 row
between midrib and margin,; indusia absent or attached on one side, horseshoe-shaped or cuplike and
almost round to elongate.
There are about 180 species of Athyrium in the world but only two in the IMR.
1a. Sori round, submarginal; indusia absent or highly reduced; veinlets obscure on underside of
pinnae ................................................................................................................................................. A. alpestre
27
1b. Sori elongate, , medial; indusia usually present; veinlets evident on underside of pinnae........... A. filix-femina
40.2 Athyrium alpestre (Hoppe) Ryl. [Alpine ladyfern; IMF 1: 215–216 as A. distentifolium Tausch ex
Opiz ;FNA 2: 256; UF 7 as A. distentifolium Tausch ex Opiz] [Global distribution; Encyclopedia of Life;
Intermountain Biota]
Wet rocky slopes, talus, and alpine meadows. North American plants belong to Athyrium alpestre var.
americanum Butters. It differs from var. alpestre (Hoppe) Ryl. in several qualitative features.
40.3 Athyrium filix-femina (L.) Roth [Ladyfern; IMF 1:215 [Global distribution; Encyclopedia of Life;
Intermountain Biota]
Moist woods, meadows, and stream banks below timberline. Two varieties grow in the IMR. They grow
in similar habitats but different, but overlapping, geographic regions.
1a. Lobes of pinnae narrowly deltate to oblong-lanceolate, base about as long as the sides ......... var. cyclosorum
1b. Lobes of pinnae linear-oblong to linear lanceolate, base shorter than the sides ..................... var. californicum
Athyrium filix-femina var. californicum Butters [Southwestern ladyfern] grows in the southwestern
US, extending north through southern Washington and Idaho. Athyrium filix-femina var. cyclosorum
Rupr. grows in northwestern North America, extending from Oregon, Idaho, and Montana to Alaska
and Yukon Territory.
Dryopteridaceae Herter [Wikipedia]
Plants growing on the ground, rocks, or trees. Rhizomes short or long, erect to horizontal, sometimes
climbing or scarmbling, with scales at the apices; scales not lattice-like. Leaves usually monomorphic,
sometimes dimorphic, overwintering or not, sometimes bearing scales or glandular hairs; petioles 1/4–
2/3 the length of the blades, with numerous, round, vascular bundles arranged in a ring; blades usually
1–2-pinnate. Sori usually round; indusia absent or, if present, round to reniform, sometimes peltate;
sporangia stalked.
There are 40–45 genera and 1700 species in the Dryopteridaceae. Two genera grow in the IMR. The
indusia in both genera in the IMR tend to fall off rapidly so care must be taken to search for them on
young sori.
1a. Veins anastomosing (coming together); rachis groove not open to grooves from midrib of pinnae Cyrtomium
1b. Vein ends free, rachis groove open to grooves from the midrib of pinnae ....................................................... 2
2a. Indusia attached on one side of the sori, round-reniform; blades widest near the middle; pinnae
pinnatifid ............................................................................................................................................. Dryopteris
2b. Indusia attached in the center, round; blades widest near the base; pinnae entire or pinnatifid
only at the base ................................................................................................................................ Polystichum
Cyrtomium C. Presl Holly fern [Wikipedia]
Plants terrestrial.
There are 15-20 species of Cyrtomium. They are very similar to, and possibly should be included in
Polystichum.
Cyrtomium falcatum (L. f.) C. Presl [Global distribution; Encyclopedia of Life; Intermountain Biota]
This species is native to China, Japan, and Korea.
28
Dryopteris Adans. [Wikipedia]
Plants usually terrestrial, rarely on rock. Rhizomes short, not developing stolons. Leaves monomorphic,
clustered; petioles 1/4-2/3 blade length; blades deltate-ovate to lanceolate, 1–3-pinnate-pinnatifid,
lower pinnae as long as or shorter than those at midlength, lower surfaces with linear to ovate scales.
Sori in 1 row between margins and teeth, round; indusia round-reniform, attached at a narrow sinus,
persistent or rapidly deciduous.
There are about 250 species of Dryopteris in the world but only one grows in the IMR.
Dryopteris filix-mas (L.) Schott [Male fern; IMF 1: 215–217, FNA 2: 284; UF 10] [Global distribution;
Encyclopedia of Life; Intermountain Biota]
Dense woods and thickets, often along streams.
Polystichum Roth [Wikipedia]
Plants terrestrial. Rhizomes horizontal to erect, Leaves usually monomorphic, overwintering; petioles
1/9–1 times the length of the blades; blades linear to broadly lanceolate, 1–3-pinnate, with linear to
lanceolate scales on the lower, sometimes also the upper, surfaces. Sori usually in 1 row between the
midribs and the margins, usually round and distinct, sometimes confluent; indusia peltate, persistent
or not.
Polystichum includes about 180 species; three grow in the IMR.
1a. Pinnae upswept with respect to the blade rachis, spinulose, with one lobe at the base.. Polystichum lonchitis
1b. Pinnae straight with respect to the blade rachis, crenate and mucronate to shortly spinulose,
lower pinnae with more than one basal lobe .................................................................................................... 2
2a. Lower pinnae elongate, length at least three times the width, their lobes and tip tending to be
acute ............................................................................................................................ Polystichum kruckebergii
2b. Lower pinnae oblong, length 2-3 times the width, their lobes and tips rounded ........ Polystichum scopulinum
Polystichum kruckebergii D.H. Wagner [Kruckeberg’s swordfern; IMF: 210–211, FNA 2: 296; UF 12]
[Global distribution; Encyclopedia of Life; Intermountain Biota]
Crevices and talus slopes in subalpine to alpine habitats.
Polystichum lonchitis (L.) Roth [Holly fern; IMF 1: 210–211, FNA 2: 297; UF 12] [Global distribution;
Encyclopedia of Life; Intermountain Biota]
Rock crevices and at the base of boulders in boreal, subalpine coniferous forests and in subalpine
regions.
Polystichum scopulinum (D.C. Eaton) Maxon [Rock swordfern; IMF 1: 210–211, FNA 2:298; UF 12
2: 297–298] [Global distribution; Encyclopedia of Life; Intermountain Biota]
Rock crevices and open rocky slopes, usually in full sun, at middle and upper elevations but below
timberline.
Davalliaceae M.R. Schromb. [Wikipedia]
Davallia Sm. [Haresfoot fern, Rabbitsfoot fern; information from Wikipedia and Bailey’s
Taxonomy of Cultivated Plants; Wikipedia]
29
Plants terrestrial or epiphytic. Stems conspicuous above ground, thick, covered in scales. Blades
triangular or ovate, 1-3-pinnate; sori at or near the margins, indusial tubular or semi-cylindric and
opening towards the margin, attached at the base and the side.
There are no native or established species of Davallia in the Intermountain Region. They are, however,
relatively common in greenhouses. The following key to is taken from Morton, C.V. 1957. Observations
on cultivated ferns, IV. The species of Davallia. American Fern Journal 47:143-148.
1a.
2a.
2b.
3a.
3b.
4a.
4b.
1b.
5a.
5b.
6A.
6b.
7a.
7b.
Scales of the rhizomes toothed but not hairy, the teeth short, composed of two marginal cells. ................... 2
Petioles less than 1 mm in diameter................................................................................................... D. mariesii
Petioles 1-1.5 mm in diameter .......................................................................................................................... 3
Leaves with stripes (“false veins” between the true veins ............................................................D. denticulata
Leaves without stripes between the veins ........................................................................................................ 4
Rhizome scales 1 mm wide or more, gradually tapering to the tip ................................................. D. divaricate
Rhizome scales less than 1 mm wide, abruptly narrowed about the base............................. D. trichomanoides
Scales od the rhizomes bearing long, multicellular hairs on the margins neat the tip, sometimes
also with a few teeth ......................................................................................................................................... 5
Rhizomes 8-13 mm think covered with persistent, large scales, these with brown bases and
centers and wide, pale margins ..................................................................................................... D. canariensis
Rhizomes usually 5-7 mm in diameter, the scales with a blackish base that persists after the rest
of the scale has fallen off6
Blades finely divided, the ultimate segments linear and 1-veined ................................................... D. Fejeensis
Blades coarser, the ultimate segments severa-veined ...................................................................................... 7
Indusium cuplike, barely longer than wide ........................................................................................ D. pyxidata
Indusium tubular, much longer than wide ............................................................................................ D. solida
Davallia canariensis Smith [Canary Island Rabbitsfootfern]
Encyclopedia of Life]
This species is native to Grand Canary Island, Tereiffe, Madeira, and Portugal.
[Global distribution;
Davallia denticulata (Burm. f.) Mett. ex Kuhn [Global distribution; Encyclopedia of Life;
Intermountain Biota]
Morton described it as a large and elegant species.
Davallia divaricata Blume [Global distribution; Encyclopedia of Life]
Another species described as large and elegant. In 2010, its value for treating osteoporosis was being
studied at the University of Taiwan.
Davallia fejeensis Hook. [Fijian Rabbitsfootfern] [Global distribution; Encyclopedia of Life
Native to the Fijian Islands.
Davallia mariesii T. Moore ex Baker [Global distribution; Encyclopedia of Life]
This species is native to Japan. According to Morton, it used to be imported bound into wires in the
form of a monkey – with a porcelain monkey’s head on top. One them hung the “monkey” up on a
porch or in a greenhouse and started watering it – which would makes its leaves appear.
Davallia pyxidata Cav. [Global distribution; Encyclopedia of Life]
This is an Australian species which was not commonly cultivated when Morton was writing.
Davallia solida (G. Forst.) Sw. [Global distribution; Encyclopedia of Life]
Described my Morton as having a coarse aspect. It is native to Polynesia.
30
Polypodiaceae Bercht. & J. Presl [Wikipedia]
Plants usually epiphytic or epipetric [on rocks], sometimes terrestrial. Rhizomes horizontal, bearing
scales. Leaves monomorphic or dimorphic; petioles often breaking off near their bases; blades usually
simple to pinnatifid or 1-pinnate, sometimes 2- or more pinnate, surfaces sometimes with hairs or
scales, sometimes glandular. Sori usually round to oblong or elliptic, sometimes elongate or sporangia
densely distributed over a portion of the blades, not forming sori; indusia absent, young sori
sometimes covered by scales that rapidly fall off; sporangia stalked, often intermixed with paraphyses.
Smith et al. (2006), whose family treatment is adopted here, include about 56 genera and 1200 species
in the Polypodiaceae but, in older works (such as IMF) it included almost all leptosporangiate ferns.
One genus, Polypodium, is native to the IMF. A second, Platycerium and Phlebodium are grown in
greenhouses.
1a. Leaves dimorphic, round leaves appressed to the tree or support and deeply lobed,
photosynthetic leaves; sporangia densely distributed over distal portion of the photosynthetic
leaves; plants indoor ornamentals in the IMR .................................................................................. Platycerium
1a. Leaves all alike, green; sporangia in circular to oval sori; plants native or indoor ornamentals ....................... 2
2a. Rhizomes green, 5-7 mm in diameter, with scattered dark scales; blades deeply pinnatifid .......... Microsorum
2b. Rhizomes brown and thinner, scales varied; blades pinnatifid to 1-pinnate..................................................... 3
3a. Veins free or forming 1 row or areoles between the midvein and margins of the pinnae; sori at
end of a veinlet ending in an areole or on a forked free vein ........................................................... Polypodium
3b. Venation highly reticulate, with 3-4 rows of areoles between midvein and margins of the pinnae;
sori usually at the end of 2 veinlets that end in an areole................................................................ Phlebodium
Microsorum Link [Wikipedia]
Plants usually epiphytic or lithophytic, occasionally terrestrial; Rhizomes scaly, central cells of scales
with dark borders. Petioles jointed to the short stalks. Blades simple, entire, variously lobed or deeply
1-pinnatifid; veinlets ending in hydathodes. Sori round or somewhat elongate, superficial or impressed
into the blade.
Microsorum, as interpreted here, has about 50 species, all native to the Old World. The species in the
greenhouse of USU’s Logan campus is sometimes placed in Phymatosorus Pic. Serm. Bostock (1998) in
the Flora of Australia gave reasons for not doing so. His treatment is accepted here.
Microsorum grossum (Langsd. & Fisch.) S.B. Andrews ] [Global distribution; Encyclopedia of Life]
Leaves 8-90 cm long. Blade deeply pinnatifid, with 5-10(15) pairs of pinnae. Sori in (1)2(3) rows on each
side of the midvein, slightly impressed into the blade. This species is sometimes confused with
Microsorum scolopendria (Burm. F.) Copel. which usually has smaller (5-28(40) cm long) leaves with
fewer (1-4(5)) pairs of pinnae.
Phlebodium (R. Br.) J. Sm. [Wikipedia]
Plants epiphytic. Rhizomes branched, scales uniformly colored. Petiole readily breaking off, dark
brown, round in cross section, with 2 grooves; blades with less than 20 pairs of pinna, glaucous, usually
glabrous. Veins free near the margins, highly reticulate elsewhere, forming elongate areoles, those
cosest to the midvein including 1-3 veinlets. Sori on veinlets in areoles. There are 2-4 species.
31
Phlebodium aureum J. Sm. [Global distribution; Encyclopedia of Life]
Native to Florida and Georgia. The other species are more tropical in their distribution.
Platycerium Desv. [Wikipedia]
Plants with tufted roots from a short rhizome bearing two types of leaves, basal and fertile. Basal
leaves sterile, shield or kidney shaped, appressed to the tree, their upper margins catching litter and
water; fertile leaves dichotomously lobed (antler-shaped), horizontal or pendent. Sporangia densely
clustered in large sori located on the lobes or on the sinus between the lobes.
There are about 18 species of Platycerium. They are native in southeast Asia, Australia, Southern
Africa, and South America. Some species are solitary having only one rhizome. Other species form
colonies when their rhizomes branch or when new rhizomes are formed from root tips. If the
conditions are right the spores will germinate naturally on surrounding trees. Platycerium
gametophytes have a small heart shaped thallus.
A mature staghorn can grow more than a meter wide. When positioned well, Platycerium species are
able to add privacy and a natural look to a garden. Some species have Crassulacean Acid Metabolism,
an adaptation to hot, dry climates. [Information on Platycerium from wikipedia7 February 2011].
Platycerium bifurcatum (Cav.) C. Chr.. ] [Global distribution; Encyclopedia of Life]
This is the species present in the greenhouse of the Logan Campus of USU. It is the most frequently
cultivated species.
Polypodium L. [Wikipedia]
Plants usually on rock, sometimes terrestrial or epiphytic. Rhizomes horizontal, usually branched, scaly;
scales uniformly colored or bicolored, lanceolate to ovate- acuminate, sometimes lattice-like. Leaves
monomorphic, clustered or not, to 90 cm long; petioles straw-colored, winged distally, readily breaking
off; blades broadly ovate to deltate, pinnatifid or 1-pinnate at the base, usually with fewer than 25
pairs of pinnae, midveins sometimes sparsely scaly on lower surface, scales ovate- lanceolate to linear;
segments entire to erose, apices rounded to attenuate. Sori often restricted to distal portion of leaves,
circular to oval, borne at the tips of single veins, in 1–3 rows on either side of the midrib.
There are about 100 species of Polypodium; only one grows in the IMR.
Polypodium hesperium Maxon [Licorice fern, Western polypody; IMF 1:208–209, FNA 320] [Global
distribution; Encyclopedia of Life; Intermountain Biota]
Sporulating summer-fall. Wet crevices and ledges on cliffs, rarely on limestone.
32
REFERENCES
This is a preliminary list. It contains the primary sources of information. I shall add other references
later. For now, I have to develop a key for the monocots that, like this treatment, includes the plants
grown in the greenhouse of the main campus of Utah State University.
Christenhulz, M.J.M., X-C Zhang, and H. Schneier. 2011. A linear sequence of extant families and genera
of lycophytes and ferns.Phytotaxa 19:7-54.
Cronquist, A., A.H. Holmgren, N.H. Holmgren, and J.L. Reveal. 1972. Intermountain Flora, vol. 1. New
York Botanical Garden, The Bronx, New York.
Flora of North America Editorial Committee. 1993. Flora of North America north of Mexico, vol. 2.
Oxford University Press, New York.
Smith, A.R., K.M. Pryer, E. Schuettpelz, P. Korall, H. Schneider, and P.G. Wolf. 2006. A classification for
extant ferns. Taxon 55: 505–731.
Welsh, S.L., N.D. Atwood, S. Goodrich, and L.C. Higgins (Eds.). 2008. A Utah Flora. Printing Services,
Brigham Young University, Provo, Utah.
33
INDEX
My apologies. Accepted names are supposed to be in bold but I find Word a bit idiosyncratic when it
comes to formatting the index.
Adiantum aleuticum (Rupr.) Paris, 16
Adiantum capillus-veneris L., 17
Adiantum L., 16
Adiantum pedatum Paris, 16
Adiantum raddianum C. Presl, 17
Argyrochosma (J. Sm.) Windham, 17
Argyrochosma jonesii (Maxon) Windham, 17
Argyrochosma limitanea (Maxon) Windham, 17
Argyrochosma limitanea (Maxon) Windham subsp.
limitanea, 17
Aspidotis Copeland, 17
Aspidotis densa (Brackenb.) Lellinger, 17
Aspleniaceae Newman, 23
Asplenium adiantum-nigrum L., 24
Asplenium L., 24
Asplenium nidus L., 24
Asplenium resiliens Kunze, 24
Asplenium septentrionale (L.) Hoffm., 24
Asplenium trichomanes L., 24
Asplenium trichomanes L. subsp. trichomanes, 24
Asplenium trichomanes-ramosum L., 25
Asplenium viride Huds., 25
Athyriaceae Alston, 26
Athyrium alpestre (Hoppe) Ryl., 27
Athyrium alpestre (Hoppe) Ryl. var. alpestre, 27
Athyrium alpestre var. americanum Butters, 27
Athyrium distentifolium Tausch ex Opiz, 27
Athyrium filix-femina (L.) Roth, 27
Athyrium filix-femina var. californicum Butters, 27
Athyrium filix-femina var. cyclosorum Rupr., 27
Athyrium Roth, 26
Azolla Lam., 13
Azolla mexicana Presl, 13
Bird’s-nest fern, 24
Bladder-fern
Mountain, 23
Blechnaceae Newman, 26
Botrychium adscendens W.H. Wagner, 10
Botrychium crenulatum W.H. Wagner, 10
Botrychium hesperium (Maxon & R.T. Clausen) W.H.
Wagner & Lellinger, 10
Botrychium lanceolatum (S.G. Gmel.) Ångström, 11
Botrychium lanceolatum (S.G. Gmel.) Ångström subsp.
lanceolatum, 11
Botrychium lunaria (L.) Sw., 11
Botrychium matricariifolium (Döll) A. Braun ex W.D.J.
Koch, 10
Botrychium minganense Vict., 11
Botrychium multifidum (S.G. Gmel.) Rupr., 11
Botrychium paradoxum W.H. Wagner, 11
Botrychium pinnatum H. St. John, 11
Botrychium simplex E. Hitchc., 11
Botrychium Sw., 9
Botrychium virginianum (L.) Sw., 11
Bracken, 15
Brake fern, 22
Brittlefern, 23
Mackay’s, 23
Southwestern, 23
Utah, 23
Chainfern
Giant, 26
Cheilanthes covillei Maxon, 18
Cheilanthes eatonii Baker, 18
Cheilanthes feei Moore, 18
Cheilanthes gracillima D.C. Eaton, 19
Cheilanthes intertexta (Maxon) Maxon, 19
Cheilanthes parryi (D.C. Eaton) Domin, 19
Cheilanthes Sw., 18
Cheilanthes wootonii Maxon, 19
Cibotiaceae Korall, 14
Cibotium glaucum (Sm.) Hook. & Arn., 14
Cibotium Kaulf, 14
Cliff fern
Mountain, 26
Oregon, 25
Plummer’s, 25
Cliffbrake
Brewer's, 20
Delicate, 19
Purple cliffbrake, 20
Slender, 19
Spiny, 21
Suksdorf's, 20
Wright’s, 21
Cloakfern
Jones’, 17
Waxy, 17
Clubmoss
Bristly, 4
Club-moss
Prairie, 6
Cryptogramma acrostichoides R. Br., 19
Cryptogramma crispa var. acrostichoides (R. Br.) C.B.
Clarke, 19
Cryptogramma R. Br., 19
34
Cryptogramma stelleri (Gmel.) Prantl, 19
Cyathea cooperi (F. Muell.) Domin, 14
Cyatheales A.B. Frank, 14
Cyrtomium C. Presl, 27
Cyrtomium falcatum (L. f.) C. Presl, 27
Cystopteridaceae Smakov, 22
Cystopteris Bernh., 22
Cystopteris bulbifera (L.) Bernh., 23
Cystopteris fragilis (L.) Bernh., 23
Cystopteris fragilis var. mackayi G. Laws., 23
Cystopteris reevesiana Lellinger, 23
Cystopteris tenuis (Michx.) Desv., 23
Cystopteris utahensis Windham & Laufler, 23
Davallia canariensis Smith, 29
Davallia denticulate (Burm. f.) Mett. ex Kuhn, 29
Davallia divaricata Blume, 29
Davallia fejeensis Hook., 29
Davallia mariesii T. Moore ex Baker, 29
Davallia pyxidata Cav., 29
Davallia Sm., 28
Davallia solida (G. Forst.) Sw., 29
Davalliaceae M.R. Schromb., 28
Dennstaedtiaceae Lotsy, 14
Dryopteridaceae Herter, 27
Dryopteris Adans., 28
Dryopteris filix-mas (L.) Schott, 28
Equisetaceae Michx. ex DC., 8
Equisetidae Warm, 8
Equisetopsida C. Agardh, 6
Equisetum ×ferrissii Clute, 8
Equisetum arvense L., 8
Equisetum hyemale L., 9
Equisetum hyemale subsp. affine (Engelm.) Calder & R.L.
Taylor, 9
Equisetum L., 8
Equisetum laevigatum A. Br., 9
Equisetum variegatum Schleich. ex F. Weber & D. Mohr, 9
Equisetum variegatum Schleich. ex F. Weber & D. Mohr
subsp. variegatum, 9
Fern
Bird's-nest, 24
Four-leaf clover, 12
Goldback, 22
Haresfoot, 28
Holly, 27, 28
Lace, 19
Licorice, 31
Maidenhair, 16
Mosquito, 13
Rabbitsfoot, 28
Rattlesnake, 11
Silverback, 22
Grape-fern
Common, 11
Leather, 11
Haresfoot fern, 28
Holly fern, 27, 28
Horsetail, 8
Common, 8
Horsetails, 8
Indian’s dream, 17
Isoëtaceae Dumort., 4
Isoëtes bolanderi Engelm., 4
Isoëtes echinospora Durieu, 4
Isoëtes howellii Engelm., 4
Isoëtes L., 4
Isoëtes lacustris L., 5
Isoëtes melanopoda Gay & Durieu, 5
Isoëtes occidentalis L.F. Henders., 5
Lace-fern
Dense, 17
Ladyfern, 27
Alpine, 27
Southwestern, 27
Lemna L., 13
Licorice fern, 31
Lipfern
Coastal, 19
Coville’s, 18
Eaton’s, 18
Lace, 19
Parry's, 19
Slender, 18
Wooton’s, 19
Lycopodiaceae P. Beauv. ex Mirb., 3
Lycopodiopsida Bartl, 3
Lycopodium annotinum L., 4
Lycopodium L., 3
Maidenhair fern
Southern, 17
Western, 16
Male fern, 28
Marsilea L., 12
Marsilea oligospora Goodd., 12
Marsilea quadrifolia L., 12
Marsilea vestita Hook. & Grev., 13
Marsileaceae Mirb., 12
Microlepia C. Presl, 15
Microlepia platyphylla (Don) J. Sm., 15
Microsorum grossum (Langsd. & Fisch.) S.B. Andrews, 30
Microsorum Link, 30
Microsorum scolopendria (Burm. F.) Copel., 30
Moonwort
Common, 11
Dainty, 10
Least, 11
35
Mingan Moonwort, 11
Northwestern, 11
Paradox, 11
Triangle, 11
Upswept, 10
Western, 10
Mosquito fern, 13
Mountain-moss
Prickly, 6
Notholaena jonesii Maxon, 17
Notholaena limitanea Maxon, 17
Notholaena parryi D.C. Eaton, 19
Ophioglossaceae Martinov, 9
Ophioglossidae Klinge, 9
P. triangularis subsp. maxon (Weath.) Yatsk., Windham &
E. Wollenw., 22
Parsleyfern
American, 19
Pellaea atropurpurea (L.) Link, 20
Pellaea breweri D.C. Eaton, 20
Pellaea glabella Mett. ex Kuhn, 20
Pellaea glabella subsp. occidentalis (E.E. Nelson)
Windham, 20
Pellaea glabella subsp. simplex (Butters) Á. Löve & D.
Löve, 20
Pellaea jonesii (Maxon) Morton, 17
Pellaea limitanea (Maxon) Morton, 17
Pellaea Link, 19
Pellaea mucronata (D.C. Eaton) Windham subsp.
mucronata, 21
Pellaea mucronata (D.C.Eaton) Windham, 21
Pellaea mucronata subsp. californica (Lemmon)
Windham, 21
Pellaea rotundifolia (Forst.) Hook., 21
Pellaea truncata Goodd., 21
Pellaea wrightiana Hook., 21
Pentagramma triangularis (Kaulf.) Yatsk., 22
Pentagramma triangularis (Kaulf.) Yatsk. subsp.
triangularis, 22
Pentagramma Yatsk., Windham & E. Wollenw., 21
Pepperwort, 13
Phlebodium (R. Br.) J. Sm., 31
Phlebodium aureum J. Sm., 31
Phymatosorus Pic. Serm., 30
Pillwort, 13
American, 13
Pilularia americana A. Braun, 13
Pilularia L., 13
Pityrogramma Link, 21
Pityrogramma triangularis (Kaulf.) Maxon;, 22
Platycerium bifurcatum (Cav.) C. Chr., 31
Platycerium Desv., 30
Polypodiaceae Bercht. & J. Presl, 30
Polypodiidae Cronquist, Tahkt. & Zimmerm., 12
Polypodium hesperium Maxon, 31
Polypodium L., 31
Polystichum kruckebergii D.H. Wagner, 28
Polystichum lonchitis (L.) Roth, 28
Polystichum Roth, 27, 28
Polystichum scopulinum (D.C. Eaton) Maxon, 28
Psilotaceae J.W. Griff. & Henfr., 11
Psilotum nudum (L.) P. Beauv., 12
Psilotum Sw., 11
Pteridaceae E.D.M. Kirchn., 15, 21
Pteridium aquilinum (L.) Kuhn, 15
Pteridium aquilinum var. pubescens L., 15
Pteridium Gled. ex Scop., 15
Pteris cretica L., 22
Pteris L., 22
Pteris vittata L., 22
Quillwort
Black-footed, 5
Bolander’s, 4
Spiny-spored, 4
Western, 4, 5
Rabbitsfoot fern, 28
Canary Island, 29
Fijian, 29
Resurrection plant, 6
Salvinia minima Baker, 14
Salvinia molesta D.S. Mitch., 14
Salvinia natans All., 14
Salvinia Ség., 13
Salviniaceae Martinov, 13
Scouring rush, 9
Scouring-rush
Smooth, 9
Variegated, 9
Selaginella densa Rydb., 6
Selaginella lepidophylla (Hook. & Grev.) Spring, 6
Selaginella leucobryoides Maxon, 6
Selaginella mutica D.C. Eaton ex Underw., 6
Selaginella mutica D.C. Eaton ex Underw. var. mutica, 6
Selaginella selaginoides (L.) Link, 6
Selaginella underwoodii Hieron, 6
Selaginella utahensis Flowers, 6
Selaginella watsonii Underw., 6
Selaginellaceae Willk., 5
Southwestern ladyfern, 27
Spike-moss
Awnless, 6
Mojave, 6
Northern, 6
Rocky Mountains, 6
Underwood’s, 6
Utah, 6
36
Watson’s, 6
Spleenwort
Black-stemmed, 24
Forked, 24
Green, 25
Maidenhair, 24
Triangular;, 24
Swordfern
Kruckeberg’s, 28
Rock, 28
Water-clover, 13
Waterfern, 13
Western polypody, 31
Totals: 19 families, 32 genera, and 98 species.
Woodsia oregana D.C. Eaton, 25
Woodsia oregana D.C.Eaton subsp. oregana, 25
Woodsia oregana subsp. cathcartiana (B.L.Rob.)
Windham, 25
Woodsia plummerae Lemmon, 25
Woodsia R. Br., 25
Woodsia scopulina D.C. Eaton, 26
Woodsia scopulina D.C. Eaton subsp. scopulina, 26
Woodsia scopulina subsp. laurentiana Windham, 26
Woodsiaceae Herter, 25
Woodwardia fimbriata Sm., 26
Woodwardia Sm., 26