CROSSOSOMA

]ournal of the Southern California Botanists , Inc.

Volume 35, Number 1

Spring-Summer 2009

Southern California Botanists, Inc.

- Founded 1927 -
http://www. socalbot. org

CROSSOSOMA (ISSN 0891-9100) is published twice a year by Southern Cali-
fornia Botanists, Inc., a California nonprofit organization of individuals devoted
to the study, conservation, and preservation of the native plants and plant com-
munities of southern California.

SCB Board of Directors for 2009

President Gary Wallace

Vice President Naomi Fraga

Secretary Carrie Kiel

Treasurer Alan P. Romspert

Webmaster Naomi Fraga

Editors of Crossosoma Scott D. White and Michael Honer

Editor of Leaflets Kerry Myers

Directors-at-large

David Bramblet Gilberto Ocampo

Sara Baguskus Linda Prince

Elizabeth Delk Gina Richmond

Erica Gardner Fred Roberts

Charlie Hohn Darren Sandquist

Diane Menuz Allan Schoenherr

Orlando Mistretta Jonathan Snapp-Cook

Bart O’Brien Sula Vanderplank

and Katie Vinzant

Ex officio Board Member Sula Vanderplank (Past President)

Articles, book reviews, or other items for submission to CROSSOSOMA can be sent to
the editor Scott White (scottbioservices@verizon.net) or 201 N. First Ave., #102, Upland,
CA., USA, 91786. Electronic submission is preferred. Please see our website, www.
socalbot.org, for format guidelines. Notices of a time-dated nature (field trips, work-
shops, symposia, etc.) to be included in the newsletter Leaflets should be submitted to
Kerry Myers, Editor of Leaflets, kerrymyers@fs.fed.us, or mail to: Kerry Myers, Botanist,
SBNF Mountaintop Ranger Dist., 42300 North Shore Dr., Fawnskin, CA 92333-04004.

Views published in CROSSOSOMA are those of the contributing author(s) and are not
necessarily those of the editors, the membership of Southern California Botanists, Inc., or
the SCB Board of Directors, unless specifically stated.

Copyright © 2009 by Southern California Botanists, Inc. All rights reserved.
Permission to reproduce items in CROSSOSOMA, in whole or part,
should be requested from the Editor.

Crossosoma 35(1), Spring-Summer 2009

1

CRYPTANTHA OF SOUTHERN CALIFORNIA

Michael G. Simpson and Kristen E. Hasenstab

Department of Biology
San Diego State University
San Diego, California 92 1 82 USA
msimpson@sunstroke.sdsu.edu; kristen.hasenstab@cgu.edu
(Current address for K. Hasenstab: Rancho Santa Ana Botanic Garden, Claremont,
1500 N. College Ave., California 91711)

ABSTRACT: The genus Cryptantha (Boraginaceae) contains 202 species, with 49
species and 56 taxa (including varieties) occurring in Southern California, defined here
as including the entire Southwestern California region and Tehachapi Mountain region
of the California Floristic province, the entire Desert province, and most of the White
and Inyo Mountain subregion of the Great Basin province. The purposes of this article
are 1) to summarize the current taxonomy of Cryptantha species and infraspecies
in Southern California; 2) to provide taxonomic keys and images illustrating the
diagnostic features for identification; and 3) to review the distribution, environmental
factors, and current conservation status of these taxa.

KEYWORDS: Cryptantha , Boraginaceae, taxonomy, identification,

INTRODUCTION

Taxonomic History and Nomenclature

Cryptantha Lehmann ex G. Don, commonly known as “popcorn flower” or “cat’s
eye,” is a genus within the family Boraginaceae. The circumscription of this family
has changed repeatedly over the last twenty years [Engler and Prantl 1897, Heywood
et al. 2007, Gottschling et al. 2001, Angiosperm Phylogeny Group (APG II) 2003],
with various authors recognizing either a broad or narrow family concept. Here
we accept the APG II (2003) system of classification, which recognizes a broad
Boraginaceae. As treated in this manner, the family may be divided into subfamilies
Boraginoideae, Cordioideae, Eretioideae, Heliotropoideae, Hydrophylloideae,
and (possibly) Lennoideae (see Stevens 2001 onwards). Despite these changes in
classification, Cryptantha firmly belongs in subfamily Boraginoideae, characterized
by an inflorescence that is a scorpioid cyme (Buys and Hilger 2003), a deeply 4-lobed
ovary with a gynobasic style, and a fruit that is a schizocarp of nutlets.

Cryptantha is an amphitropically distributed genus, with taxa in both western North
America and western South America, but lacking populations in between. The genus is
currently catalogued at 202 species. The greatest species diversity, 136 species, occurs
in western North America, with distributions from Alaska to southern Mexico, and

2

Crossosoma 35(1), Spring-Summer 2009

as far east as Texas (Johnston 1925; Payson 1927; Higgins 1971; Mabberley 2008).
Approximately 69 species occur in western South America, in Peru, Chile, Bolivia,
and east to Argentina (Reiche 1915; Johnston 1927; Schwarzer 2007; Zuloaga et al.
in prep). Three species, C. albida, C. circumscissa, and C. maritima , are distributed in
both North and South America.

Members of the genus are annual, biennial, or perennial herbs. Stems are simple
to highly branched, generally ascending to erect, and densely covered in trichomes
(Johnston 1925; Payson 1927; Munz and Keck 1959, 1968; Higgins 1971; W.A.
Kelley and Wilken 1993, Mabberley 2008; R.B. Kelley in prep). Often in perennial
taxa, a basal rosette of robust leaves is present before and during inflorescence growth.
Among annuals, the basal rosette is common early in development but is lost as the
internodes of the aerial shoots (including inflorescence) elongate. Cauline leaves
range from opposite to alternate in arrangement. Leaf shape is spatulate, lanceolate,
oblanceolate, but most often linear. Leaves have acute to obtuse bases. Leaf vestiture
is strigose, “rough hairy,” or “bristled.” The largest, relatively stout trichomes, which
are often designated as bristles, often have bulbous bases, described as pustulate
(giving a tessellate appearance to the leaf surface). These pustules are composed of
epidermal cells arranged in a ring around the trichome base; the epidermal cells are
elevated, opaque, and silicified and are thought to be cystolith in origin (Johnston
1925). The trichomes are most often translucent and white, but can be yellow-brown
in some taxa. The inflorescence is characteristic of subfamily Boraginoideae, being
a circinate (coiled), scorpioid cyme unit (often described as a helicoid cyme, but see
Buys and Hilger 2003); however, the inflorescence structure is often obscured by short
internodes and absence of floral bracts. The calyx, considered part of the fruiting unit,
is accrescent, and continues to grow as the nutlets mature. The shape and fusion of the
calyx, as well as calyx vestiture and orientation when the plant is in fruit, varies among
taxa and is often used to distinguish them. Flowers are chasmogamous (opening and
capable of cross-pollination) in most taxa, but may be cleistogamous (not opening
and self-pollinated) in members of subgenera Cryptantha and Geocarya (modified
into lenticular structures in the last). The corolla limb (the expanded portion above
the corolla tube, containing the five lobes) ranges in size from less than 1 mm wide
(measured across from lobe to lobe) to up to 2 cm wide. Corollas are almost universally
white (a few species having yellow corollas), and are rotate to salverform. The corolla
tube is usually equal to the calyx in length and bears five epipetalous stamens. At
the outer corolla throat, the tissue is invaginated, resulting in five pinched or folded
regions, known as fornices. The fornices, which enclose the throat, can vary in size,
shape, and color, but are often yellow and possibly function as nectar guides. Corolla
lobes are ovate-oblong or suborbicular, ranging from widely spreading to ascending
in chasmogamous species. As is characteristic of the subfamily Boraginoideae, the
ovary is deeply four-lobed, with a gynobasic style arising from the base and center of
the ovary lobes. The ovary lobes mature into 1-seeded units with hardened pericarps,
termed nutlets. The number of nutlets that develop to maturity can vary from 1 to
4. The tissue between the ovary lobes, interpreted as either receptacular tissue or the
style base, continues to elongate and differentiate during fruit maturation. This tissue

Crossosoma 35(1), Spring-Summer 2009

3

is often called the gynobase and is that to which the nutlets are laterally attached at
maturity. The gynobase is generally narrowly-pyramidal in form, elongating during
fruit maturation. (Note that “fruit” is used here to include the nutlets and accompanying
calyx.) Nutlets are generally ovate to lanceolate in shape. The fruits of annual taxa are
generally deciduous, and the whole unit encased by the calyx will easily detach from the
inflorescence; however, in the perennial subgenus Oreocarya, the nutlets detach easily
but the calyx and pedicel remain firmly attached to the plant. Several annual species
in North American have fruits comprised of heteromorphic nutlets. Often, there are
three smaller, easily detached nutlets, and one larger nutlet that is strongly adnate to the
gynobase. The pericarp wall is variable in sculpturing and color. The attachment scar
is generally a shallow, triangular areole (generally not rimmed or elevated), which is
continuous with the ventral groove. When describing the ventral groove, morphologists
generally consider the groove and attachment scar as one feature on the nutlet; thus the
ventral groove is often described as being basally forked.

The genus name Cryptcmtha was first used by Johann Georg Christian Lehmann in
a seed catalogue for the Hamburg Botanical Garden. In 1836, Fischer and Meyer
used Lehmann’s proposed name Cryptantha in association with two South American
species, C. glomerata and C. microcapa, but with no formal genus diagnosis. In that
same year, however, the name Cryptantha was validly published in the General System
of Gardening and Botany by Don (1836), who provided the first formal diagnosis for
the genus in association with the same two South American species (Johnston 1925).
Thus, authorship of the genus should correctly be cited as Cryptantha Lehmann ex
G. Don. Greene (1887a,b,c) was the first botanist to carefully study the borages of
tribe Eritrichieae (within which Cryptantha has been classified), which are diverse
in western North America. He authored several revisions of Eritricheae members,
including Cryptantha , in a series of papers in the journal Pittonia. Green recognized the
importance of classifying the group based on characters other than floral appearance,
noticing the great variation in nutlet characteristics among members of this tribe. He
expanded the genus Cryptantha to include North American species that had been placed
in the genus Krynitzkia Fisch. & C.A. Mey. Greene’s circumscription of Cryptantha
included strictly annual taxa with deciduous fruits. Additionally, Greene recognized
several genera that are now treated as part of Cryptantha, including Oreocarya Greene,
Piptocalyx Torr. [=Greeneocharis Giirke & Harms], and Eremocarya Greene. The
next major revision came in the dissertation research of the Boraginaceae expert I.
M. Johnston in 1925. In this monographic treatment of the North American species
of Cryptantha , the genus was circumscribed to include Eremocarya and Piptocalyx
[ : =Greeneocharis ]. Each species was revised and clarified in this work, with each of
the 57 species placed into one of fifteen sections. Shortly after the publication of his
dissertation, Johnston spent several months in South America. These studies, with new
herbarium vouchers, led Johnston to conclude that the perennial Oreocarya species
should also be included in Cryptantha to form one large, homogenous genus. In
1927, Johnston published treatments of South American Boraginaceae, recognizing
Cryptantha as having three subgenera in South America: Krynitzkia, Geocarya,
and Eucryptantha [= Cryptantha ]. Also in 1927, Payson, who was working on a

4

Crossosoma 35(1), Spring-Summer 2009

Table 1 . Subgenera of Cryptantha recognized by Johnston (1927) and Payson (1927).

Subgenus Characters delineating subgenus

Eucryptantha

[=Cryptantha]

Geocarya

Krynitzkia
(incl. Eremocarya
and Greeneocharis)
Oreocarya

Plants with cleistogamous flowers. Cleistogamous flowers similar to
chasmogamic flowers, except for closed corolla. Cleistogamous flowers
located in axiles of leaves and often throughout the inflorescence.
Distributed in South America.

Plants with cleistogamous flowers. Basal cleistogamous flowers highly
specialized into lenticular structures. Distributed in South America.
Annual plants having only chasmogamous flowers. Distributed in North
and South America.

Perennial or biennial herbs, producing only chasmogamous flowers.
Restricted to North America.

monograph of Oreocarya, agreed with Johnston that Oreocarya should be included
in Cryptantha. Payson, following Johnston’s treatments, recognized four subgenera in
Cryptantha. Oreocarya, Krynitzkia, Geocarya and Eucryptantha [=Cryptantha\. (See
Table 1 for details of the characters used to circumscribe these subgenera, and their
distributions.) Since the 1920s, systematists have made revisions among species and
their relationships within Cryptantha, but the subgenera themselves have remained
largely unchanged. Higgins, another expert on the perennial taxa, published a revised
monograph of Oreocarya in 1971, and agreed with Johnston and Payson on the
inclusion of Oreocarya within Cryptantha. However, the classification of Oreocarya
has been contentious. For example, Abrams (1951) recognized Oreocarya as genus
separate from Cryptantha in his Illustrated Flora of the Pacific States: Washington,
Oregon and California. More recently, Weber (1987) also recognized Oreocarya at the
genus level, giving no specific reason for his circumscription, but arguing that genera
should consist only of species easily recognizable by suites of characters. However,
most recent classifications follow the treatments of Johnston, Payson, and Higgins in
circumscribing Cryptantha in the broad sense, as encompassing all four subgenera. For
Southern California, these include the treatments of Cryptantha in Munz (1974), The
Jepson Manual (W.A. Kelley and Wilken 1993), and the draft treatment for the second
edition of The Jepson Manual (R.B. Kelley in prep).

Hasenstab (2009), in a molecular phylogenetic analysis of Cryptantha and relatives,
concluded that the genus Cryptantha as currently defined appears to be paraphyletic,
warranting the resurrection of several genera, including Eremocarya, Greeneocharis,
and Oreocarya. Although we feel it premature to divide the genus for this article, the
key we have prepared reflects (in part) a more natural grouping of taxa.

Definition of the Southern California Region

As this article is on the Cryptantha of Southern California, we must define what we
mean by that geographic designation. We use Munz’s concept of Southern California,
as he recognized it in A Flora of Southern California (1974). He defined the northern
boundary of Southern California as “Point Conception, eastward along the crest of

Crossosoma 35(1), Spring-Summer 2009

5

White &
Inyo Mtns,

Tehachapi P
? Mountains * 7

Desert

Province

San Ui iimirlMM L’si.

PL < \tnc-t ptithii

Southwestern

Region

Hnrfu*-cv

linpt-riul -ft*.

Figure 1 . Regions and provinces from The Jepson Manual (Hickman 1993), corresponding to
Southern California, sensu Munz (1974). See this work for included subregions and districts.

6

Crossosoma 35(1), Spring-Summer 2009

the Santa Ynez mountains to the Mt. Pinos region in Ventura County, Fort Tejon in
Kern County, the Tehachipi and Piute mountains, then northward to Little Lake in Inyo
County and along the east slopes of the Inyo and White mountains to the Deep Springs
region.” Munz stated that the region does not include the northern part of Santa Barbara
County, the southern end of the San Joaquin Valley, nor any of the Owens Valley, as
these regions contain more of a northern element of the flora (Figure 1).

This boundary for “Southern California” contains components of three floristic
provinces designated in The Jepson Manual (Hickman 1993): 1) The entire Southwestern
California region (including the South Coast, Channel Islands, Transverse Ranges, and
Peninsular Ranges subregions) and the Tehachapi Mountain subregion of the California
Floristic province; 2) the entire Desert province (Sonoran and Mohave deserts,
including the Desert mountains); and 3) the White and Inyo Mountain subregion of
the Great Basin province. Although “Southern California” is often viewed as a socio-
political designation, the region as defined here actually corresponds to major floristic
provinces, regions, or subregions of California and thus has some scientific meaning.
With regard to county borders, Southern California includes all of San Diego, Imperial,
Los Angeles, Orange, Riverside, Ventura, and San Bernardino counties, the extreme
southern and southeastern portion of Santa Barbara County, the extreme southern,
south-central, and eastern parts of Kern County, all except the extreme northwestern
and a small northeastern part of Inyo County, and a southeastern swath of Mono county
(Figure 1).

Identification of Southern California Cryplantha

Cryptantha species and infraspecies are generally recognized as being difficult to
identify, even by professional botanists; the taxa are often being perceived to look very
similar to one another. Perhaps the major reason for this perception is that the main
feature used in identifying flowering plants - the flower itself - is generally both obscure
and uniform among Cryptantha taxa. The corolla, the primary cue of identification
both for humans and potential pollinators, is rarely used in Cryptantha identification,
except for some variation in size and (rarely) in color. And yet, it should be pointed
out that Cryptantha taxa vary tremendously in a great number of features, only some
of which are listed in Table 2. Many of these features collectively contribute to the
reflection of light, interpreted by our brains as a pattern. Thus, despite the variation
among individual plants due to differences in habitat and developmental stage, it is
possible, with practice, to recognize Cryptantha species at a glance (Figure 2A-D).
This wholistic, pattern recognition method is how most botanists and hobbyists identify
plants.

Until one develops a wholistic pattern recognition familiarity with Cryptantha, one
must rely instead on analytic methods, especially taxonomic keys and diagnostic
descriptions. Interestingly, relatively few of the characters listed in Table 2 are actually
used in keys. Among these major features in identification are plant duration, branching
pattern and position, bract presence, pedicel length, calyx length (in flower and fruit),

Crossosoma 35(1), Spring-Summer 2009

7

Table 2. Some characters used to distinguish between Cryptantha taxa, those in bold most
often used in taxonomic keys.

Plant duration

Calyx vestiture / trichome type

Stem type

Calyx posture

Stem vestiture / trichome type

Calyx orientation

Branch number / position

Corolla tube length

Internode length

Corolla limb width

Leaf position

Corolla color

Leaf size

Fomices color

Leaf shape

Gynobase size / shape

Leaf vestiture / trichome type

Style length

Inflorescence position

Nutlet number per fruit

Flower internode length

Nutlet size

Bract presence / morphology

Nutlet shape / base / margin / apex

Pedicel length

Nutlet sculpturing

Calyx length (flower vs. fruit)

Nutlet symmetry

Calyx / Sepal shape

Nutlet ventral groove shape

calyx/sepal shape, calyx vestiture/trichome type, and corolla limb width (Table 2).
However, variation in nutlet morphology is most important in Cryptantha identification,
including nutlet number (per fruit), size, shape, and sculpturing. (Why nutlet variation
is so great in this complex is a question of great interest regarding the evolution of the
group.) Thus, it is critical to collect Cryptantha specimens at a stage when at least some
mature fruits are present. Identification of immature plants can be quite problematic.

Observations of most of these key characteristics require the use of a good dissecting
microscope and basic dissecting tools. In addition, we have found two procedures to
be of value in identification of Cryptantha (and of related taxa, such as Amsinckia ,
Pectocarya, and Plagiobothrys). First, if nutlets adhere to and are difficult to remove
from the gynobase, boiling the fruits for 1-2 minutes will usually loosen them, allowing
them to be removed intact (Figure 2E). Second, to clearly see details of the fruits or
calyx, it is valuable to place them on a glass microscope slide covered with a strip of
double-stick tape and a label with herbarium voucher information (Figure 2F). Note
that in order to get an accurate count of nutlet number, care must be taken both to choose
fruits that are not too mature (i.e., before the nutlets or entire fruits have naturally fallen
away) and to avoid dislodging nutlets prematurely during dissection.

MATERIALS AND METHODS

Herbarium specimens or live plants (subsequently vouchered) were photographed
with a Nikon Coolpix 4400 or a Sony digital camera. Close-ups of vegetative parts,
flowers, or fruits were photographed either using a Wild M3Z dissecting scope with
a Nikon Coolpix 995 digital camera or using a Visionary Digital Imaging System
(consisting of an FX2 lighting system, Infinity Optics Long Distance Microscope,
SolMate trans-illuminator, Precision Linear Camera Controller, 10 megapixel digital
camera, and a devoted computer workstation with Helicon Focus software). Appendix

8

Crossosoma 35(1), Spring-Summer 2009

Figure 2. A-D: Species of Cryptantha , illustrating that many do look different and can be site-
identified. A: C. maritima. B: C. nevadensis. C: C. racemosa. D: C. utahensis. E: Boiling of
fruits for 1-2 minutes will loosen nutlets. F: Microsope slides with double-stick tape and labels
for observing Cryptantha nutlets.

Crossosoma 35(1), Spring-Summer 2009

9

1 lists herbarium voucher accessions for all Cryptantha images (except for a few field
photographs), by figure number and letter. Dichotomous keys were modified from pre-
existing keys from Munz and Keck (1959), Munz (1974), W.A. Kelley and Wilken
(1993), and R.B. Kelley (in prep). For current nomenclature and descriptions, we are
especially appreciative of an early version of R.B. Kelley’s treatment of the genus
for the upcoming second edition of The Jepson Manual. Information on bioregions
was obtained from R.B. Kelley {in prep) and from W.A. Kelley and Wilken (1993).
County and regional distributions were obtained from herbarium vouchers listed on the
Consortium of California Herbaria (2009; see Rosatti 2003) for the region of Southern
California as defined in this paper; however, identifications were not confirmed by
us, except in a few cases. Information on sensitive or rare taxa was obtained from
the California Native Plant Society Inventory (2009), cited in the text as a CNPS
listing, both in descriptions and the Conservation Status and Ecology of Southern
California Cryptantha section.

The following keys, plus multi-entry (polythetic) keys, numerous images, distribution
maps, and nomenclatural information, are accessible at our Cryptantha web site:
http://www.scLsdsu.edu/plants/cryptantha. Our aim is to provide a service for both
professionals and amateurs in the identification of this fascinating group of plants.
We provide an initial key to previously recognized groups (variously treated in the
past as subgenera or separate genera): Eremocarya, Greeneocharis, Krynitzkia,
and Oreocarya. We do not include detailed descriptions of taxa, but these will be
forthcoming in the second edition of The Jepson Manual (R.B. Kelley in prep). We cite
only the diagnostic features of each taxon, its general and county distribution, pertinent
ecological information, and conservation status, where applicable. Species diagnoses
and plates appear in the order presented in keys and not in alphabetical order so as to
more clearly illustrate and compare diagnostic features. For ease of use, each major
group and individual taxa are numbered in the keys and descriptions. Technical terms
that we use include dorsal, for the back side of a nutlet, i.e., the side facing away from
the gynobase, and ventral, for the inner side of a nutlet, facing the gynobase. (See
Cryptantha web site: Keys for a complete list of terminology and their definitions.)
Corolla width is always measured from lobe to lobe of the corolla limb, the expanded
portion at the apex of the tube or throat. We do not include distribution maps (due to
space concerns), but encourage all to do a specimen search for selected taxa on the
Consortium of California Herbaria.

10

Crossosoma 35(1), Spring-Summer 2009

KEYS TO CRYPTANTHA OF SOUTHERN CALIFORNIA

The 49 species and 56 taxa (including varieties) of Cryptantha occurring in Southern
California are listed in Table 3. The initial key, below, segregates taxa into “groups”
that correspond to previously recognized genera or subgenera. The Krynitzkia group is
further divided into two keys: one for smooth nutlets and one for rough nutlets.

Table 3. The 49 species and 56 taxa of Cryptantha in Southern California. Taxonomy after R. B.
Kelley, in prep. *= Eremocarya ; **= Greeneocharis: ***= Oreocarya ; all others “Krynitzkia” .
CNPS Inventory listings in bold.

C affinis

C. gracilis

C. nubigena ***

C. angustifolia

C. hoffinannii ***

C. oxygona

C. barbigera var. barbigera

C. holoptera 4.3

C. pterocarya var. cycloptera

C. barbigera var. fergusoniae

C. humilis ***

C. pterocarya var. pterocarya

C. cinerea var. abortiva ***

C. inaequata

C. pterocarya var. purpusii

C. circumscissa var. circumscissa **

C. intermedia var. intermedia

C. racemosa

C. clevelandii var. clevelandii

C. leiocarpa

C. recurvata

C. clevelandii var. florosa

C. lepida *

C. roosiorum *** 1B.2

C. clokeyi 1B.1

C. maritima var. maritima

C. scoparia 4.3

C. confertiflora ***

C. micrantha *

C. similis **

C. corollata

C. micromeres

C. simulans

C. costata 4.3

C. microstachys

C. sparsiflora

C. decipiens

C. mohavensis

C. torreyana

C. dnmetorum

C. muricata var. denticulata

C. traskiae 1B.2

C. echinella

C. muricata var. jonesii

C. tumulosa *** 4.3

C. flaccida

C. muricata var. muricata

C. utahensis

C. flavoculata * * *

C. nemaclada

C. virginensis ***

C. gander i 1B.1

C. nevadensis var. nevadensis

C. watsonii

C. glomeriflora 4.3

C. nevadensis var. rigida

Key to Cryptantha Groups

1. Plants annual, gen wider than tall, often rounded to cushion-like; taproot red or purple when dry
(often staining herbarium paper); flower bracts present

2. Calyx basally fused, tubular, tube circumscissile in fruit Group 1: Greeneocharis , p. 11

2’ Calyx of distinct sepals, intact in fruit Group 2: Eremocarya , p. 13

1’ Plants annual, biennial, or perennial, gen taller than wide, rarely rounded to cushion-like;
taproot usually not red or purple when dry; flower bracts absent or rarely present

3. Plants biennial or perennial; vegetative leaves basal or tufted; nutlets smooth, rugose, or
scabrous, apically broadly rounded to obtuse; ventral groove apex well below nutlet apex
Group 3: Oreocarya , p. 13, 15

3’ Plants annual or rarely perennial; vegetative leaves cauline; nutlets smooth, papillate,
tuberculate, or muricate, apically narrowly acute to acuminate; ventral groove apex almost
to nutlet apex Group 4: Krynitzkia , p. 20

Crossosoma 35(1), Spring-Summer 2009

11

Group 1: Greeneocharis [ Piptocalyx \

The two species of Greeneocharis (named after Edward Lee Greene, 1843-1915) are
readily distinguished by the combination of red to purple-pigmented roots, bracteate
flowers, and a calyx that is basally synsepalous (sepals fused), splitting transversely at
fruit maturity, a condition termed circumscissile dehiscence. Interestingly, the only
other close relatives with such a calyx dehiscence are two species of Plagiobothrys: P.
arizonicus and P. nothofulvus. See Figures 29A,B for comparative nutlet images in the
Greeneocharis group.

1. Corolla 1-2 mm wide, fomices tiny, white 1. C. circumscissa var. circumscissa

V Corolla 3-6 mm wide, fomices evident, yellow 2. C. similis

1. Cryptantha circumscissa var. circumscissa (Hook. & Arn.) I.M. Johnst.

(Figure 3A-E, p.12)

Common name: Circumscissile cryptantha. Distribution in Southern California:

Southwestern California region (eastern South Coast, Transverse Ranges, and
eastern Peninsular Ranges), Desert province, White and Inyo mountains; Imperial,
Inyo, Kern, Los Angeles, Mono, Riverside, Santa Barbara, San Bernardino, San
Diego, and Ventura counties. Conservation Status: None. Diagnosis: Plants (at
maturity) generally wider than tall with red-purple roots and bracteate flowers,
distinguished from the Eremocarya group in having circumscissile calyces and
from C. similis by the smaller (1-2 mm wide) corolla. Note: C. circumscissa var.
rosulata, the ground rose cryptantha (CNPS IB. 2 listing), has a limited distribution
in subalpine habitats of the Sierra Nevada, Inyo and Tulare Cos., outside Southern
California.

2. Cryptantha similis K. Matthew & PH. Raven (Figure 3F-J, p.12)

Common name: Dome cryptantha. Distribution in Southern California:

Southwestern California region (northern San Gabriel and western San Bernardino
mountains), southwestern Mohave Desert region; Kern, Los Angeles, and San
Bernardino counties. Conservation Status: None. Diagnosis: Plants (at maturity)
generally wider than tall with red-purple roots and bracteate flowers, distinguished
from the Eremocarya group in having circumscissile calyces and from C.
circumscissa by the larger (3-6 mm wide) corolla. [C. circumscissa (Hook. & Am.)
I.M. Johnst. in part , sensu W.A. Kelley and Wilken 1993]

These two taxa are distinguished only by corolla size and color of fomices. The range
of Cryptantha similis is much more restricted, but appears to overlap, with that of
C. circumscissa.

12

Crossosoma 35(1), Spring-Summer 2009

calyx

circumscissile

floral

■bract-

vircumscissai

xirjcumseissa.

circumscissa

calyx

circumscissile

\ : V

i.

■

5 mm

fiTirafl

Figure 3. A-E: Cryptantha circumscissa. A: Whole plant, showing small height relative to
width. B-C: Inflorescence and flowers. Note bracts and small corolla. D: Circumscissal calyx.
E: Nutlet, dorsal (left), ventral (middle), and lateral (right) views. F-J: C. similis. F: Whole
plant. G: Flowering shoot. Note wide (3-6 mm) corolla and floral bracts. H: Circumscissal calyx.
I: Stem close-up, showing appressed and spreading trichomes. J: Nutlet, dorsal (left) and ventral
(right) views.

Crossosoma 35(1), Spring-Summer 2009

13

Group 2: Eremocarya

The two species of Eremocarya (from eremo , desert + carya, fruit) are distinguished
by the combination of red to purple-pigmented roots, bracteate flowers, and an
aposepalous calyx (sepals not fused). Thus, it is the latter feature that distinguishes this
group from Greeneocharis. See Figures 29C,D for comparative nutlet images in the

Eremocarya group.

1. Corolla limb 1-3.5 mm wide 3. C. lepida

V Corolla limb 0.5-1 .2 mm wide 4.C. micrantha

3. Cryptantha lepida (A. Gray) I.M. Johnst. (Figure 4A-E, p.14)

Common name: Elegant cryptantha. Distribution in Southern California:

Southwestern California region (Peninsular and Transverse Ranges), Desert
province; Imperial, Kem, Los Angeles, Riverside, San Bernardino, San Diego
counties. Conservation Status: None. Diagnosis: Plants (at maturity) generally
wider than tall with red-purple roots and bracteate flowers, distinguished from the
Greeneocharis group in having non-circumscissile calyces and from C. micrantha
in having larger (1-3.5 mm wide) corollas. [C. micrantha (Torrey) I.M. Johnst. in
part , sensu W.A. Kelley and Wilken 1993]

4. Cryptantha micrantha (Torrey) I.M. Johnst. (Figure 4F-J, p.14)

Common name: Purple Root cryptantha. Distribution in Southern California:

mostly Desert province; Imperial, Inyo, Los Angeles, Riverside, San Bernardino,
San Diego counties. Conservation Status: None. Diagnosis: Plants (at maturity)
generally wider than tall with red-purple roots and bracteate flowers, distinguished
from the Greeneocharis group in having non-circumscissile calyces and from C.
lepida in having smaller (0.5- 1.2 mm wide) corollas.

Note that some overlap occurs in the overall corolla width of these two species.
However, within a plant, most flowers fit the range of their respective species. The
two taxa also seem to occupy different habits, C. lepida in montane and desert
transition regions, occasionally cismontane washes, and C. micrantha in lower
elevation desert regions, generally flats and alluvial slopes, often sandy soils, with
creosote bush scrub.

Group 3: Oreocarya

The species of Oreocarya ( oreo , mountain + carya , fruit) are distinctive in being
biennial or perennial taxa with a generally well formed and persistent rosette of leaves.
Many of these taxa occur at high elevations. In addition, the nutlets in this group tend
to be larger than other Cryptantha species, usually with characteristic rounded apices
and ventral grooves that do not extend to the nutlet apex. See Figure 26 for comparative
nutlet images in the Oreocarya group.

14

Crossosoma 35(1), Spring-Summer 2009

Figure 4. A-E: Cryptanthalepida. A: Whole plant. B: Inflorescence; note floral bracts. CiFlower,
showing larger (1-3.5 mm wide) corolla. D: Fruit, with 4 nutlets removed; note unfused sepals
(calyx aposepalous). E: Nutlet, dorsal, ventral, and lateral views (left to right). F-J: C. micrantha.
F: Whole plant, wider than tall. G: Inflorescence, with floral bracts. H: Flower, showing small
(0.5-1 mm wide) corolla. I: Fruit, with unfused sepals. J: Nutlets, two forms: rough (left, dorsal
and ventral views) and smooth (right, dorsal and ventral views).

Crossosoma 35(1), Spring-Summer 2009

15

Key to Oreocarya Group

1. Corolla tube > sepals

2. Corolla and fomices yellow; nutlet back smooth, groove closed, ventral groove margins ±

overlapping 5. C. confertiflora

T Corolla white, fomices light yellow; nutlet back rough, groove open, narrowed near

middle, ventral groove margins elevated 6. C. flavoculata

V Corolla tube = or < sepals

3. Stems prostrate to ascending; nutlet dorsal side strongly convexly curved, surface smooth;

groove closed, margins generally overlapping 7. C. cinerea var. abortiva

3’ Stems ± erect (generally cespitose in C. humilis ); nutlet back relatively straight, surface
smooth or rough; groove closed or open, margins not overlapping

4. Stems 1-2 cm tall, < or = basal leaves; basal leaves generally < 1 cm .... 8. C. roosiorum
4’ Stems > 2 cm tall, > basal leaves; basal leaves generally » 1 cm

5. Nutlet lanceolate to lance-ovate, inner surface (between groove and margin) generally

smooth, shiny 9. C. nubigena

5’ Nutlet ovate, inner surface generally rough or wrinkled

6. Edge of nutlet groove not elevated; nutlet sculpturing finely rugose-tuberculate on

dorsal and ventral surfaces 10. C. humilis

6’ Edge of nutlet groove elevated; nutlet sculpturing roughly rugose

7. Pedicel 0.5-2 mm in fruit; calyx lobes 4.5-8 mm in fruit; trichomes on stem and

leaves white 11. C. hoffmannii

T Pedicel 2-6 mm in fruit; calyx lobes 7-11 mm in fruit; trichomes on stem and
leaves yellow to brown

8. Biennial or short-lived perennial herbs; caudex not woody; nutlets

prominently dorsally ridged or keeled 12. C. virginensis

8’ Long-lived perennial herbs; caudex woody; nutlets not prominently keeled,
dorsal ridge rounded 13. C. tumuiosa

5. Cryptantha confertiflora (Greene) Pay son (Figure 5A-B, p.16)

Common name: Yellow-flowered cat’s eye. Distribution in Southern
California: Southwestern California region (San Bernardino Mountains), Desert
mountains; Inyo, Kern, Mono, San Bernardino counties. Conservation Status:
None. Diagnosis: Corolla yellow, tube exceeding calyx in length; nutlets smooth
and shiny with closed ventral groove with overlapping margins.

6. Cryptantha flavoculata (A. Nelson) Payson (Figure5C-D, p.16)

Common name: Yellow-eyed cat’s eye. Distribution in Southern California:

Desert mountains; Inyo, Mono, San Bernardino counties. Conservation Status:
None. Diagnosis: Corolla white with prominent, bright yellow fomices, tube
exceeding calyx in length; nutlets with rough sculpturing and open ventral groove
with elevated margins.

7. Cryptantha cinerea var. abortiva (Greene) Cronquist (Figure 5E-H, p.16)

Common name: Bow-nut cat’s eye. Distribution in Southern California: San

Bernardino, Desert mountains; Inyo, Los Angeles, Mono, San Bernardino counties.
Conservation Status: None. Diagnosis: Stems prostrate to ascending; nutlets
strongly convex-curved dorsally, smooth and shiny with a closed nutlet groove. [C.
jamesii (Torrey) Payson var. abortiva (Greene) Payson]

16

Crossosoma 35(1), Spring-Summer 2009

Figure 5. A-B: Cryptantha confertiflora. A: Inflorescence, showing corolla tube longer than
calyx. B: Nutlet, dorsal (left) and ventral (right) views, showing smooth surface and closed
groove. C-D: C. flavoculata. C. Inflorescence, showing long corolla tube. D: Nutlet, dorsal (left)
and ventral (right) views, showing rough surface and open groove. E-H: C. cinerea var. abortiva.
E: Plant, showing prostrate to ascending stem habit. F: Nutlet, side view, showing strongly
curved dorsal surface. G: Nutlet, dorsal view. H: Nutlet, ventral view. Note: images at A, C, and
E courtesy of Gary Monroe.

Crossosoma 35(1), Spring-Summer 2009

17

8. Cryptantha roosiorum Munz (Figure 6A-B, p.18)

Common name: Bristlecone cat’s eye. Distribution in Southern California:

Inyo Mountains; Inyo County. Conservation Status: CNPS List 1B.2. Diagnosis:
Stems short, often equal in length with the basal leaves and generally no greater
than 2 cm tall; basal leaves no greater than 1 cm long. It has a very restricted
distribution in high elevation limestone soils of bristle-cone pine and limber pine
forests.

9. Cryptantha nubigena (Greene) Payson (Figure 6C-D, p.18)

Common name: Sierra cat’s eye. Distribution in Southern California: White
and Inyo mountains, Desert mountains; Inyo, Mono, San Bernardino counties.
Conservation Status: None. Diagnosis: Stems elongate, much longer than leaves;
nutlets lanceolate to lance-ovate with smooth, shiny dorsal and ventral surfaces.

10. Cryptantha humilis (Greene) Payson (Figure 6E-F, p.18)

Common name: Low cat’s eye. Distribution in Southern California: northern
Desert mountains; Inyo, Mono counties. Conservation Status: None. Diagnosis:
Stems elongate, much longer than leaves; nutlets ovate, sculpturing finely rugose
on dorsal and ventral surfaces, margin of the ventral groove non-elevated.

11. Cryptantha hoffmannii I.M. Johnston (Figure 7A-D, p.19)

Common name: White Mountain cat’s eye. Distribution in Southern California:

White and Inyo mountains, northern Desert mountains; Inyo, Mono, San
Bernardino counties. Conservation Status: None. Diagnosis: Biennial or short-
lived perennial; pedicel 0.5-2mm in fruit; calyx 4.5-8 mm long; nutlet coarsely
rugose, ventral groove margin elevated. It is distinguished from the closely
associated species C. virginensis and C. tumulosa by the snowy-white trichomes
and the smaller pedicel and fruiting calyx. It is distinguished from C. humilis by the
coarsely rugose sculpturing on the nutlet and elevated margin of the nutlet groove.
[C. virginensis (M.E. Jones) Payson in part, sensu W.A. Kelley and Wilken 1993]

12. Cryptantha virginensis (M E. Jones) Payson (Figure 7E-G, p.19)

Common name: Virgin River cat’s eye. Distribution in Southern California:

White and Inyo mountains, northern and eastern Mojave Desert mountains;
Inyo, Mono, San Bernardino counties. Conservation Status: None. Diagnosis:
Biennial or short lived perennial; caudex slender and short-lived; pedicel 2-5 mm
long in fruit; calyx 7-11 mm long; nutlet roughly rugose, dorsally carinate, ventral
groove margin elevated. It differs from C. hoffmannii in its tawny, yellow-brown
trichomes, the longer pedicel and calyx, and the dorsally carinate (keeled) nutlet.

13. Cryptantha tumulosa (Payson) Payson (Figure 7H-I, p.19)

Common name: New York Mountain cat’s eye. Distribution in Southern
California: eastern Mojave Desert, northern and eastern Desert mountains; Inyo,
Mono, San Bernardino counties. Conservation Status: CNPS List 4.3. Diagnosis:
Perennial; caudex woody; pedicel 3-6 mm long in fruit, calyx 7-10 mm long in
fruit, nutlet roughly rugose, not carinate, ventral groove margin elevated. It is

18

Crossosoma 35(1), Spring-Summer 2009

leaves
< 1 cm

stems

1-2 cm

mosiomm

Inutl ets landeolate to ■
j^ance-ovate; ventra^^M
surface smooth, sl^^»

C. nubigena

edge of groove ‘
not elevated

humili

Figure 6. A-B: Cryptantha roosiorum. A: Whole plant, showing short stature and small leaves.
B: Nutlet, dorsal (left) and ventral (right) views. C-D: C. nubigena. C: Whole plant. D: Nutlet,
dorsal (left) and ventral (right) views. Note smooth, shiny ventral surface. E-F: C. humilis.
E: Whole plant. F: Nutlet, dorsal (left) and ventral (right) views. Note finely rugose surface, edge
of ventral groove not appreciably elevated.

Crossosoma 35(1), Spring-Summer 2009

19

Figure 7. A-D: Cryptantha hoffmannii. A: Whole plant. B: Stem trichomes, white in color.
C: Calyx in fruit, 4.5-8 mm long. D: Nutlet, dorsal (left) and ventral (right) views. Note elevated
ventral groove edge. E-G: C. virginensis. E: Whole plant. F: Stem trichomes, yellow to brown
in color. G: Nutlet, dorsal (left) and ventral (right) views. Note dorsal keel. H-l: C. tumulosa.
H: Whole plant. I: Nutlet, dorsal (left) and ventral (right) views. Note absence of dorsal keel.

20

Crossosoma 35(1), Spring-Summer 2009

distinguished from closely associated C. hoffmannii, by its tawny, yellow-brown
trichomes, and the longer (in fruit) pedicel and calyx. It differs from C. virginensis
in having a prominently woody caudex and nutlets with a rounded dorsal ridge.

Group 4: Krynitzkia

The species of Krynitzkia (origin unknown, presumably named as a commemorative),
as circumscribed by Johnston (1927) and Payson (1927), are almost certainly an
unnatural, paraphyletic group (Hasenstab 2009). Almost all members are annuals (very
rarely perennial) with mostly cauline leaves. The nutlets in this group are quite variable
in sculpturing, number, and heteromorphism, but all have an acute to narrowly acute
apex and a ventral groove that extends almost to the apex. See Figures 27, 28, 29E-H,
and 30 for comparative nutlet images in the Krynitzkia group. For ease of identification,
Krynitzkia is divided into two groups, based on general nutlet sculpturing.

Key to Groups of Krynitzkia

1. All nutlets of fruit smooth and shiny Krynitzkia Smooth Nutlet Group, below

V At least one nutlet of fruit rough: papillate, tuberculate, and/or muricate

Krynitzkia Rough Nutlet Group, p. 29

Krynitzkia - Smooth Nutlet Group (See Figure 27 for comparative nutlet images)
[Note: an asterisk indicates that taxon also appears in the Krynitzkia Rough Nutlet key.]

1. Nutlet asymmetric, ventral groove off-center from nutlet body

2. Nutlet ventral groove strongly off-center, appearing marginal; nutlets 4 per fruit; flowers in

elongate inflorescence units 14. C. afftnis

2’ Nutlet attachment scar edges slightly off-center, nearer one margin than the other; nutlet 1
per fruit; flowers in globose inflorescence units 15. C. glomeriflora

V Nutlet symmetric, ventral groove central

3. Some calyx trichomes apically hooked, sometimes basally encrusted with exudate; nutlet 1
per fruit

4. Nutlet lance-ovate with swollen base, rounded margin, and long acuminate (caudate)

apex; corolla limb 1-5(6) mm wide; calyx trichomes usually basally encrusted with
exudate; common, widespread 16. C. flaccida

4’ Nutlet compressed ovate with dorsally flattened base, sharp angled margin, and ± acute
apex; corolla limb 0. 5-1(1. 5) mm wide; calyx trichomes not basally encrusted with

exudate; uncommon 17. C. sparsiflora

3’ Calyx trichomes ± straight, rarely wavy, either erect, spreading, or reflexed, not apically
hooked nor basally encrusted with exudate; nutlets 1-4 per fruit

5. Calyx sepal tips with reflexed bristles; corolla limb 0.5-1 mm wide .... 18. C. nemaclada
5’ Calyx sepal tips with bristles mostly spreading or ascending; corolla limb 0.5-7.5 mm

wide

6. Sepals 6-10 mm in fruit; longest sepal bristles 3-4 mm in fruit 19. C. ganderi

6 ’ Sepals 1-6 mm in fruit; longest sepal bristles < 3 mm in fruit
7. Flowers gen subtended by bracts

8. Stems gen prostrate to decumbent, greenish when in fruit; nutlets 3-4 per fruit

(all smooth) 20. C. leiocarpa

8’ Stems gen erect to ascending, reddish brown when in fruit; nutlet 1 per fruit and
smooth (or 2 per fruit, 1 smooth, 1 rough) * 21. C. maritima

Crossosoma 35(1), Spring-Summer 2009

21

7’ Flower bracts absent

9. Calyx obconic at base; calyx trichomes dense, straight, ascending, relatively

soft, sepal midvein hairs slightly longer, but not bristly 22. C. gracilis

9’ Calyx rounded at base, with both appressed, soft trichomes and ascending to
spreading, stiff, bristle-like trichomes

10. Sepals < 1 mm in ft, 1-2 mm in fruit 23. C. microstachys

10’ Sepals > 1 mm in ft, >3 mm in fruit

11. Nutlets 1-1.5 mm long, 1-4 per fruit C. cleveiandii

12. Pedicels 0-0.5 mm, becoming erect, ± appressed to stem in fruit; calyx
(2. 5)3-3. 5 mm in fruit, narrow at base, sepals linear in fruit; corolla
limb 1-2.5 mm wide; corolla fornices pale yellow; nutlets 1-2 per fruit
24. C. cleveiandii var. cleveiandii

12’ Pedicels 0.5-1 mm, ± inclined, not appressed to stem in fruit; calyx
3 — 4(4.5) mm in fruit, ± wide at base, sepals ovate in fruit; corolla limb
(2)2. 5-5 mm wide; corolla fornices yellow; nutlets 3-4(1 ) per fruit

25. C. cleveiandii var. florosa

11’ Nutlets 1.5-2 mm long, 4 per fruit

13. Nutlet margin gen rounded throughout, sometimes angled below

middle 26. C. torreyana

13’ Nutlet margin angled (obtuse in cross-section), esp near tip

14. Corolla limb (4)5-7. 5 mm wide; nutlets ovate to lance-ovate

27. C. mohavensis

14’ Corolla limb 2-3 mm wide; nutlets lanceolate to lance-ovate

28. C. watsonii

14. Cryptantha affinis (A. Gray) Greene (Figure 8A, p.22)

Common name: Side-grooved cryptantha. Distribution in Southern California:

Southwestern California region (Transverse and Peninsular Ranges); Kern, Los
Angeles, Mono, San Bernardino, San Diego, and Ventura counties. Conservation
Status: None. Diagnosis: Nutlets 4 per fruit with a very eccentric ventral groove
appearing to be marginal; inflorescences with straight clusters of flowers.

15. Cryptantha glomeriflora Greene (Figure 8B,C, p.22)

Common name: Trukee cryptantha. Distribution in Southern California: White
mountains; Mono County. Conservation Status: CNPS List 4.3. Diagnosis: Nutlet
1 per fruit with a slightly off-center ventral groove; inflorescence with globose
clusters of flowers.

16. Cryptantha flaccida (Lehmann) Greene (Figure 8D-E, p.22)

Common name: Weak-stemmed cryptantha. Distribution in Southern
California: Southwestern California region (western Transverse Ranges, eastern
San Bernardino mountains, South Coast, Peninsular Ranges), Tehachapi mountains;
Inyo, Kem, Los Angeles, Riverside, Santa Barbara, San Diego, Ventura counties.
Conservation Status: none. Diagnosis: Calyx apically hooked, trichomes usually
basally encrusted with exudate; nutlet 1 per fruit, lance-ovoid with a swollen base,
rounded margin, and long acuminate apex.

22

Crossosoma 35(1), Spring-Summer 2009

Figure 8. A: Cryptantha affinis, nutlet, ventral view, showing strongly off-center ventral groove.
B-C: C. glomeriflora. B: Nutlet, ventral view, showing slightly off-center (asymmetric) ventral
groove. C: Inflorescence unit, a tightly clustered, globose circinate cyme. D-E: C. flaccida.
D: Sepal, showing hooked (uncinate) trichomes (arrow). E: Nutlet, dorsal, ventral, and lateral
views (left to right). Note lanceolate shape, globose base, rounded margin, and elongate,
acuminate (caudate) apex. F-G: C. sparsiflora. F: Sepal, showing hooked (uncinate) trichomes
(arrow). G: Nutlet, dorsal, ventral, and lateral views (left to right). Note ovate shape, dorsally
flattened base, sharp-angled margin, and short, acute apex.

Crossosoma 35(1), Spring-Summer 2009

23

17. Cryptantha sparsiflora (Greene) Greene (Figure 8F-G, p.22)

Common name: Few-flowered cryptantha. Distribution in Southern California:

Southwestern California region (South Coast), Tehachapi mountains; Kern, Ventura
counties. Conservation Status: none. Diagnosis: Calyx trichomes apically hooked,
not basally encrusted with exudate; nutlet 1 per fruit, ovate with dorsally flattened
base, sharp-angled margin, and acute apex. [ C.flaccida (Lehmann) Greene in part ,
sensu W.A. Kelley and Wilken 1993]

18. Cryptantha nemaclada Greene (Figure 9A-B, p.24)

Common name: Colusa cryptantha. Distribution in Southern California:

Southwestern California region (western Transverse Ranges), Tehachapi mountains;
Kem, Santa Barbara, Ventura counties. Conservation Status: none. Diagnosis:
Sepal apices with reflexed trichomes; nutlets 1-2 (4) per fruit.

19. Cryptantha ganderi I.M. Johnst. (Figure 9C-D, p.24)

Common name: Gander’s cryptantha. Distribution in Southern California:

western Sonoran Desert; San Diego County (Borrego Valley). Conservation
Status: CNPS List 1B.1. Diagnosis: Sepals relatively long (6-10 mm) with long
(3-4 mm) sepal bristles; nutlets large (2-2.5 mm long).

20. Cryptantha leiocarpa (Fisher & C. A. Meyer) Greene (Figure 9E, 9G, p.24)
Common name: Beach cryptantha. Distribution in Southern California:
Southwestern California region (northern South Coast); Los Angeles, Orange,
Santa Barbara, and Ventura counties. Conservation Status: none. Diagnosis: Plant
decumbent; stems greenish at maturity; flowers bracteate; nutlet 1 per fruit.

21. Cryptantha maritima (Greene) Greene var. maritima

(Figure 9F, 9H, 19A-C, p.24, 40)
Common name: Maritime cryptantha. Distribution in Southern California:

Southwestern California region (South Coast, Channel Islands, Transverse Ranges,
Peninsular Ranges), Desert province; Imperial, Inyo, Los Angeles, Riverside, Santa
Barbara, San Bernardino, San Diego, and Ventura counties. Conservation Status:
none. Diagnosis: Plants erect; stems reddish at maturity; flowers bracteate; nutlets
either 1 per fruit and smooth or 2 per fruit with one smooth and one tuberculate.
[C. maritima var. pilosa l.M. Johnston is purported to differ from C. maritima var.
m. in having dense tufts of soft (pilose) trichomes arising from the calyx, but this
feature forms a grade is not diagnostic, sensu R.A. Kelly (in prep ); C. maritima var.
cedrocensis (Greene) I.M. Johnston, endemic to Cedros Island, Baja California,
Mexico, differs in having 4 nutlets (Wiggins 1980)]

22. Cryptantha gracilis Osterh. (Figure 10A-B, p.26)

Common name: Slender cryptantha. Distribution in Southern California: Desert
province (eastern Mojave Desert and mountains), White and Inyo mountains; Inyo,
Mono, San Bernardino counties. Conservation Status: none. Diagnosis: Calyx
obconical, with ascending, soft trichomes; nutlet 1 (2,3) per fruit, 3-sided.

24

Crossosoma 35(1), Spring-Summer 2009

Figure 9. A-B: Cryptantha nemaclada. A: Calyx, showing short, reflexed bristles near apex.
B: Nutlet, ventral (left) and dorsal (right) views. C-D: C. ganderi. C: Calyx. Note long length.
D: Nutlet, dorsal view. E&G: C. leiocarpa. E: Inflorescence branch, showing bracteate flowers.
G : Plant from field, showing decumbent branching. Inset: Nutlet, dorsal view. F&H : C. maritima.
F: Inflorescence, showing bracteate flowers. H: Plant, showing erect to ascending branches.
Inset: Nutlet, dorsal view.

Crossosoma 35(1), Spring-Summer 2009

25

23. Cryptantha microstachys (A. Gray) Greene (Figure 10C-E, p.26)

Common name: Tejon cryptantha. Distribution in Southern California:

Southwestern California region (Transverse Ranges, Peninsular Ranges, South
Coast, Santa Catalina Island); Kem, Los Angeles, Orange, Riverside, Santa
Barbara, San Bernardino, San Diego, and Ventura counties. Conservation Status:
none. Diagnosis: Calyx small, short (< 1 mm in fl, 1-2 mm in fr); nutlet usually
solitary.

24. Cryptantha clevelandii Greene var. clevelandii (Figure 10F-H, p.26)

Common name: Cleveland’s cryptantha. Distribution in Southern California:

Southwestern California region (South Coast); Los Angeles, Santa Barbara, San
Diego counties. Conservation Status: none. Diagnosis: Flowers ebracteate;
pedicels absent to short (0-0.5 mm), becoming appressed to the stem in fruit; calyx
basally narrowed with straight, non-reflexed, soft and bristle-like trichomes, the
calyx (2. 5)3-3. 5 mm in fruit; corolla 1-2.5 mm wide; nutlets 1-2 per fruit (1-1 .5 mm
long).

25. Cryptantha clevelandii Greene var florosa I.M. Johnst. (Figure 11A-D, p.27)
Common name: Coastal cryptantha. Distribution in Southern California:

Southwestern California region (South Coast, Channel Islands); Kern, Los
Angeles, Orange, Riverside, Santa Barbara, San Bernardino, San Diego, and
Ventura counties. Conservation Status: none. Diagnosis: This variety differs in
having longer (0.5-1 mm) pedicels that become inclined (not appressed) to the stem
in fruit, a longer (3-4 mm long), basally wide calyx (in fruit), a wider (2.5-5 mm)
corolla limb, and nutlets 3-4(1) per fruit. [C. clevelandii Greene in part , sensu W.A.
Kelley and Wilken 1993]

26. Cryptantha torreyana (A. Gray) Greene (Figure 11E-J, p.27)

Common name: Torrey’s cryptantha. Distribution in Southern California:

Southwestern California region (western Transverse Ranges), White mountains;
Inyo, Kern, and Ventura counties. Conservation Status: none. Diagnosis: Flowers
ebracteate; calyx with a rounded base and straight, non-reflexed trichomes; nutlets
symmetrical (1.5-2 mm long), 4 per fruit, with rounded margins.

27. Cryptantha mohavensis (Greene) Greene (Figure 12A-B, p.28)

Common name: Mohave cryptantha. Distribution in Southern California:

Tehachapi mountains, western Mohave Desert, edge of desert only, mostly found
along the desert slope of mountains; Inyo, Kern, Los Angeles, San Bernardino
counties. Conservation Status: none. Diagnosis: Flowers ebracteate; calyx with
a rounded base (1-6 mm long), trichomes straight, non-reflexed; corolla limb
relatively large (4-7.5 mm wide); nutlets symmetrical, ovate to lance-ovate (ca. 2
mm long), 4 per fruit, with distinctly angled margins.

26

Crossosoma 35(1), Spring-Summer 2009

Ja-jfmwrostach^

Wn microstachys

yar-./c/ eve/anaii

nutlets
-1.5 mm
long

C. gracilis

sepals
1-2 mm
in fr

sepals

1mm

in 11

C. microstachys

corolla
1-2.5 mm
wide

fruits

appressed

fliTlTTil

C. clevelandii var. clevelandii

var. clevelandii

Figure 10. A-B: Cryptantha gracilis. A: Calyx, showing characteristic obconic base
and long, soft trichomes. B: Nutlet, dorsal, ventral, and lateral views (left to right).
C-E: C. microstachys. C: Flower. Note short calyx. D: Fruit, also with short calyx.
E: Nutlet, dorsal (left) and ventral (right) views. F-H: C. clevelandii var. clevelandii. F: Flower,
showing small corolla. G: Fruit. Note appressed orientation. H: Nutlet, dorsal (left) and ventral
(right) views.

Crossosoma 35(1), Spring-Summer 2009

27

nutletsjf

Rffrait

r @fc/eve/andii
var. florosa

C. Jorreyana<

E C. torreyana

F W^VPrreydna

C. torreyana

B. fruits
^inclined,
nut appressed

torreyanm I C. torreyana

margin «
rounded A jw

nutlet
triangular
in cross-section

margin

rounded

Figure 11. A-D: Cryptantha clevelandii var. ft or os a A: Fruit, showing inclined orientation.
B: Corolla, relatively wide in size. C: Fruit, showing 4 nutlets. D: Nutlet, dorsal (left) and ventral
(right) views. E-J: C. torreyana. E: Whole plants from herbarium specimen. F: Close-up of leaf
(left) and stem (right). G: Young inflorescence. H: Flower. I: Fruit. J: Nutlet, dorsal, ventral,
lateral, and cross-sectional views (left to right). Note triangular shape and rounded margins.

28

Crossosoma 35(1), Spring-Summer 2009

Figure 12. A-B: Cryptantha mohavensis. A: Nutlet, dorsal, ventral, lateral, and
cross-sectional views (left to right). Note ovate shape and rounded ventral face (far
right) and sharp-angled margins. B: Corolla, with wide limb. C-D: C. watsonii.
C: Corolla, with short limb. D: Nutlet, dorsal, ventral, lateral, and cross-sectional views (left to
right). Note lanceolate shape and sharp-angled margins.

Crossosoma 35(1), Spring-Summer 2009

29

28. Cryptantha watsonii (A. Gray) Greene (Figure 12C-D, p.28)

Common name: Watson’s cryptantha. Distribution in Southern California: White
and Inyo mountains; Inyo, Mono counties. Conservation Status: none. Diagnosis:
Flowers ebracteate; calyx with a rounded base (1-6 mm long), trichomes straight,
non-reflexed; corolla relatively small (limb 1-2 mm wide); nutlets lanceolate to
lance-ovate (1.5-2 mm long), 4 per fruit, with distinctly angled margins.

Krynitzkia - Rough Nutlet Group (See Figure 28, 29E-H, 30 for nutlet images)
[Note: an asterisk indicates that taxon appears in the Krynitzkia Smooth Nutlet key or
in more than 1 couplet.]

1. One or more nutlets of fruit with margin sharp-angled and flattened as a narrow rim or a
narrow to broad wing

2. Pedicels 1-4 mm long in fruit; plant annual or perennial

3. Nutlets homomorphic, dorsally tuberculate-papillate, margin with narrow, flattened rim

or wing; plant usu. annual, with major central axis; inflorescence in fruit dense
29. C. holoptera

3’ Nutlets heteromorphic, 1 longer and dorsally papillate to papillate-tuberculate, 2-3
shorter and dorsally papillate-tuberculate, margins with very narrow, flattened rim; plant
perennial (but may flower first year), without major, central axis; inflorescence in fruit

open 30. C. racemosa

T Pedicels 0-1 mm long in fruit; plant annual

4. Nutlet 1 (rarely 2) per fruit; fruit sepals ovate-elliptic, nutlet margin sharp-angled to

rimmed, dorsal tubercles minutely spinulose 31. C. utahensis

4’ Nutlets usu. 4 per fruit; fruit sepals linear-lanceolate, nutlet margin winged along entire
margin, dorsal tubercles, if present, not spinulose

5. At least some nutlets with prominent, strongly lobed wings C pterocarya

6. Nutlets heteromorphic, 3 nutlets winged, 1 not winged

7. Wing of winged nutlets very broad 32. C. p. var. pterocarya

T Wing of winged nutlets narrow, knifelike 33. C p. var. purpusii

6’ Nutlets homomorphic, all 4 nutlets with prominent wings

34. C. p. var. cycloptera

5’ Nutlets sharp-angled, rimmed, or narrowly winged (not strongly lobed)

8. Nutlets homomorphic

9. Corolla limb 4-9 mm wide; calyx in fruit 3-4 mm long; nutlets 2-2.5 mm

long, narrowly winged (wings sometimes toothed), dorsal surface coarsely
tuberculate; flowers ebracteate 35. C. oxygona

9’ Corolla limb 1-3 mm wide; calyx in fruit 4-6 mm long; nutlets 1.5-2 mm long,
narrowly winged (wings sharp, never toothed), dorsal surface finely tuberculate;

flowers bracteate (bracts deciduous) 36. C. costata

8’ Nutlets heteromorphic by size, one distinctly larger than the other three

10. Nutlet margin rimmed and knife-like to narrowly winged; small nutlets ca. 1.3

mm long 37. C. inaequata

10’ Nutlet margin sharp-angled to slightly rimmed (usually rounded); small nutlets

ca. 1 mm long * 38. C. angustifolia

V All nutlets of fruit with margin rounded, ridged, or sharp-angled (acute to obtuse), but not
flattened as a rim or wing

11. Nutlets 2-4 per fruit, heteromorphic by size, one nutlet larger than the other(s)

12. Nutlets 4 per fruit, similar in sculpturing

13. Plants erect; nutlets lance-ovate, small nutlets ca. 1 mm long, finely papillate,
ventral groove narrow * 38. C. angustifolia

30

Crossosoma 35(1), Spring-Summer 2009

13’ Plants sprawling; nutlets narrowly lanceolate, small nutlets ca. 2 mm long,

tuberculate-papillate, ventral groove wide 39. C. dumetorum

12’ Nutlets 2-4 per fruit, different in sculpturing, the larger nutlet smooth to mostly
smooth, the smaller nutlet(s) papillate to tuberculate

14. Nutlets 4 per fr, < 1 mm long, triangular-ovate; flower bracts absent; calyx in fruit

sub-globose with hooked trichomes, sepals 1-1.5 mm long 40. C. micromeres

14’ Nutlets 2 per fruit, 1 .5-2 mm long, oblong-lanceolate; flower bracts present, calyx in

fruit oblong with straight trichomes, sepals 2-3 mm long * 21. C. maritima

11’ Nutlets 1-4 per fruit, homomorphic in size and sculpturing

15. Nutlets lanceolate (length: width ratio >3:1)

16. Nutlet 1 (-2) per fruit

17. Mature fruit strongly recurved from stem; nutlet incurved 41. C. recurvata

17’ Mature fruit not recurved from stem; nutlet(s) not incurved

18. Corolla limb 1-1.5 mm wide, deserts 42. C. decipiens

18’ Corolla limb 3-5 mm wide, cismontane 43. C. corollata

16’ Nutlets usu. 4 per fruit

19. Calyx 6-10 mm long; nutlets coarsely tuberculate C. nevadensis

20. Plants 20-60 cm tall; secondary branches sprawling; sepals strongly recurved

at tips; corolla limb 1-2 mm wide 44. C. nevadensis var. nevadensis

20’ Plants <20 cm tall; secondary branches ascending; sepals erect to slightly

recurved at tips; corolla limb 2-4 (5) mm wide 45. C. nevadensis var. rigida

19’ Calyx 4-6 mm long; nutlets finely tuberculate-spinulose 46. C. scoparia

15’ Nutlets lance-ovate, ovate, or triangular-ovate (length:width ratio < 3: 1)

21. Nutlet dorsal surface with at least a faint median ridge C. muricata

22. Corolla limb 1-2.5 mm wide; nutlets brownish

23. Stem bearing, from top to below middle, many short, flower-bearing
branchlets from a larger, primary axis, forming an elongate, leafy, thyrse

inflorescence 47. C. muricata var. jonesii

23’ Stem loosely and sparsely branched, without a larger primary axis

48. C. muricata var. denticulata

22’ Corolla limb 3-8 mm wide, nutlets grayish 49. C. muricata var. muricata

21’ Nutlet dorsal surface lacking median ridge

24. Corolla limb 3-9 mm wide

25. Sepals 2-3 mm in flower, 3 .5-5 mm in fruit

50. C. intermedia var. intermedia

25’ Sepals 4—6 mm in flower, 5-10 mm in fruit

51. C. barbigera var. fergusoniae

24’ Corolla limb 1-2 mm wide

26. Nutlets 1—1.5 (1.8) mm long; flowers bracteate; southern Channel Islands
52. C. traskiae

26’ Nutlets 1.5-3 mm long; flowers ebracteate; mainland

27. Nutlet dorsal face flattened to low-convex, densely papillate, sparsely

tuberculate 53. C. simulans

27’ Nutlet dorsal face convex, densely tuberculate, papillate or not

28. Nutlets ca. 3 mm long, turbercle tips translucent 54. C. clokeyi

28’ Nutlets 1 .5-2 mm long, turbercle tips not translucent

29. Calyx in fruit 5-6 mm long; nutlets gen 4, ovate, sculpturing finely
tuberculate and papillate; corolla limb 1-1.5 mm wide

55. C. echinella

29’ Calyx in fruit 5-10 mm long; nutlets 1-4, lanceolate to ovate,
sculpturing coarsely tuberculate to muricate, not papillate;
corolla limb 1-2.5 mm wide 56. C. barbigera var. barbigera

Crossosoma 35(1), Spring-Summer 2009

31

29. Cryptantha holoptera (A. Gray) J.F. Macbr. (Figure 13A-C, p.32)

Common name: Winged cryptantha. Distribution in Southern California:

eastern Mohave Desert, Sonoran Desert; Imperial, Inyo, Los Angeles, Riverside,
San Bernardino, and San Diego counties. Conservation Status: CNPS List 4.3.
Diagnosis: Annual duration (sometimes perennial); stems with a major axis
(inflorescence “dense”); pedicels long (1-4 mm); nutlets homomorphic, tuberculate-
papillate, narrowly winged (4 per fruit).

30. Cryptantha racemosa (S. Watson) Greene (Figure 13D-G, p.32)

Common name: Bushy cryptantha. Distribution in Southern California:

Southwestern California region (northern San Bernardino mountains, eastern
Peninsular Ranges), Desert province, northern White and Inyo mountains; Imperial,
Inyo, Los Angeles, Mono, Riverside, San Bernardino, San Diego counties.
Conservation Status: none. Diagnosis: Perennial duration; stems without a single,
major axis (inflorescence “open”); pedicels long (1-4 mm); nutlets heteromorphic,
papillate to tuberculate-papillate, narrowly winged (4 per fruit).

31. Cryptantha utahensis (A. Gray) Greene (Figure 14A-D, p.33)

Common name: Utah/scented cryptantha. Distribution in Southern California:

Desert province, White and Inyo mountains; Inyo, Kem, Los Angeles, Riverside,
San Bernardino, San Diego counties. Conservation Status: none. Diagnosis:
Pedicels short (<1 mm); nutlet 1 per fruit, with distinctive tuberculate-spinulose
processes and a sharp-angled to rimmed margin.

32. Cryptantha pterocarya (Torrey) Greene var. pterocarya

(Figure 14E-G, 15A-B, p.33,34)
Common name: Fringe nut cryptantha. Distribution in Southern California:

Southwestern California region (eastern Peninsular Ranges), Desert province,
White and Inyo mountains, often found in rocky uplands; Imperial, Inyo, Kern, Los
Angeles, Mono, Riverside, San Bernardino, San Diego counties. Conservation
Status: none. Diagnosis: Pedicels short (<1 mm); nutlets 4 per fruit, at least some
of which have distinctive, strongly lobed wings. This variety is distinguished from
the other two by the heteromorphic nutlets, three of which have broad (>0.5 mm),
strongly lobed wings and one of which is unwinged.

33. Cryptantha pterocarya (Torrey) Greene var. purpusii Jeps. (Figure 15D-E, p.34)
Common name: Argus Mountains cryptantha. Distribution in Southern
California: Southwestern California region (Peninsular Ranges, Transverse
Ranges), Desert province, White and Inyo mountains; Imperial, Inyo, Kern, Los
Angeles, Riverside, San Bernardino, San Diego counties. Conservation Status:
none. Diagnosis: Pedicels short (<1 mm); nutlets 4 per fruit, at least some of which
have distinctive, strongly lobed wings. This variety is distinguished from the other
two in having heteromorphic nutlets, three of which have narrow (<0.5 mm), lobed
wings and one of which is unwinged. [C. pterocarya (Torrey) Greene in part , sensu
W. A. Kelley and Wilken 1993]

32

Crossosoma 35(1), Spring-Summer 2009

pedicels
> 1 m nil

[(SjJffiloptew ]

pedicels
> 1 mm

C. racemosa

@$wcemosa

\r L aeem'osa\

C. holoptera

Figure 13. A-C: Cryptantha holoptera. A: Fruit, showing relatively long pedicel. B: Nutlet, dorsal
(left) and ventral (right) views. Note ovate- triangular shape, narrow winge and scattered tubercles.
C: Whole plants, showing single, major axis. D-G: C. racemosa. D: Whole plants, showing dense
banching with no major primary axis. E: Fruit, showing long pedicel. F: Inflorescence. G: Nutlets
from a single fruit, dorsal view. Note size heteromorphism.

holoptera

Crossosoma 35(1), Spring-Summer 2009

33

[G\utahe'misi

utanerms

G. utahensh

nw Rtemcarva

nutlets 1(2)
per fruit

C. utahensis

Cmb pterocarya

mewmma G fe- pterocarya

Figure 14. A-D: Cryptantha utahensis. A: Mature, elongate inflorescence. B: Inflorescence unit.
Note narrowly elliptic sepals. C: Fruit, showing single nutlet. D: Nutlet, dorsal, ventral, lateral,
and cross-sectional views (left to right). Note sharp-angled ridge and spinulose sculpturing.
E-G: C. pterocarya var. pterocarya. E: Inflorescence, showing swollen, ovoid calyx and small
corollas. F: Mature fruit, with four nutlets. G: Close-up of stem (below and right) and leaf (upper
left).

34

Crossosoma 35(1), Spring-Summer 2009

nutlets heteromorphic

C. pterocarya var. pterocarya

nutlets heteromorphic

Figure 15. A-B: Cryptantha pterocarya var. pterocarya. A: Heteromorphic nutlets of same fruit.
Left: one of three winged nutlets (dorsal and ventral views). Right: Single unwinged nutlet, dorsal
and ventral views. B: Nutlets of a common fruit, heteromorphic: three winged, one unwinged.
C: C. pterocarya var. cycloptera. Nutlets homomorphic, all winged. D-E: C. pterocarya var.
purpusii. D: Nutlet, dorsal view. Note narrow, lobed wing. E: Nutlets of a common fruit,
heteromorphic: three winged, one unwinged.

Crossosoma 35(1), Spring-Summer 2009

35

34. Cryptcintha pterocarya (Torrey) Greene var. cycloptera (Greene) J.F. Macbr.

(Figure 15C, p.34)

Common name: Tucson cryptantha. Distribution in Southern California:

Southwestern California region (eastern Peninsular Ranges), Desert province,
White and Inyo mountains, lower elevations of the desert; Imperial, Inyo, Riverside,
San Bernardino, San Diego counties. Conservation Status: none. Diagnosis:
Pedicels short (<1 mm); nutlets 4 per fruit, at least some of which have distinctive,
strongly lobed wings. This variety is distinguished from the other two in having
homomorphic nutlets, all of which have broad (>0.5 mm), strongly lobed wings.
[C. pterocarya (Torrey) Greene in part , sensu W.A. Kelley and Wilken 1993]

35. Cryptantha oxygona (A. Gray) Greene (Figure 16A-E, p.36)

Common name: Sharp nut cryptantha. Distribution in Southern California:

Southwestern California region (Transverse and northern Peninsular Ranges),
western Mohave Desert, Tehachapi mountains. White and Inyo mountains; Inyo,
Kern, Los Angeles, Mono, Riverside, San Bernardino, Santa Barbara, and Ventura
counties. Conservation Status: none. Diagnosis: Pedicels short (<1 mm); corolla
large (4-9 mm wide); nutlets 4 per fruit (2-2.5 mm long), homomorphic, and
narrowly winged (wings sometimes toothed but not strongly lobed).

36. Cryptantha costata Brandegee (Figure 16F-K, p.36)

Common name: Ribbed/ashen cryptantha. Distribution in Southern California:

Desert province. White and Inyo mountains, in sand dunes; Imperial, Inyo,
Riverside, San Bernardino, San Diego counties. Conservation Status: CNPS List
4.3. Diagnosis: Pedicels short (<1 mm); corolla small (1-3 mm wide); nutlets 4 per
fruit (1.5-2 mm long), homomorphic, narrowly winged (wings sharp, never toothed
or lobed).

37. Cryptantha inaequata I.M. Johnst. (Figure 17A-F, p.37)

Common name: Panamint cryptantha. Distribution in Southern California:

Mohave Desert, especially eastern; Imperial, Inyo, Riverside, San Bernardino
counties. Conservation Status: none. Diagnosis: Pedicels short (<1 mm); nutlets
4 per fruit, tuberculate-papillate, rimmed to narrowly winged (wings sharp, never
toothed or lobed), heteromorphic by size, 1 slightly larger than the other 3.

38. Cryptantha angustifolia (Torrey) Greene (Figure 17G-K, p.37)

Common name: Narrow-leaved cryptantha. Distribution in Southern California:
Desert province; Imperial, Inyo, Riverside, San Bernardino, San Diego counties.
Conservation Status: none. Diagnosis: Pedicels short (<1 mm); nutlets 4 per fruit,
tuberculate-papillate, margin usually rounded (rarely rimmed to narrowly winged,
wings sharp, never toothed or lobed), usually heteromorphic by size, 1 slightly
larger than the other 3 (rarely homomorphic). In addition to variation in nutlet
morphology in this species, two forms have been noted that vary in corolla limb
width and trichome density (personal communication).

36

Crossosoma 35(1), Spring-Summer 2009

Figure 16. A-E: Cryptantha oxygon a. A: Whole plant from specimen. B: Flower, showing
relatively wide corolla. C: Fruit. D: Fruit, with 4, homomorphic nutlets removed. E: Nutlet,
dorsal, ventral, and side views (left to right). Note narrow, toothed wing. F-K: Cryptantha costata.
F: Whole plant from specimen. G: Plant in field. Note large, lanceolate inflorescence bracts.
H: Close-up of leaf (left) and stem (right). I: Fruit. J: Fruit with sepals removed, showing four
nutlets. K: Nutlet, dorsal (left) and ventral (right) views.

Crossosoma 35(1), Spring-Summer 2009

37

C. inaequata

[(f^in^quataj

[inaequataj

kF. angustifolia

l^^n'gustimlia )

G^mgustifdl la

small nutlet

margin narrowly
winged

C. inaequata

C. inaequata

Figure 17. A-F: Cryptantha inaequata A: Whole plants from specimen. B: Close-up of
stem (left) and leaf (right). C: Flower, showing wide corolla limb. D: Heteromorphic nutlets,
one larger. E: Large nutlet, dorsal, ventral, and side views (left to right). F: Small nutlet,
dorsal, ventral, and side views (left to right). Note ovate-triangular shape and narrow wing.
G-K: C. angustifolia. G: Whole plant in field. H: Inflorescence close-up. I: Close-up of stem
(left) and leaf (right). J: Nutlets of one fruit, showing size heteromorphism. Note larger nutlet
adherent to gynobase. K: Small nutlet, dorsal, ventral, and side views (left to right).

38

Crossosoma 35(1), Spring-Summer 2009

39. Cryptantha dumetorum (A. Gray) Greene (Figure 18A-G, p.39)

Common name: Bush-loving cryptantha. Distribution in Southern California:

Desert province, Southwestern California region (northern Transverse Ranges);
Inyo, Kem, Los Angeles, Riverside, San Bernardino counties. Conservation
Status: none. Diagnosis: Stem habit sprawling, usually climbing in shrubs, foliage
very green; nutlets long (2-2.5 mm), lanceolate to lance-ovate, turberculate-
papillate, heteromorphic by size (1 larger, 3 smaller) with a wide ventral groove.

40. Cryptantha micromeres (A. Gray) Greene (Figure 18H-M, p.39)

Common name: Minute-flowered cryptantha. Distribution in Southern
California: Southwestern California region (South Coast, Channel Islands,
Peninsular Ranges, and Transverse Ranges); Los Angeles, Riverside, San
Bernardino, San Diego, Santa Barbara, Ventura counties. Conservation Status:
none. Diagnosis: Calyx sub- globose with hooked trichomes, 1-1.5 mm long;
nutlets small (<1 mm long), heteromorphic, 1 large and smooth to mostly smooth,
3 smaller and tuberculate-papillate.

41. Cryptantha recurvata Coville (Figure 19D-E, p.40)

Common name: Bent-nut cryptantha. Distribution in Southern California:

Desert province (Mohave, rare in Sonoran), White and Inyo mountains; Inyo, Mono,
San Bernardino, San Diego counties. Conservation Status: none. Diagnosis:
Nutlet 1 (-2) per fruit, strongly recurved from stem, recurved in fruit at maturity,
round-margined, lanceolate, and tuberculate-papillate.

42. Cryptantha decipiens (M.E. Jones) A. A. Heller (Figure 19F-H, p.40)

Common name: Gravel cryptantha. Distribution in Southern California: Desert
province, White and Inyo mountains; Imperial, Inyo, Kem, Los Angeles, Riverside,
San Bernardino, San Diego, Santa Barbara, Ventura counties. Conservation Status:
none. Diagnosis: Corolla small (1-1.5 mm); nutlet 1 (-2) per fruit, notrecurved from
stem at maturity, straight, round-margined, lanceolate, and tuberculate-papillate.

43. Cryptantha corollata (I.M. Johnst.) I.M. Johnst. (Figure 20A-B, p.41)

Common name: Coast range cryptantha. Distribution in Southern California:

Southwestern California region (Transverse Ranges, northern Peninsular Ranges),
Tehachapi mountains; Los Angeles, Riverside, San Bernardino, Santa Barbara,
Ventura counties. Conservation Status: none. Diagnosis: Corolla relatively large
(3-5 mm); nutlet 1 (-2) per fruit, not recurved from stem at maturity, straight, round-
margined, lanceolate, and tuberculate-papillate. [C. decipiens (M.E. Jones) A.A.
Heller in part , sensu W.A. Kelley and Wilken 1993]

44. Cryptantha nevadensis A. Nels. & PB. Kennedy var. nevadensis

(Figure 20C-H, p.41)

Common name: Nevada cryptantha. Distribution in Southern California: Desert
province, Tehachapi mountains, Southwestern California region (Transverse and
Peninsular Ranges), White and Inyo mountains; Imperial, Inyo, Kem, Los Angeles,
Mono, Riverside, San Bernardino, San Diego, Santa Barbara, Ventura counties.
Conservation Status: none. Diagnosis: Secondary branches sprawling, tending

Crossosoma 35(1), Spring-Summer 2009

39

N C. dumetorum

dumetorum

C.imicmmeres

micromeres}

microniefesl

micromeres}

large

nutlet

Wumetowm

plant lax, (sprawling

i

large, smooth nutlet

M

C. micromeres

Figure 18. A-G: Cryptantha dumetorum. A: Apical shoot of plant, from field. Stem is sprawling.
B: Stem and leaf close-up. C-D: Inflorescences. Note appressed flowers and fruits. E: Large
nutlet, dorsal (left) and ventral (right) views. Note large, open ventral groove. F: Small nutlet,
dorsal (left) and ventral (right) views. G: Two of four nutlets (small at left, large at right) still
attached to gynobase. H-M: C. micromeres. H: Whole plant, from specimen. I: Inflorescence.
J: Fruit, showing ovoid calyx and hooked trichomes. K: Fruit with four nutlets removed. Note
heteromorphism. L: Large, mostly smooth nutlet, dorsal (left) and ventral (right) views. M: Small,
turberculate nutlet, dorsal, ventral, and side views.

40

Crossosoma 35(1), Spring-Summer 2009

[C. mahtima

manitima ;

recurvata

WW^ecTpiemj

lecipiens

maturelfrum

C. recurvata

nutlets 2, heteromorphic: 1 smooth
1 rough (or nutlet l^mooth)

Figure 19. A-C: Cryptantha maritima. A: Inflorescence, showing bracteate flowers. B: Close-
up of flowers and fruits. Note dense, straight trichomes. C: Heteromorphic nutlets, dorsal view,
smooth (left) and apically rough (right). D-E: C. recurvata. D: Fruit, showing recurved calyx.
E: Nutlet close-up, dorsal (left), ventral (middle), and lateral (right) views. Note recurved apex.
F-H: C. decipiens. F: Fruit, asceding to inclined (not recurved). G: Flower, showing small
corolla. H: Nutlet, dorsal (left) and ventral (right) views.

Crossosoma 35(1), Spring-Summer 2009

41

Figure 20. A-B: Cryptaniha corollata. A: Flower, showing large corolla. B: Nutlet close-
up, dorsal, ventral, and lateral views. Note relatively straight apex. C-H: C. nevadensis var.
nevadensis. C: Herbarium specimen, showing sprawling stem habit. D: Inflorescence, showing
strongly recurved sepal tips. E: Close-up of stem (left) and leaf (lower right). F: Calyx of fruit.
Note long length. G: Flower, showing small (1-2 mm wide) corolla limb. H: Nutlet close-up, dorsal
(left) and ventral (right) views. Note coarse tuberculation.

42

Crossosoma 35(1), Spring-Summer 2009

to crawl through shrubs; calyx relatively long (6-10 mm) with strongly recurved
tips; corolla small (1-2 mm wide); nutlets round-margined, lanceolate, 4 per fruit,
straight, coarsely tuberculate.

45. Cryptantha nevadensisA. Nels. & P.B. Kennedy var. rigida I.M. Johnst.

(Figure 2 1A-E, p.43)

Common name: Rigid cryptantha. Distribution in Southern California:

Tehachapi mountains, Mohave Desert, Southwestern California region (Transverse
and Peninsular Ranges), brushy slopes along the edge of the desert; Inyo, Kern, Los
Angeles, San Bernardino, Santa Barbara, Ventura counties. Conservation Status:
none. Diagnosis: Plant +/- erect and self supporting, with ascending secondary
branches; calyx with erect to slightly recurved tips; corolla limb larger (2-4 mm
wide). [C. nevadensis A. Nels. & P.B. Kennedy in part , sensu W.A. Kelley and
Wilken 1993]

46. Cryptantha scoparia Nelson (Figure 21 F-H, p.43)

Common name: Great Basin cryptantha. Distribution in Southern California:

White and Inyo mountains; Inyo, Mono counties. Conservation Status: CNPS List
4.3. Diagnosis: Calyx smaller (4-6 mm) and round-margined; nutlets lanceolate,
4 per fruit, straight, finely tuberculate-spinulose. [C. nevadensis A. Nels. & P.B.
Kennedy in part , sensu W.A. Kelley and Wilken 1993]

47. Cryptantha muricata (Hook. & Arn.) A. Nels. & J.F. Macbr. var jonesii (A. Gray)

I.M. Johnst. (Figure 22A, 22D-F, p.44)

Common name: Jones’s cryptantha. Distribution in Southern California:
Southwestern California region (South Coast, Transverse and Peninsular Ranges);
Los Angeles, Orange, Riverside, San Bernardino, San Diego, Santa Barbara,
Ventura counties. Conservation Status: none. Diagnosis: Corolla smaller (1-2.5
mm); nutlets brownish; stems with a relatively large primary axis bearing many
short, flower-bearing branchlets, forming an elongate, leafy, thyrse inflorescence.
[C. muricata (Hook. & Am.) A. Nels. & J.F. Macbr. in part , sensu W.A. Kelley and
Wilken 1993]

48. Cryptantha muricata (Hook. & Am.) A. Nels. & J.F. Macbr. var. denticulata

(Greene) I.M. Johnst. (Figure 22B, p.44)

Common name: Prickly-nut cryptantha. Distribution in Southern California:
Southwestern California region (Transverse and Peninsular Ranges); Los Angeles,
Riverside, San Bernardino, San Diego counties. Conservation Status: none.
Diagnosis: Nutlets ovate to triangular-ovate, muricate, with at least a faint median
ridge on the dorsal surface. This variety is distinguished from the others by the
smaller (1-2.5 mm) corolla, brownish nutlets, and stems that are loosely and
sparsely branched, without a large, primary axis. [C. muricata (Hook. & Arn.) A.
Nels. & J.F. Macbr. in part , sensu W.A. Kelley and Wilken 1993]

Crossosoma 35(1), Spring-Summer 2009

43

^^neyad emis
m wvMgida

A C. nevadensis

C^neyddensis var. rigida'

corolla>2?4Imm

SlriTTiTl

C. nevadensis var. rigida

w |f w

gssanSBgg 2.

Figure 21. A-E: Cryptantha nevadensis var. rigida. A: Herbarium specimen. B: Flower close-
up, showing corolla (2-4 mm wide). C: Inflorescence with fruits. Note long calyx with slightly
recurved sepal tips. D: Fruit, with 4 nutlets removed. E: Nutlet close-up, dorsal (left), ventral
(middle), and lateral (right) views. F-H: Cryptantha scoparia. F: Herbarium specimen. G: Fruit,
showing smaller calyx. H: Nutlet close-up, dorsal (left), ventral (middle), and lateral (right) views.

44

Crossosoma 35(1), Spring-Summer 2009

C. muricata I
A var jonesii I

aC. muricata
var. denticulata

C. muricata vaM]onesii>

muricata*
E^ani 'jonesni

SQQSflfe

ft fiVrrrm

C. muricata var. murjcata

C. muricata var. jonesu

G jimermec^

C. intermedia

Figure 22. A-F: Cryptantha muricata. A: C. muricata var. jonesii. Herbarium specimen,
showing erect, primary axis bearing numerous lateral inflorescences. B: C. muricata var.
denticulata. Herbarium specimen, showing numerous ascending secondary branches and absence
of strong primary axis. C: C. muricata var. muricata. Flower close-up, showing large corolla.
D-F: C. muricata var. jonesii D: Corolla, small. E: Fruit, with 4 attached nutlets. F: Nutlet close-
up, dorsal, ventral, and lateral views. Note ovate shape and dorsal ridge. G-l : Cryptantha intermedia
var. intermedia. G: Flower with large corolla. H: Fruit, sepals removed, showing four nutlets.
I: Nutlet close-up, dorsal, ventral, and lateral views. Note absence of dorsal ridge.

Crossosoma 35(1), Spring-Summer 2009

45

49. Cryptantha muricata (Hook. & Arn.) A. Nels. & J.F. Macbr. var. muricata

(Figure 22C, p.44)

Common name: Showy prickly cryptantha. Distribution in Southern

California: Southwestern California region (Transverse and Peninsular Ranges);
Imperial, Kem, Los Angeles, Orange, Riverside, San Bernardino, San Diego, Santa
Barbara, Ventura counties. Conservation Status: none. Diagnosis: This variety is
distinguished from the others by the wider (3-8 mm) corolla and grayish nutlets.

50. Cryptantha intermedia (A. Gray) Greene var. intermedia (Figure 22G-I, p.44)
Common name: Common/Nievitas cryptantha. Distribution in Southern
California: Southwestern California region (South Coast, Channel Islands - San
Clemente and Santa Catalina Islands, Transverse and Peninsular Ranges); Imperial,
Inyo, Kern, Los Angeles, Orange, Riverside, San Bernardino, San Diego, Santa
Barbara, Ventura counties. Conservation Status: none. Diagnosis: Corolla
relatively wide (3-6 mm), showy; nutlets 4 per fruit, homomorphic, round-margined,
lance-ovate to ovate, tuberculate, lacking a dorsal ridge. Note: C. intermedia var.
grandiflora [=C. hendersonii (Nelson) Piper], not native to Southern California,
has much larger corollas.

51. Cryptantha barbigera (A. Gray) Greene var. fergusoniae J.F. Macbr.

(Figure 23A-E, p.46)

Common name: Palm Springs cryptantha. Distribution in Southern California:

western Colorado Desert, especially Coachella Valley; Imperial, Riverside, San
Bernardino San Diego counties. Conservation Status: none. Diagnosis: This
variety is largely limited in distribution to the Coachella Valley and periphery. It
is distinguished from C. intermedia by the significantly larger (4-6 mm in flower,
5-10 mm in fruit) calyx and from C. barbigera var. barbigera by the larger (4-9
mm wide) corolla. [C. barbigera (A. Gray) Greene in part , sensu W.A. Kelley and
Wilken 1993]

52. Cryptantha traskiae I.M. Johnst. (Figure 23F-H, p.46)

Common name: Trask’s cryptantha. Distribution in Southern California:

Southwestern California region (Channel Islands - San Clemente, San Nicolas
Islands); Los Angeles, Ventura counties. Conservation Status: CNPS List IB. 2.
Diagnosis: Corolla small (1-2 mm wide), nutlets round-margined, lance-ovate to
ovate, tuberculate, 1-1. 5(1. 8) mm long, lacking a dorsal ridge.

53. Cryptantha simulans Greene (Figure 24A-C, p.47)

Common name: Pine cryptantha. Distribution in Southern California:

Southwestern California region (Transverse and Peninsular Ranges); Kem, Los
Angeles, Riverside, San Bernardino, San Diego, Santa Barbara, Ventura counties.
Conservation Status: none. Diagnosis: Corolla small (1-2 mm wide); nutlets 4
per fruit, round-margined, ovate with dorsal face flattened to low-convex, densely
papillate - sparsely tuberculate, ca. 2 mm long, lacking a dorsal ridge.

46

Crossosoma 35 ( 1 ), Spring-Summer 2009

(fjmarbigera

vmifergusoniae

Mjambigma H
v arf t^gusoma^Sj

WSMbmb isera

^wTfergusahiae

[Gfbmmgeral

g ^ar ^Jergusaniae<

i F C. traskiae

Kf traskiae

C. traskiae

[corollal4r9^mm\wide]

C. barbigera var. fergusoniae

Figure 23. A-E: Cryptantha barbigera var. fergusoniae. A: Whole plant from herbarium sheet.
B: Fruit. C: Close-up of stem (center) and leaf (upper right). D: Flower, showing large corolla.
E: Nutlet close-up, dorsal (left), ventral (middle), and lateral (right) views. F-H: C. traskiae.
F: Whole plant from herbarium specimen, showing short plant stature. G: Inflorescence.
H: Nutlet close-up, dorsal (left), ventral (middle), and lateral (right) views.

Crossosoma 35(1), Spring-Summer 2009

47

Ci simulans \

C. sirmUanSj

GWmom*

[(f&Iokeyil

lokeyil

fewlokey.il

C. simulans

nutlet tubercles
translucent

C. clokeyi

Figure 24. A-C: Cryptantha simulans. A: Whole plants from herbarium specimens.
B: Fruit close-up. C: Nutlet close-up, dorsal (left), ventral (middle), and lateral (right) views.
D-l: Cryptantha clokeyi. D: Whole plant from herbarium sheet. E: Close-up of stem (right)
and leaf (far left). F: Flower close-up. Note small corolla. G: Fruit. H: Nutlet close-up, dorsal
(left), ventral (middle), and lateral (right) views. I: Nutlet close-up, showing rounded, translucent
tubercles.

54. Cryptantha clokeyi I.M. Johnst. (Figure 24D-I, p.47)

Common name: Clokey’s cryptantha. Distribution in Southern California:

Mohave Desert; Inyo, Los Angeles, San Bernardino counties. Conservation
Status: CNPS List 1B.1. Diagnosis: Corolla small (1-2 mm wide); nutlets 4 per
fruit, ca. 3 mm long, round-margined, ovate, densely tuberculate with tubercle tips
translucent, lacking a dorsal ridge.

48

Crossosoma 35(1), Spring-Summer 2009

Figure 25. A-C: Cryptantha echinella. A: Whole plant from herbarium specimen.
B: Inflorescence. C: Nutlet close-up, dorsal (left), ventral (middle), and lateral (right) views.
D-H: C. barbigera var. barbigera. D: Whole plant in field. E: Inflorescence close-up. Note small
corollas. F: Fruit, showing long calyx. G: Nutlet close-up, dorsal (left) and ventral (right) views.
H: Flower, showing small corolla.

Crossosoma 35(1), Spring-Summer 2009

49

55. Cryptantha echinella Greene (Figure 25A-C, p.48)

Common name: Spiny cryptantha. Distribution in Southern California:

Tehachapi mountains, Southwestern California region (Transverse Ranges),
Mohave Desert; Inyo, Kern, Los Angeles, Mono, San Bernardino, Santa Barbara,
Ventura counties. Conservation Status: none. Diagnosis: Calyx 5-6 mm long in
fruit; corolla small (1-1.5 mm wide); nutlets 4 per fruit, ca. 1.5-2 mm long, round-
margined, ovate, finely tuberculate and papillate, lacking a dorsal ridge.

56. Cryptantha barbigera (A. Gray) Greene var. barbigera (Figure 25D-H, p.48)
Common name: Bearded cryptantha. Distribution in Southern California: Desert
province, Tehachapi mountains; Imperial, Inyo, Kern, Los Angeles, Riverside, San
Bernardino, San Diego, Santa Barbara, Ventura counties. Conservation Status:
none. Diagnosis: Calyx in fruit long (5-10 mm); nutlets 1-4 per fruit, 1.5-2 mm
long, round-margined, ovate, densely tuberculate-muricate, lacking a dorsal ridge.
This variety is distinguished from C. b. var. fergusoniae by the relatively small
corolla (1-2.5 mm wide). [C. barbigera, in part, sensu W.A. Kelley and Wilken
1993]

NUTLET COMPARISON WITHIN MAJOR GROUPS

For comparison of nutlets within major groups, see Figures 26-30 (at end of article)
for images of all Southern California Cryptantha taxa, arranged as follows: Oreocarya
Group (Figure 26), Krynitzkia Group, smooth nutlets (Figure 27), Krynitzkia Group,
nutlets with ridged or winged margin (Figure 28), Greeneocharis Group (Figure
29A,B), Eremocarya Group (Figure 29C,D), Krynitzkia Group, heteromorphic nutlets
with rounded margin (Figure 29 E-H), Krynitzkia Group, rough nutlets with rounded or
sharp-angled margin (Figure 30).

CONSERVATION STATUS AND ECOLOGY OF SOUTHERN CALIFORNIA
CRYPTANTHA

Nine species of Southern California Cryptantha are included in the 2009 California
Native Plant Society (CNPS) Inventory as sensitive or rare taxa. Of these, two species,
Cryptantha clokeyi and C. ganderi, are on CNPS List IB. 1, which signifies that they
are “rare, threatened, or endangered in California and elsewhere” and “seriously
endangered in California (over 80% of occurrences threatened/high degree and
immediacy of threat).” Cryptantha clokeyi is found in “sandy or gravelly soil” (R.B.
Kelley in prep) in the Mohave Desert region, in three counties (Inyo, Los Angeles, and
San Bernardino). Only 16 original collections are cited in the Consortium of California
Herbaria at the time of this writing. Cryptantha ganderi is found in “stabilized, +- silty,
fine sand deposits, creosote scrub association” (R.B. Kelley, in prep.), solely in parts of
the Borrego Valley of San Diego County. Only 9 original collections are currently cited
in the Consortium of California Herbaria.

50

Crossosoma 35(1), Spring-Summer 2009

Two Southern California Cryptantha species are on CNPS List 1B.2, meaning “rare,
threatened, or endangered in California and elsewhere” and “fairly endangered
in California (20-80% occurrences threatened).” One of these, C. roosiorum, is a
distinctive member of Oreocarya. It is known from “rocky, silty soils, dry meadows
in open bristlecone pine-limber pine forest, limestone endemic” (R.B. Kelley, in prep.)
of only three nearby localities in the Inyo Mountains (Inyo County). Only 5 original
collections are cited in the Consortium of California Herbaria. The second CNPS List
1B.2 species is C. traskiae, which is found only on two of the Channel Islands: San
Clemente Island (Los Angeles County) and San Nicolas Island (Ventura County), both
of the southern group. However, C. traskiae has been collected considerably more, as
60 original collections (almost two-thirds by S. A. Junak) are cited in the Consortium
of California Herbaria.

The five other Cryptantha species included in the CNPS Inventory - C. costata, C.
glomeriflora, C. holoptera, C. scoparia , and C. tumulosa - are on List 4.3, meaning they
are relatively rare, of “limited distribution (watch list)” and are “not very endangered in
California (<20% of occurrences threatened or no current threats known).”

Cryptantha taxa are found in a variety of substrate types and habitats. Most members
of the Eremocarya, Greeneocharis, and Krynitzkia groups grow in sandy or gravel
soil. Cryptantha fiavaculata, C. hoffmannii, C. humilis, C. roosiorum, C. tumulosa,
and C. virginensis of the Oreocarya group and C. inaequata of the Krynitzkia group
occur occasionally or solely in limestone-based substrates. Interestingly, members of
the perennial Oreocarya group tend to occur in high elevation, mountainous regions;
the taxa in Southern California are found at elevations ranging from 840-3600 meters
(2,700-11,800 feet), with a median elevation of 2,300 meters (7,700 feet). In addition,
two common Cryptantha species - C.flaccida and C. intermedia (no variety indicated)
- are cited as “weak” or “indifferent” indicators of ultramafic substrates (Safford et al.
2005).

TAXONOMIC PROBLEMS IN CRYPTANTHA

Several Cryptantha taxa of Southern California warrant further studies, both
quantitative statistical studies of morphology and genetic studies at the population
level. Comparative studies of biogeography, ecology, and reproductive biology of
the Greeneocharis species pair (C. circumscissa and C. simitis ) are needed to better
understand how they may be related to one another and how they diverged. Within the
Krynitzkia group, the varieties of C. clevelandii need to be properly documented with
respect to biogeographic distribution, habitat differences, and variation. Cryptantha
angustifolia is reported to have variants, including forms with homomorphic and
winged nutlets, in contrast to the typical heteromorphic, round-margined nutlets,
and forms differing in corolla width and vestiture. The complex of varieties in C.
muricata are at times difficult to identify and appear to intergrade, particularly in the
southern part of the range. Additional research is needed on this group (R. B. Kelley,
pers. comm. 2008). What is called C. intermedia var. intermedia shows considerable

Crossosoma 35(1), Spring-Summer 2009

51

variation in leaf morphology from coastal to montane habitats. Cryptantha corollata
shows considerable variation. Cryptantha maritima might be worthy of further study
with regard to variation in calyx trichome morphology, given that the synonomized
C. m. var. pilosa presumably differs in that regard. Both Cryptantha maritima and C.
circumscissa would be fruitful to study with respect to potential differences between
populations in North versus South America and the assessment of the direction
and timing of their dispersal events. In general, it is likely that many collections of
Cryptantha across California need careful annotation, using the taxonomy and keys
in the upcoming The Jepson Manual (R.A. Kelley, in prep). Just before this article
went to press, we received notification of a population of Cryptantha fendleri (not
previously known in California) collected this year in the Little Cowhorn Valley of
the Inyo Mountain range, thus technically part of the Southern California region.
Cryptantha fendleri, a member of the smooth nutlet Krynitzkia group, will be included
in the second edition of The Jepson Manual (R. A. Kelley, in prep) and is currently
being considered for CNPS List 2.3 [“rare, threatened, or endangered in California,
but more common elsewhere; not very endangered in California (<20% of occurrences
threatened or no current threats known)”]. Finally, Southern California Cryptantha
that are potentially new to science are currently being evaluated (Jim Andre, personal
communication; authors of this paper), which could add even more to the Southern
California region’s diversity of this fascinating group.

ACKNOWLEDGEMENTS

We are greatly indebted to the herbaria at COLO, RSA-POM, SD, SDSU, and UC-
JEPS for specimen loans, to all the herbaria that are members of the Consortium
of California Herbaria (CAS-DS, CDA, CHSC, DAV, HSC, IRVC, OBI, PGM,
RSA-POM, SBBG, SD, SDSU, SJSU, UC-JEPS, UCR, UCSB, and UCSC, as of
this writing) for access to their data, and to UC-JEPS in particular for creating and
maintaining the Consortium data interface. We thank Ron B. Kelley for an early version
of his descriptions and keys of Cryptantha, to be incorporated in the second edition of
The Jepson Manual. We thank Anna Schochenmaier for photographing several of the
high resolution images of nutlets. We thank Gary Monroe for allowing us to use field
images of C. cinerea var. abortiva, C. confertiflora, and C. flavoculata. We sincerely
thank Andy Sanders and Leroy Gross for reviewing this paper and giving useful
suggestions (but of course we take responsibility for any errors). Finally, we thank the
editors of Crossosoma for inviting us to publish this article, which is an offshoot of a
talk given at the 2008 Southern California Botanists meeting.

LITERATURE CITED

Abrams, L. 1951. Illustrated flora of the Pacific States. Vol. Ill Stanford University
Press, Stanford, California.

Angiosperm Phylogeny Group. 2003. An update of the Angiosperm Phylogeny Group
classification for the orders and families of flowering plants: APG II. Botanical
Journal of the Linnean Society 141:399-436.

52

Crossosoma 35(1), Spring-Summer 2009

Buys, M. H. and H. H. Hilger. 2003. Boraginaceae cymes are exclusively scorpioid and
not helicoid. Taxon 52:719-724.

California Native Plant Society (CNPS). 2009. Inventory of Rare and Endangered
Plants (online edition, v7-09b). California Native Plant Society. Sacramento,
California.

Consortium of California Herbaria. 2009. Data provided by the participants of the
Consortium of California Herbaria, http://ucjeps.berkeley.edu/consortium.

Don, G. 1836. A General System of Gardening and Botany: Founded upon Miller’s
Gardener ’s Dictionary, and Arranged according to the Natural System, volume
4. London, UK.

Engler, A., and K. Prantl. 1897. Die natiirlichen Pflanzenfamilien. Engelmann, Leipzig,
Germany.

Gottschling, M., H. H. Hilger, and N. Diane. 2001. Secondary structure of the
ITS 1 transcript and its application in a reconstruction of the phylogeny of the
Boraginales. Plant Biology 3:629-636.

Greene, E. L. 1887a. Some west American Asperifoliae I. Pittonia 1 : 8-23.

Greene, E. L. 1887b. Some west American Asperifoliae II. Pittonia 1: 55-60.

Greene, E. L. 1887c. Some west American Asperifoliae III. Pittonia 1: 77-120.

Hasenstab, K. 2009. Phylogenetic systematics of the genus Cryptantha (Boraginaceae).
Master of Science Thesis, San Diego State University, San Diego, California.

Heywood, V. H, R. K. Brummitt, A. Culham, and O. Seberg. 2007. Flowering Plant
Families of the World. Royal Botanic Gardens, Kew, Richmond, Surrey.

Hickman, J. C. (ed.). 1993. The Jepson Manual: Higher Plants of California. University
of California Press, Berkeley and Los Angeles.

Higgins, L. C. 1971. A revision of Cryptantha subgenus Oreocarya. Brigham Young
University Science Bulletin Biological Series 8: 1-62.

Johnston, I. M. 1925. The North American species of Cryptantha. Dissertation: The
Gray Herbarium of Harvard University. Cambridge, Massachusetts.

Johnston, I. M. 1927. A revision of the South American Boraginoideae. Contributions
of the Gray Herbarium Harvard University 78:2-1 18.

Kelley, R. B. Cryptantha. In The Jepson Manual: Higher Plants of California, second
edition, eds. Bruce G. Baldwin, Steve Boyd, D. J. Keil, R. W. Patterson, T. J.
Rosatti, and D. H. Wilken. University of California Press, Berkeley. In prep.
(Note: title and editors subject to change.)

Kelley, W. A., and D. Wilken. 1993. Cryptantha. In The Jepson Manual: Higher
Plants of California, ed. J. C. Hickman, 369-378. University of California Press,
Berkeley.

Mabberley, D. J. 2008. Mabberley’s Plant-Book: A Portable Dictionary of the Higher
Plants, Their Classification and Uses. 3rd edition. Cambridge University Press,
Cambridge.

Munz, P. A. and D. D. Keck. 1959. A California Flora. University of California Press,
Berkeley

Munz, P. A. and D. D. Keck. 1968. Supplement to A Flora of California. University of
California Press, Berkeley.

Crossosoma 35(1), Spring-Summer 2009

53

Munz, P. A. 1974. A Flora of Southern California. University of California Press,
Berkeley.

Payson, E. B. 1927. A monograph of the section Oreocarya of Cryptantha Annals of
the Missouri Botanical Garden 14:211-358.

Reiche, K. 1915. Estudios Criticos de la Flora de Chile. Anales de Universidad de
Chile. Santiago, Chile.

Rosatti, T. J. 2003. Jepson Online Interchange for California Floristics. Fremontia 31:
23-29.

Safford, H. D., J. H. Vires, and S. P. Harrison. 2005. Serpentine Endemism in the
California Flora: A Database of Serpentine Affinity. Madrono 52:222-257.

Schwarzer, C. 2007. Systematische Untersuchungen an den peruanischen Vertretem der
Gattungen Pectocarya D.C. ex Meisn., Amsinckia Lehm., Plagiobothrys Fisch.
& C.A. Mey. und Cryptantha Fehm. ex G. Don (Boraginaceae). Dissertation:
Biology Institut fur Biologie Systematische Botanik und Pflanzengeographie,
Freie Universitat Berlin, Germany.

Steven, P. 2001 onwards. Angiosperm Phylogeny Website. Version 9, June 2008
http://www. mobot. org/MOBOT/research/APweb

Weber, W. A. 1987. Colorado Flora: Western Slope. Colorado Associated University
Press, Boulder, Colorado.

Wiggins, I. F. 1980. Flora of Baja California. Stanford University Press, Stanford,
California.

Zuloaga, F., O. Morrone, and M. Belgrano (eds). Catalogo de las Plantas Vasculares
del Cono Sur (Checklist of the Southern Cone), volumes 1-3. Boraginaceae,
Cryptantha treatment. In prep.

54

Crossosoma 35(1), Spring-Summer 2009

Figure 26. Oreocarya group nutlets (dorsal view). A: Cryptantha cinerea var. abortiva.
B: C. confertiflora. C: C. flavoculata. D: C. hoffmannii. E: C. humilis. F: C. nubigena.
G: C. roosiorum. H: C. tumulosa. I: C. virginensis. All images to scale.

Crossosoma 35(1), Spring-Summer 2009

55

Figure 27. Krynitzkia group, smooth nutlets (dorsal view). A: Cryptantha affinis.
B: C. clevelandii var. clevelandii. C: C. clevelandii var. florosa. D: C. flaccida. E: C. ganderi.
F: C. glomeriflora. G: C. gracilis. H: C. leiocarpa. I: C. maritima. J: C. microstachys.
K: C. mohavensis. L: C. nemaclada. M: C. sparsiflora. N: C. torreyana. O: C. watsonii.
All images to scale.

56

Crossosoma 35(1), Spring-Summer 2009

Figure 28. Krynitzkia group, nutlets with a ridged or winged margin (dorsal view). A: Cryptantha
costata. B: C. holoptera. C: C. inaequata. D: C. oxygona. E: C. pterocarya xnr.pterocarya, winged
nutlet (left), unwinged (right). F: C. pterocarya var. purpusii, winged nutlet. G: C. racemosa.
H: C. utaliensis. All images to scale.

Crossosoma 35(1), Spring-Summer 2009

57

Greeneocharis Group Eremocarya Group

Krynitzkia Group: Heteromorphic Nutlets, Rounded Margin

Figure 29. A-B: Greeneocharis group nutlets (dorsal view). A: Cryptantha circumscissa.
B: C. similis. C-D: Eremocarya group nutlets (dorsal view). C: C. lepida. D: C. micrantha.
E-H: Krynitzkia group, heteromorphic nutlets with a rounded margin (dorsal view, except large
nutlet of C. angustifolia in lateral view). E: C. angustifolia. F: C. dumetorum. G: C. maritima.
H: C. micromeres. All images to scale.

58

Crossosoma 35(1), Spring-Summer 2009

Figure 30. Krynitzkia group, rough nutlets with a rounded or sharp-angled margin (dorsal view).
A: Cryptantha barbigera var. barbigera. B: C. barbigera var. fergusoniae. C: C. clokeyi.
D: C. corollata. E: C. decipiens. F: C. echinella. G: C. intermedia var. intermedia.
H: C. muricata var. denticulata. I: C. muricata var. jonesii. J: C. muricata var. muricata.
K: C. nevadensis var. nevadensis. L: C. nevadensis var. rigida. M: C. recurvata. N: C. scoparia.
O: C. simulans. P: C. traskiae. All images to scale.

Crossosoma 35(1), Spring-Summer 2009

59

Appendix 1. Voucher information for Cryptantha specimens examined and photographed in this
study.

Cryptantha affinis (A. Gray) Greene: Figure 8A, 27A(SDSU 5410); Cryptantha angustifolia (Torrey) Greene:
Figure 17G,H (SDSU 18676), Figure 171 (SDSU 13030), Figurel7J, 29E (SDSU 5376); Cryptantha barbigera
(A. Gray) Greene var. barbigera. Figure 24D,E (SDSU 18686), Figure 24F,H (SDSU 9986), Figure 24G, 30A
(SDSU 5378); Cryptantha barbigera (A. Gray) Greene var . fergusoneae J.F. Macbr.: Figure 25A-C,E, 30B
(RSA 712692), Figure 25D (RSA 713328); Cryptantha cinerea (Greene) Cronquist var. abortiva (Greene)
Cronquist: Figure 5F-H, 26A (RSA 46663); Cryptantha circumscissa (Hook. & Am.) I.M. Johns! var.
circumscissa Figure 3A,B (SDSU 18697), Figure 3C-E, 29A (SD 28881); Cryptantha clevelandii Greene var.
clevelandii. Figure 10F,G (SDSU 14050), Figure 10H, 27B (SDSU 14059); Cryptantha clevelandii Greene var.
ft orosa I.M. Johns!: Figure 11A (SDSU 18342), Figure 11B-D, 27C (SDSU 5417); Cryptantha clokeyi I.M.
Johns!: Figure 23G-L, 30C (RSA 665270); Cryptantha confertiflora (Greene) Payson: Figure 5B, 26B (RSA
196047), ; Cryptantha corollata (I.M. Johns!) I.M. Johns!: Figure 20A,B, 30D (POM 368220); Cryptantha
costata Brandegee: Figure 16F,H-J (SD 113106), Figure 16G(SDSU 17342), Figure 16K, 28A(SD 113307);
Cryptantha decipiens (M.E. Jones) A.A. Heller: Figure 19F,H, 30E (SDSU 18645), Figure 19G (SDSU 16365);
Cryptantha dumetorum (A. Gray) Greene: Figure 18A-D (SDSU 18694), Figure 18E-G, 29F (RSA 553891);
Cryptantha echinella Greene: Figure 24A-C, 30F (RSA 682890); Cryptantha flaccida (Lehmann) Greene:
Figure 8D(SD 120625), Figure 8E, 27D (SDSU 17311); Cryptantha flavoculata (A. Nels.) Payson: Figure 5D,
26C (SD 21927) ; Cryptantha ganderi I.M. Johns!: Figure 9C,D, 27E (SD 115050); Cryptantha glomeriflora
Greene: Figure 8B, 27F (SD 87724), Figure 8C (RSA 625075); Cryptantha gracilis Osterh.: Figure 10A(RSA
709650), Figure 10B, 27G (SD 4205); Cryptantha hoffmannii I.M. Johns!: Figure 7A,C (SD 81222), Figure 7B
(SDSU 5495), Figure 7D, 26D (RSA 79771 ); Cryptantha holoptera (A. Gray) J.F. Macbr.: Figure 13A,B, 28B
(SD 120959), Figure 1 3C (SDSU 13036); Cryptantha humilis (A. Gray) Payson: Figure 6E (SD 8761 8), Figure
6F, 26E (RSA 516990); Cryptantha inaequata I.M. Johns!: Figure 17A-F, 28C (RSA 173141); Cryptantha
intermedia (A. Gray) Greene var. intermedia. Figure 22G (SDSU 16074), Figure 22H,1, 30G (SDSU 14564);
Cryptantha leiocarpa (Fisher & C A. Meyer) Greene: Figure 9E,G, 27H (SDSU 19212); Cryptantha lepida
(A. Gray) I.M. Johns!: Figure 4A-C (SDSU 17572), Figure 4D-E, 29C (SDSU 17281); Cryptantha maritima
(Greene) Greene var. maritima. Figure 9F,H (SDSU; inset SD 113111), Figure 19A-C(left), 271, 29G (right) (SD
113111), Figure 19C(right), 29G (left) (SDSU 5460); Cryptantha micrantha (Torrey) I.M. Johns!: Figure 4F
(SDSU 5388), Figure 4G, 29D (SDSU 5421), Figure 4H*I (SDSU 5431), Figure 4J (SDSU 5419); Cryptantha
micromeres (A. Gray) Greene: Figure 18H,J (SDSU 16695), Figure 18K (SDSU 15465), Figure 18L-M, 29H
(SDSU 12173); Cryptantha microstachys (A. Gray) Greene: Figure 10C,D (SD 58176), Figure 10E, 27J (SD
44966); Cryptantha mohavensis (Greene) Greene: Figure 12A, 27K (SD 129726), Figure 12B (RSA 712528);
Cryptantha muricata (Hook. & Arn.) A. Nels. & J.F. Macbr. var. denticulata (Greene) I.M Johns!: Figure 22B,
30H (SD 161808), ; Cryptantha muricata (Hook. & Am.) A. Nels. & J.F. Macbr. var. jonesii (A. Gray) I.M.
Johns! . Figure 22A,E (SDSU 16714), Figure 22D (17512), Figure 22F, 301 (SDSU 5413); Cryptantha muricata
(Hook. & Am.) A. Nels. & J.F. Macbr. var. muricata. Figure 22C, 30J (SDSU 19186); Cryptantha nemaclada
Greene: Figure 9A,B, 27L (RSA 195150); Cryptantha nevadensis A. Nels. & P.B. Kennedy var. nevadensis.
Figure 20C-H, 30K (SDSU 13032); Cryptantha nevadensis A. Nels. & P.B. Kennedy var. rigida I.M. Johns!.
Figure 21A-E, 30L (RSA 580318), Figure 21 B (POM 287290); Cryptantha nubigena (Greene) Payson: Figure
6C (JEPS 90327), Figure 6D, 26F (SD 87723); Cryptantha oxygona (A. Gray) Greene: Figure 16A-D (RSA
717219), Figure 16E, 28D (RSA 709140); Cryptantha pterocarya (Torrey) Greene var. cycloptera (Greene)
J.F. Macbr.: Figure 15C (SDSU 17298); Cryptantha pterocarva (Torrey) Greene var. pterocarya. Figure 14E
(SDSU 18692), Figure 14F,G (SDSU 12435), Figure 15A, 28C (SDSU 5500), Figure 15B (SDSU 17304);
Cryptantha pterocarya (Torrey) Greene var. purpusii Jeps.: Figure 15 D,E, 28F (SD 91861); Cryptantha
racemosa (S. Watson) Greene: Figure 13D (SDSU 5494), Figure 13E,G, 28G (SD 42216), Figure 13F
(SDSU 18710); Cryptantha recurvata Coville: Figure 19D (RSA 370268), Figure 19E, 30M (RSA 250147);
Cryptantha roosiorum Munz: Figure 6A,B, 26G (RSA 346621); Cryptantha scoparia Nelson: Figure 21F,G
(RSA 641530), Figure 21H, 30N (POM 170650); Cryptantha similis K. Mathew & PH. Raven: Figure 3F-I
(RSA 272392), Figure 3J, 29B (RSA 660941); Cryptantha simulans Greene: Figure 23D,E (SD 70318), Figure
23F, 300 (SDSU 5488); Cryptantha sparsiftora (Greene) Greene: Figure 8F (SD 40827), Figure 8G, 27M (RSA
191197); Cryptantha torreyana (A. Gray) Greene: Figure 11E-I (JEPS 102382), Figure 1 1 J, 27N (SD 129490);
Cryptantha traskiae I.M. Johns!: Figure 23A,B (JEPS 93692), Figure 23C, 30P (SD 90374); Cryptantha
tumulosa (Payson) Payson: Figure 7H (SD 107276), Figure 71, 26H (SD 58351); Cryptantha utahensis (A.
Gray) Greene: Figure 14A (SDSU 18688), Figure 14B-D, 28H (SD 100153); Cryptantha virginensis (M E.
Jones) Payson: Figure 7E (SD 107269), Figure 7F (SDSU 5501), Figure 7G, 26F (RSA 340713); Cryptantha
watsonii (A. Gray) Greene: Figure 12C (COLO 503943), Figure 12D, 270 (RSA 51987),

Become a Member of
Southern California Botanists

Today!

| | Individual (family)

$15.00

| | New Member

| | Organization

$25.00

| | Renewal

Name

Address 1

Address 2

City

State

Zip Code Phone (optional)

Email address

In addition, I would like to give $ to help support SCB.

Make Checks payable to Southern California Botanists and mail to:

Treasurer, Southern California Botanists
Rancho Santa Ana Botanic Garden
1500 North College Avenue
Claremont, C A. 91711

Southern California Botanists, Inc.

- Founded 1927-

http.V/www.socalbot. org

Membership, Subscriptions, and Back Issues

Individual and Family Memberships in SCB are $15 per calendar year domes-
tic, and $20 per year to foreign addresses. Membership includes two issues of
CROSSOSOMA, and 5 or 6 issues of Leaflets , the newsletter of SCB. Leaflets
provides time-dated information on activities and events that may be of interest to
our membership. A subscription to CROSSOSOMA is available to libraries and
institutions at the domestic rate of $25 per calendar year, and $30 to foriegn insti-
tutions. Back issues (Volume 18 - present) are available for $5 each, or $10 for
the volume, postpaid. Prior to Volume 18, CROSSOSOMA included time-dated
notices to the membership and was published six times a year. These back issues
of Volumes 1 - 17 are $1 each, or $6 per volume, postpaid. Some back issues that
are out of stock may be provided as photocopies.

Available SCB Special Publications

No. 1 A Flora of the Santa Rosa Plateau , by Earl W. Lathrop and Robert F.
Thorne, 30 pp $7.00

No. 3 Endangered Plant Communities of Southern California , Proceedings of the
15 th Annual SCB Symposium, edited by Allan A. Schoenherr, 114 pp $12.00

No. 4 Flora and Ecology of the Santa Monica Mountains , 194 pp $40.00

The following article has been reprinted separately, with covers, and is available
for plant collecting workshops:

Herbarium Specimens as Documents: Purposes and General Collecting Tech-
niques , by T. S. Ross [from Crossosoma 22(1): 3-39, 1996]

$3.95 each; 10 for $22.50

Prices include California State sales tax, handling, and domestic postage.

Applications for membership, requests for purchases of Special Publications and
back issues, name or address corrections, and requests for replacement of lost or
damaged CROSSOSOMA issues should be sent to:

Southern California Botanists, Rancho Santa Ana Botanic Garden
1500 North College Avenue, Claremont, CA. 91711, USA

Consult our website for current contact information: http://www.socalbot.org
Make checks out to Southern California Botanists, or SCB.

Index to Cryptantha of Southern California

TEXT

IMAGES

KEY to Cryptantha Groups

p. 10

C. qffinis

P-21

22

C. angustifolia

P-35..

37

C. barbigera var. barbigera

p. 49

48

C. barbigera var. fergusoneae

p. 45 ..

46

C. cinerea var. abortiva

P-15

16

C. circumscissa var. circumscissa

P-11 -

12

C. clevelandii var. clevelandii

P- 25

26

C. clevelandii var.florosa

P-25..

27

C. clokeyi

p. 47

47

C. confertiflora

P-15-.

16

C. corollata

p. 38

41

C. costata

P-35-

36

C. decipiens

p. 38

40

C. dumetorum

p. 38 ..

39

C. echinella

p. 49

48

C.flaccida

P-21 ••

22

C.flavoculata

P-15

16

C. ganderi

p. 23 ..

24

C. glomeriflora

P-21

22

C. gracilis

p. 23 ..

26

C. hoffmannii

P-17

19

C. holoptera

P-31-

32

C. humilis

P-17

18

C. inaequata

P-35..

37

C. intermedia var. intermedia

p. 45

44

C. leiocarpa

P-23 ..

24

C. lepida

P-13

14

C. maritima var. maritima

P-23 ..

24, 40

C. micrantha

P-13

14

C. micromeres

p. 38 ..

39

C. microstachys

p. 25

26

C. mohavensis

p. 25 ..

28

C. muricata var. denticulata

p. 42

44

C. muricata var. jonesii

p. 42 ..

44

C. muricata var. muricata

p. 45

44

C. nemaclada

p. 23 ..

24

C. nevadensis var. nevadensis

p. 38

41

C. nevadensis var. rigida

p. 42 ..

43

C. nubigena

P-17

18

C. oxygona

P-35..

36

C. pterocarya var. cycloptera

P- 35

34

C. pterocarya var. pterocarya

P-31-

33, 34

C. pterocarya var. purpusii

p. 31

34

C. racemosa

P-31-

32

C. recurvata

p. 38

40

C. roosiorum

P- 17 -

18

C. scoparia

p. 42

43

C. similis

P-11 -

12

C. simulans

p. 45

47

C. sparsiflora

p. 23 ..

22

C. torreyana

p. 25

27

C. traskiae

p. 45 ..

46

C. tumulosa

P-17

19

C. utahensis

p. 31 ..

33

C. virginensis

P-17

19

C. watsonii

p. 29 ..

28

Front Cover:

Nutlet dorsal images of all 49 species and 56 taxa (including varieties)

of Cryptantha in Southern California.

All images to scale.