©The World Botanical Associates Web Page
Prepared by Richard W. Spjut
May 2004; Oct 2006, June 2007, Jan 2011, Feb 2012, Jan 2013, Mar 2013,
July 2014, April-May 2015, Jun 2016, Jan 2024, Feb 2024


Ephedra antisyphilitica
Sanderson Co., TX
Spjut & Marin15146, Nov 2002




Ephedra arenicola
Piute Mt Region, Kern Co., CA
Squirrel Canyon, Apr 2014.compared  with isotype (US) from Apache Co, AZ to show similarity in the distinctive 1-3 nerved seed cone bracts, and with E. aspera to show similarity to leaves..


Ephedra aspera
Brewster Co., Big Bend Region,
Black Gap Wildlife Refuge, TX. |
Spjut & Marin 14444,
Oct 2001


Ephedra aspera
CA: E Sierra Nevada, Inyo Co.,
SW of Bishop, Coyote Valley, Oct 2006



Ephedra californica
Kern Co., CA.  Mouth of the Kern River Canyon, near top of river bank, 26 Mar 2012.  Right image of stems from steep banks east of Caliente, July 2016


Ephedra foliata
Southern  Sierra Nevada, Kern Co.,
Pacific Crest Trail, northeast of Tehachapi,
Juniper-Joshua Tree Woodland,
5,000 ft.7 June 2012.

Ephedra foliata
Cultivated: University of California Davis  Arboretum. Wikimedia Commons (Ephedra sp.) last accessed Jan 22,  2024. Cones appearing terminal, whorled branches lacking.


Ephedra foliata
 Piute Mountains, Kern Co.:  Base of limestone rock on steep west-facing slope Erskine Canyon, May 2012.  Cones at ends of short to long stalk-like branches. Native to northern Africa, southern Europe and southwestern Asia. Compare with E. foliata, Flora of Qatar online


Ephedra californica
Vizcaíno Desert, BCN
Third photo shows branch with
three persistent recurved leaves.

Ephedra californica
Carrizo Plain, Kern Co., CNPS
Kern Chapter Field Trip, 14 March 2015


Ephedra viridis
CA: Kern Co. SE Sierra Nevada,
Jawbone Canyon-Kelso Valley Rd,
12 May 2011


Ephedra funerea
CA: Death Valley NP, Dante's View
Feb 2012, sterile. May 2016,
pollen and seed strobili,
last image showing winged bracts

Ephedra funerea
(left and center back) occurring with
Ephedra nevadensis (center right foreground) and right. Pahrump Valley, NV, Feb 2012


Ephedra funerea

NV: S of Hwy 190, West of Death Valley Jct., 2,580 ft, SPJ-16865. Third image:
Death Valley Pass, Feb 2012 


Ephedra funerea
NV: Bob  Roud Memorial Hwy,
2,730 ft, Feb 2012, SPJ-16863




Ephedra funerea
Mesquite Mts., Mojave Desert, CA
Apr 2005


Ephedra nevadensis
5 mi N of Walker Lake, NV
June 2007


Ephedra nevadensis
E Sierra Nevada, Kern Co., |
Short Canyon, CA
Plant photo Feb 2012.  Pollen cones photo Apr 2019 from same or different plant.


Ephedra nevadensis
Walker Lake, NV, Sep 2006.
 Plants generally intricately and shortly
branched, stems grayish-green to glaucous.



Ephedra nevadensis
Inyo Co., CA. Northeast base of
Coso Mountains
7 May 2015


Ephedra nevadensis
S Sierra Nevada, Kern Co.,
Pacific Crest Trail, northeast of Tehachapi,
Juniper-Joshua Tree Woodland, 5,000 ft
7 June 2012.


Ephedra nevadensis
E Sierra Nevada near Bishop, CA
May 2008.  Long stalked cones in left image.


Ephedra cf. nevadensis
S Sierra Nevada, Kern Co.,
West Mojave Desert, near Mojave, May 2016. Stems notably sticky. Seed cones on short curved stalks, bracts incurved.

Ephedra cf. nevadensis
S Sierra Nevada, Kern Co.,
Kern River, rocky bench, near southern shore of Lake Isabella, 22 Mar 2012. Close up of specimen scanned 9 May 2014. Note woody stolon network from which erect photosynthetic stems arise over a broad area. Fourth image shows E. cf. nevadensis on left, and E. cf. viridis on right. See also E. cf. viridis below which grows with the species at same location. 


Ephedra torreyana
Valley of the Gods, Utah
20 May 2008 


Ephedra torreyana
Glen Canyon Natl. Park, Utah
21 May 2008



Ephedra trifurca
New Mexico: Left Hidalgo Co., near Hachita, right, Dona Ana Co.; Oct 2007




Ephedra viridis
S Sierra Nevada, Kern Co.,
Piute Mt., CA. Apr 2005


Ephedra viridis
Mesquite Mts., Mojave Desert, CA
Apr 2005. Plants generally with erect yellowish green branches and dark brown leaf collars


Ephedra viridis
S Sierra Nevada, Kern Co., CA
Piute Mt. Region, north of Caliente. July 2015


Ephedra viridis

S Sierra Nevada, Kern Co., Piute Mt,
Saddle Springs Rd, CA.  June 2016

Ephedra cf. viridis
Kern Co.: Slopes bordering Cuddy Valley, southeastern slopes of Tecuya Mountain. .
26 June 2012


Ephedra cf. viridis
S Sierra Nevada, Kern Co.,
Kern River, rocky bench, near southern shore of Lake Isabella, May 2012
Scanned image from specimen obtained 9 May 2014 showing leaves incurved as they dry, to 5 mm long, but also shorter leaves near base of shoots, seeds 1 or 2. Last two images from Erskine Creek Canyon, south of Lake Isabella, May 2016


Ephedra viridis
Kern Co., CA: Piute Mts.,
E slopes facing Kelso Valley,
gray pine wd. 14 June 2016

Ephedra viridis
Zion Natl. Park, Utah, 23 May 2008
Leaves similar to the plants at Lake Isabella,
Kern Co., CA. 


Ephedra viridis
Kern Co., E. viridis Alliance, NE
of Tehachapi, photo from Sand
Canyon looking west, 30 April 2012


Ephedra sp. A
Kern Co., CA: Piute Mts., near Stevenson Peak on Cattani Ranch. Two plants growing near each other on the same rock. Male on right similar to E, viridis in the scale-leaves but older stem broken off is clearly square (4-angled). Female on left like E. nevadensis in wide spreading green stems, but differs in stems appearing 4-angled and in their absence of secondary branches, which in E. nevadensis often develop in whorls from a node.  Seed cones developing near base of plant on thick long stalks; seeds 2..
5 May 2010


Ephedra sp. A.
S Sierra Nevada/Piute Mts., Kern Co.,
Near Caliente on steep rocky slopes,  CNPS/TNC Chapter field trip
10 April 2013




Ephedra sp. A
Kern Co., CA: Piute Mts., near Stevenson Peak on Cattani Ranch. Plant with mature pollen cones, similar to E. virids in the yellowish green color stems but differing in that the stems are  4-angled and spread widely  and irregularly in various directions, often abruptly bent from one node to the next. 5 May 2010


Ephedra sp. A.
S Sierra Nevada/Piute Mts., Kern Co.,
Erskine Canyon, base of steep rocky slopes,
CNPS Chapter field trip
8 May 2012


Ephedra sp. B.
Inyo Co., CA, just west of Death Valley NP, road to Darwin.. Isolated shrub in low scrub. Intermediate between E. viridis and E. nevadensis. Also similar to E. cutleri, which differs by the more setaceous leaves near tips of shoots.
7 May 2015

Ephedra sp. B.
Kern Co., CA: Piute Mts., near Stevenson Peak on Cattani Ranch.  Similar to E, viridis but distinct for the leafy stems with whorls of short branchlets that appear deciduous. [Similar to CalPhoto image by Gary Monroe, identified E. viridis, from
Washoe Co., NV]. 5 May 2010.



Ephedra sp.
S Sierra Nevada, Kern Co.,
Piute Mt., CA. Apr 2005
(E. viridis x E. nevadensis?)


Ephedra sp.
Tejon Ranch Conservancy, CNPS Field Trip 17  Apr 2016
Stems flaccid. spreading; without dotted lines but with conspicuous abundant stomata; leaves opposite, green without swollen base. Pollen strobili clustered; bisporangiate anthers 4–7.

Ephedra sp.
Piute Mountains, Kern Co., CA
Squirrel Canyon. April 2014. Stoloniferous, monoecious with bisexual cone, pollen cones with creamy white sporangia. Occurring in population with E. arenicola and E. viridis.



Trees and Shrubs of Kern County (Jan 2013, May 2014, July 2015, Jan 2024)

Ephedra. Much-branched shrubs, subshrubs, or climbers, or small trees to 4.5 m (Price 1996) with simple or usually branched photosynthetic (green) stems. Basal woody stems when present erect or prostrate, the erect woody stems simple or irregularly branched above base; prostrate woody stem simple or branched forming a network from which photosynthetic (green) stems arise at regular intervals, the stems varying in color from yellow green to dark green, jointed, longitudinally grooved, often with whitish dot-like lines along the intervening ridges; branchlets appearing opposite along a main axis, or in whorls, or fascicles, or dichotomously divided. Leaves seasonal, appearing with cones or after cones, opposite and decussate (each successive pair at right angles) or whorled, scale-like, partially scarious and connate, with or without terminal green photosynthetic segment, 2–3 (-4) per node, variable on a plant, or on different plants of a species (Foster 1972), base often persistent, aging brown, collar or scar-like. Male (pollen) and female (ovulate) cones usually on separate plants (dioecious), or plants rarely monoecious, the cones usually unisexual, very rarely bisexual, or in some species plants are predominantly monoecious (Price 1996); male cones (1-) 2–10 at branch nodes, each cone with opposite or whorled sets of scale-like bracts overlapping in a graduated series, except for the lowermost (proximal), each bract overlying two fused bracteoles (calyx-like or perianth) from within which arises 2–8 “stamens,” with filaments united (sporangiophore), terminating in a cluster of 1–8 bilocular pollen sacs, often exserted slightly beyond the cone bract; female cones up to 10 per node, with fewer scales than males, uppermost (distal) pair or whorl fertile, ovules 1–4, usually 1–2 (-3) mature, each enclosed by two integuments (hardened bracteoles), elongated to apex in a style-like segment that forms a pollen chamber. Cone fruit with 1–2 or rarely 3 seeds, generally known as an arcesthida  (Spjut 1994), or may be classified as an ephedroid carpidium when dry and not winged, or apterocarpidium if winged, or an ephedroid sarcocarpidium when fleshy (white, yellow, red, purple (Spjut 2014 umpubl.). Species 55–60 (-71); ~30 in Mediterranean N Africa, Europe, and SW Asia extending east in dry subtropical and temperate regions across Asia; ~14 western N America (including Mexico), ~12 in the Andes (Price 1990). California has 7or more native and 2 nonnative. Kern County generally recognized with 3 in flora (Moe 2016), or perhaps 5 to as many 9 species may be recognized, appearing to be a center of diversity for dry bracteate fruit types of Ephedra species.  Images for 50 species on SEINet.

Note: Abbreviation cf. (confer) is indicated when plant photos and/or specimens differ from  species as circumscribed below based on their type specimen (holotype, lectotype, neotype). Taxonomic keys to species of Ephedra have been regarded artificial (Cutler 1939). Populations studies are needed to identify morphological character attributes that relate to molecular differences. This is complicated by polyploidy that is reportedly common in Ephedra (Ickert-Bond et al. 2020; Hu et al. 2021; Rydin et al. 2021; Yu et al. 2023).  

     The genus Ephedra (Ephedraceae) is a gymnosperm whose species are commonly known as joint firs. Fossilized ovulate cones date back to early Cretaceous (Puebla et al. 2017),  ~100 million years (mya), while pollen date earlier to late Triassic, 200 mya (Puebla et al. 2017, citations); yet, DNA studies of the extant species indicate a slow evolution only since the Eocene (~34 mya). Despite the long evolutionary history of Ephedra, only about 50 (56 -71) species are accepted of the more than 100 species that have been described (Tropicos, 285 names under Ephedra). The DNA phylogeny indicates Ephedra species dispersed from the Mediterranean (Africa-Europe) east to Asia, then on to North America (NA) and from there to South America (Ickert Bond and Renner 2015, Ickert-Bond 2004; Rydin et al. 2021). Although the NA route via Beringia seems reasonable, the Tertiary occurrences of pollen in eastern and southeastern US (PALYNODATA) could also indicate an Europe-NA direction of dispersal as suggested for Taxus (Spjut 2007). However, this may also be a secondary route of dispersal from an earlier presence of Ephedra or Ephedra-like species reported from seed and pollen found in Portugal and eastern North America" during the early Cretaceous (Rydin et al. 2009). The age gap between Cretaceous fossils and later Tertiary diversification might be best explained by the Cretaceous-Tertiary extinction (Bolinder et al. 2016) followed by a second Eocene-Oligocene extinction from which the surviving species in the Mediterranean regions then diversified and spread.

     Phylogenetic studies of Ephedra have not fully resolved species relationships using plastid and nuclear ribosomal DNA data (Rydin et al. 2021; Ickert-Bond 2004), which have included representative individuals of most species. Least resolved are the southwestern North American species.  It has been concluded that "many species appear affected by a history of hybridization/introgression and/or polyploidy, but other processes may result in similar patterns and reasons for the detected incongruence's must be further analyzed, preferably using population-level sampling and low-copy nuclear data" (Rydin et al. 2021).

     Ephedra species have been used for approximately 5,000 years in Chinese medicine.  Perhaps most documented is “Ma-huang” that includes E. sinicaE. equisetina, and E. intermedia, the stems employed for treating fever, nasal congestion, and asthma (Caveney et al. 2001). In North  America, seven species have been used medicinally by native Americans (Moerman 1998).  Ephedra californica and E. nevadensis have been used in Baja California for treating kidney ailments (Villanueva-Almanza 2011).  The alkaloid ephedrine has been employed for treating asthma, while also used as a stimulant by athletes (Mabberley 1987). Ephedrine is commercially extracted from stems of Eurasian species; but not the New World species because they lack ephedrine alkaloids.  A review of recent studies on biological activity of Ephedra extracts is given by Elhadef et al. (2020). Caveney et a. (2001) also reported that the "Distachyae group of Eurasian species (Freitag and Maier-Stolte, 1994)...contains the richest natural source of ephedrine and pseudoephedrine" and that the "E. major (syn. E. distachya) complex in the group is rich in both ephedrine and pseudoephedrine (Qazilbach, 1971)."

     Seventh-three (73) extracts of Ephedra species were screened by the National Cancer Institute during 1960–1982. Seven were found active from samples collected in Pakistan, Turkey, Arizona, and California. Activity was mostly from aqueous extracts in tumors sensitive to tannins.

     Neuroactive amino acids with cyclopropyl ring structures and quinoline-related tryptophan derivative—that have been reported in Eurasian and New World species—may account for their historical use in Old and New World traditional medicines (Caveney et al. 2001). Also, a diversity of endophytes thrive in the Ephedra stems that include gall midges (Lasioptera spp., Boeklen & Hoffman 1993) and bioactive fungi (Huang et al. 2008). Fusarium oxysporum was isolated from Ephedra fasciculata and found to contain a depsipeptide (beauvericin 29) active against four different cancer cell lines, NCI-H460 (human non-small-cell lung cancer), MIA Pa Ca-2 (human pancreatic carcinoma), MCF-7 (human breast cancer) and SF-268 human CNS cancer—glioma (Turbyville et al. 2006). Another fungal endophyte, Chaetomium chiversii, isolated from the same Ephedra species, contained radicicol 38 that has antiproliferative activity against breast cancer cell line MCF-7 (Zhan et al. 2007).

     The taxonomy of the California Ephedra by Ickert-Bond (2012 in JM2; eflora accessed 12/13/2023) expands previous interpretations of Ephedra funerea and E. aspera by Cutler (1939). for  example E. funerea "Death Valley Region, California to Nevada" (Cutler 1939), Munz & Keck (1959, Death Valley region; sw Nev), Stevenson (1993 in FNA, not detailed) and Griffin (1993 in JM1, "DMoj; to w NV").  The same characters still apply with leaves and cones. Here other character attributes are recognized  for Kern County plants such as seasonal differences in development of leaves and cones; for example, leaves appear with cones in Ephedra cf. viridis plants near Lake Isabella or after cones in plants near Caliente (Spjut 2015).  This is also seen in angiosperms in which species of willows (Salix) differentiated by whether catkins develop before leaves or with  the leaves. Coning periods for North American species have been reported (Stevenson in FNA 1993), but not for seasonal development of leaves.  Examples of leaf variation can be seen on SEINet specimens for E. aspera,  Correll & Johnston 18332 (det. B. L. Turner) collected in Big Bend Natl. Park, Chisos Mtns., July 22 (1957), and by E. Palmer 1288 (syntype, det. S. Ickert-Bond) collected in Mexico, Feb-Oct 1880. 

      Seed characters such as number per cone (1 or 2–3) and their shape help  differentiate species of female plants, while number of microsporangia seems to have little taxonomic utility at the species level.  The number of male strobili per node, which does not always correlate with the number of leaves per node, appears variable in a species, or perhaps the variation may relate to different species rather than a variable feature of a species.

      Cutler (1939) reported species of Ephedra can be identified by stem anatomical characters such as number of vascular bundles, number of hypodermal fiber strands, number of cells in a fiber strand, and number of stomata per sq mm; however, counting stomata per sq mm such as 108 cited for E. torreyana vs. 84 for E. funerea, for example, does not seem practical, while species variation of these character features were not described.  Pant and Verma (1974)  also reported on Ephedra stomata data  that seems to support Cutler's (1939) observations in that species can be distinguished by number of stomata bands per stomatiferous internodal and nodal areas that ranged  from 1–4 bands  in E. alata , or up to 19 bands in E. chilensis and 17 in E. californica. Variation in papillae on subsidiary cells in the internodal and nodal regions as described by Pant and Verma (1964) also appear to have taxonomic value. Species differences in number of leaf stomata rows and of adjacent rows of marginal cells lacking papillae in the conifer genus Taxus were employed by Spjut (2007) to distinguish its species. Nuclear and chloroplast DNA phylogenies largely support Spjut's taxonomy of Taxus.

      Although California is reported to have six native species (Ickert-Bond in JM2), E. torreyana is an additional species that has been found in the Death Valley region (Sanders et al., CCH specimens, 2011, 2012). Also, Villanueva-Almanza and Fonseca  (2011) reported E. antisyphilitica in Mexico near the California border—Mexicali and Ensenada; thus, it too may be in California.  Ephedra torreyana, which reportedly hybridizes with other species (Stevenson 1993), is distinguished by the midnerve on ovulate bracts pale orange sword-shaped (lanceolate) nerve (or orange yellow to greenish yellow, Cutler 1939) on the ovulate bracts and scale-leaves, in contrast to the broad elliptical  pale orange-red nerve in seed cones of E. funerea.  Leaves of E. trifurca, which has a similar orange colored nerve, differ in being narrower and longer and not recurved  from the tip with age as in E. torreyana; instead, it shreds with age as in E. funerea. Ephedra trifurca is also be distinguished by the rigid branches ending in a sharp spine and by the relatively long linear leaves. Ephedra antisyphilitica, not likely to occur in Kern County, is distinguished by fleshy red bracts that surround the seed.

       In Kern County, three species are fairly common, Ephedra californica, E. nevadensis, and E. viridis. A helpful character to separate them is the angle of branching, nearly at right angles in E. nevadensis, ~30° in E. viridis in which branches appear erect and nearly parallel to one another.  The angle of branching in E. californica, ~45° (Cutler 1939; Stevenson 1993), is thus intermediate between the two other species, while their erect to spreading growth  resembles E. viridis; however, the leaves, when present—that occur in three's—easily identifies E. californica.  Nevertheless, the reported occurrences of E. californica east of the Southern Sierra foothills need study, especially where the species overlaps with E. funerea in Inyo County. The latter can be distinguished by grayish green color of the stem ending in sharp spine. Color of stems also separates E. nevadensis, gray to grayish green, from E. viridis, green to yellow green.

     Ephedra aspera has been reported near the Kern County line along Hwy 58, while I have only seen E. nevadensis in the region—where locally common in creosote scrub (Larrea tridentata shrubland alliance). Shrubs with well-developed erect woody stems and orange pollen cones along the Pacific Crest Trail near Walker Pass and in upper Jawbone Canyon have persistent leaf characteristics similar to an illustration of E. aspera in Powell (1998), described in the key below as cuspidate-recurved, while another kind of leaf also seems to develop as seen in the isolectotype (US), but not evident in a  male syntype.   Variation in leaf development on an Ephedra plant can vary considerably (Ickert-Bond and Renner 2015), and for some plants leaf development appears to be of two kinds such as shown for E. viridis near Lake Isabella.

     A fourth species in Kern County, Ephedra arenicola, was discovered in Squirrel Canyon in the Piute Mountains (Spjut 2015). The identification was made by comparison to the type specimen collected from Apache County, AZ. The species was described by Cutler as a hybrid between E. torreyana and “E. coryi var. viscida” [E. cutleri]; however, the unusual character feature of the distal bracts of seed cones appearing with raised mid and lateral nerves, and its disjunct geographical disjunction, would seem to justify species status without hybrid designation. The leaves of the E. arenicola type are remarkably similar to those seen on the E. aspera type.  Based on this and the rough appearance of the dark green stems, E. arenicola would seem to be a distinct species that was more widely distributed and has since evidently hybridized with E. aspera in Apache County, AZ and E. viridis in Kern County, CA. Although originally thought to be a hybrid, the occurrence of a second collection from a distant disjunct location, is viewed as an amendment to the circumscription of a species, thus the hybrid designation is not necessary; it is not uncommon for species to have their circumscriptions amended based records from new geographical areas. 

     Ephedra funerea has also been reported in Kern County based on identification from a software photo determination, Plantnet (Calflora with reference to iNatualist addition 11/26/2008, but photo not included, observer Del Faverno (Bristlecone Chapter, location accuracy 50,000 sq km). Another Kern County collection reported by Bartholomew and Boufford 04/20/1985, 28 km E of Interstate Highway 5 on State Highway 138 (CCH1, herbarium specimen) was searched for in 2013 without success . This location, which is about 3 km south of the Kern County line in Los Angeles County, appears to have had its vegetation cleared at sometime in the past.

          An ephedra found along Erskine Creek in the Piute Mountains and in the southern Sierra Nevada along the Pacific Crest Trail northeast of Tehachapi has a vine-like habit with long pedunculate (stalked) cones (Erskine Creek plants), originally thought to be E. viridis. These features suggest E. pedunculata, a species known from southern Texas and northern Mexico; however, upon further review (since 2016) as of Dec 2023, it is referred to Ephedra foliata, a widely distributed Mediterranean species that extends from the desert regions of northern Africa to western India. Here it may be noted that the Mediterranean E. foeminea, an Eurasian-north African species, was discovered growing in Santa Barbara County where possibly naturalized (CCH2, identified by Ickert-Bond). It is distinguished from all other Kern County species by the red fleshy bracts (JM2), which are likely dispersed by birds (see photo of species by Ori Fragman-Sapir, JungleDragon.com). 

     Along the Kern River near Lake Isabella is an Ephedra with an erect pale green stems arising from a much branched network of horizontal, rhizomatous-like, woody stems, spreading over 10 m, tentatively assigned to Ephedra cf. nevadensis. It differs from the typical form of the Nevada ephedra by the flexuous (not rigid) whorled branches not as wide spreading. Occurring with this (Ephedra cf. nevadensis) is Ephedra cf. viridis, differing slightly from the common form by shortly stalked seed cones, aborted development of one of two seeds per cone, and by  development of long narrow linear green leaf segments, either from the axils of nonphotosynthethic scale-leaves, or from their tips, which appear to soon wither and drop.  These plants may possibly be hybrids.  They both cone during the May.

          Two proposed species of Ephedra in Kern County are referred to as Ephedra sp. A (Erskine Creek in the Piute Mountains, and north of Caliente below Stevenson Peak, and Ephedra sp. B photographed in Box Canyon near Stevenson Peak. The latter also recognized in Inyo County near Darwin (pers. obs. May 2015), and from images on Calphotos of one reportedly taken in Washoe County, NV.  Young branches or branchlets were frequently present on Arizona specimens reviewed of E. fasciculata on SEINet. 

         The following key is tentative due to species concepts in Ephedra that have varied among the treatments over time (Cutler 1939; Munz & Keck 1959; Stevenson in FNA 1993; Griffin in JM1 1993; Ickert-Bond in JM2).  Although the number of scale-leaves imply a diagnostic feature for recognizing species, its application probably does not reflect the natural relationships as evident from  their variation in shape, texture, color, connation, and persistence observed within a “species” from different locations and also from comparing species descriptions among the floristic treatments.  Further, species of Ephedra do not generally fall into the ecogeographic patterns of other plant species as noted for E. californica in MCV2.  Detailed field studies during coning periods are needed.  As reported by Cutler (1939) about two-thirds of the herbarium specimens he studied could not be confidently identified, while he also indicated that his key was “artificial.”

Key to Species of Ephedra

1. Green stems creeping, vine like; branches below apex often opposite
at 90 degree angles; rare, Erskine Creek, Eastern Sierra 
Nevada along Pacific Crest Trail northeast of Tehachapi ............ Ephedra foliata

1. Green stems mostly upright, branches below apex often in whorls
at  28–90 degree angles................................................................ .......................2

    2. Green stems mostly parallel to one another, at least above mid region
 of plant; branching not more than at 45 degree angle except below mid region ..... 3

    2. Green stems wide spreading above mid region, branching at ± 45 degrees
or more .............................................................................................................. 8

3. Leaves soon deciduous, white to gray, scarcely thickened at base;
stems not closely parallel to one another, often crisscrossing; 1 of 2 seeds
often aborting
; Kern River, south end of Lake Isabella (possibly a hybrid
from growing in association with E. viridis)...................... Ephedra cf. nevadensis

3. Leaves often with persistent collar at base............................................................... 4

    4. Leaf bases yellow orange to brownish orange, often puckered;
seeds  usually two, sometimes 1 aborting, two angled,
rounded on back, flat on inner surface......................................... .........................5

    4. Leaf base mostly a thick brownish collar, or absent; seeds 1-2, variable
in shape .............................................................................................................. 7

5. Main stems with whorls of erect short leafy shoots; leaves at short
intervals, overlapping nearer apex;
cones solitary, sessile, oriented
perpendicular to stem; Piute Mtns. (Stevenson Peak, Piute Mt., also Inyo Co
near Death Valley NP and Washoe Co., NV (CalPhotos)
................ Ephedra sp. B

5. Main stems with long branches, not always in whorls; leaves at
regular intervals .................................................................................................... 6

Leaves similar throughout, tightly clasping stems, often with sheath and
    salmon-orange mid-nerve, puckered at base, connate to above middle,
    obtusely rounded to apex; seed cone bract not fleshy orange;
    widely distributed in Kern Co..................................................... Ephedra viridis
6.  Leaves on stems, with sheath and with or without long
    linear terminal green segment.................................................. Ephedra cf. viridis

6.  Leaves connate halfway, without sheath, triangular lobed
     near apex, lobes acute; seed cone bracts orange,
     apparently rare, Four Corners, not in Kern Co, sandy
     desert, wTX ............................................................................... Ephedra coryi 

 7.  Distal bracts of seed cone (1-) 3-(5) nerved, seeds 2;
     Apache Co., ne AZ near  Dennehotsco (
leaves connate to near apex),
     Kern Co. (Squirrel Canyon, leaves deciduous)...................... Ephedra arenicola

 7.  Distal bracts of seed cone not appearing nerved, margins scarious;
  seed 1; "Ft. Tejon and vicinity," juniper woodland between Tehachapi
  and Mohave................................................................................  Ephedra aspera

       8. Stems ±4-angled; yellowish green; nodes with dark
     brown collar; Piute Mts. (Stevenson Peak, near
     Caliente, Erskine Creek ................................................................ Ephedra sp. A

       8. Stems mostly round in outline, grayish green; scale-leaves gray to
white-scarious, deciduous; node collar white with
    narrow strip of ................................................................................................... 9

9. Stems rigid, grayish green; branches in whorls diverging at ± 60 degrees.................... 10

9. Stems flexuous or rigid, dark green to yellow green; branches in whorls or not,
diverging at < 60 degrees ..................... .................................................................11

     10. Seeds 2, broadly ellipsoid, 6–9 mm, ~2× longer than wide,
 brownish, flattened on inner surface; scale-leaves in 2's;
 branch tips not spiny; mostly Mojave Desert and desert slopes
 of Sierra Nevada ........................................................... ......... Ephedra nevadensis

    10. Seed 1 (-3), narrow ellipsoid to bottle shaped, ~3× longer
 than wide; scale-leaves in 3's; branches spine tipped......................... Ephedra funerea

11.   Scale-leaves in 3's (whorled)....................................................... Ephedra californica

11.  Scale-leaves opposite (in 2's, or 2's and 3's).............................................................. 12


12.  Stems yellow green to dark green; pollen cones often stalked, seed cones
shortly stalked; seeds usually one .............................................................................. 13

12.  Stems bright yellow green; pollen and seed cones stalked; seeds two.......................... 14 


13.  Stems branched at wide angles; branches often flexuous or appearing
almost filiform;  persistent parts of leaves short oval white to gray,
above collar; stems slender, cone bracts pale yellow with narrow dark
mid area; seeds 1(-2), conical, widest below mid region, long tapered to
pointed apex, longitudinally furrowed; possibly in Kern County;
eCA (
seed  <8 mm may be referred to E. clokeyi, and by cone bracts ,
widest near base); wAZ, sNV, sUT................................................. Ephedra fasciculata

13. Branches of main stems  ascending upwards, closely parallel; 
persistent scale-leaves brown cuspidate-recurved or gray-white,
or connate-clasping with short triangular acute tapered lobes
(one or both present on same plant); seeds 1(-2), widest near 
base, tapered to an obtusely rounded apex, mostly smooth; Kern County;
"Ft. Tejon and vicinity"; Sonoran, southern Mojave, and Chihuahua
deserts; between Tehachapi and Mojave (Abrams & McGregor at HUH
may be regarded E. clokelyi)................................................................. Ephedra aspera

14. Pollen cones stalked, the stalks longer towards apex of young 
shoots; seed cone bracts fleshy; orange; possibly in Kern County
but generally considered rare, sandy desert, wTX....................................... Ephedra coryi

14.   Scale-leaves long tapered (setaceous) to apex, often persistent at tips of
stems and below, cuspidate; photosynthetic stems often bright greenish
yellow; seed scales with dark central oval region; seeds dark purplish or
nearly black, exserted; plants often with many whorls of  short
shoots, possibly in Kern Co; Four Corners .............................................. Ephedra cutleri


Ephedra arenicola Cutler 1939 (“X E. arenicola,” putative hybrid between E. torreyana and E. cutleri). Shrub to 1 m, with whorled branches, spreading ~35; leaves persistent and opposite in type specimen, of two kinds as seen in E. aspera, (1) connate and slightly wedge-shaped with a prominent mid nerve, shortly broad deltoid to apex, and (2) cuspidate without mid nerve, acutely tapered to recurved apex; seed cones in pairs; lower seed bracts keeled, upper bracts 3 nerved; seeds acutely tapered to apex.
from Apache Co., AZ, 5 miles south of  Dennehotsco. Also recognized here to occur in Squirrel Canyon, Kern County; specimen from one plant collected, remarkably similar in development of paired seed cones, seed shape, and the strongly 1-3 nerved bracts (see images above).

Ephedra aspera S. Watson 1883. [Ephedra nevadensis S. Watson 1871 var. aspera (S. Watson) L. D. Benson 1943]. Includes Ephedra clokeyi Cutler 1939 [Ephedra fasciculata A. Nelson 1934 var. clokeyi (H.C. Cutler) Clokey 1945]. Rough ephedra, boundary ephedra. Shrubs with stiff erect closely parallel branches, to 1.5 m; older stems with fissured bark, photosynthetic shoots rather thick, ~ 3 mm diam, dark green, yellow with age, rough and papillate, smooth and  glaucous between ridges, branching opposite or in whorls of 3 or 4, at angles of ~35, whorls of branches often at frequent intervals; scale-leaves in 2’s (rarely 3 per node), 1–3.5 mm long, united half or more of their length, persistent in the typical form (with seed cones), or the sheath splitting, fibrous, swollen at base; leaf collar brownish; pollen cones yellow to orange, drying reddish brown, with broad rounded (elliptical) bracts; seed 1, ± broad ellipsoid, tapering to apex from above mid region, or trigonal, 5–8 mm, tan to chestnut brown. 
Widely distributed in deserts of the southwestern North America, mostly below 5,000 ft, generally Mojave and Sonoran  deserts as far south as the Magdalena Desert in Baja California (Turner et al. 1995), to the Chihuahua Chihuahua Desert Texas and south into Mexico, also reported in the western Great Basin Desert. Type from the Sierra Madre in Coahuila, 40 miles south of Saltillo, Mexico [Palmer 1288, implied by Watson but not the only specimen referred to; lectotype designated by Cutler (1939) with reference to specimen at MO (Tropicos.org. Missouri Botanical Garden. 22 May 2015 <http://www.tropicos.org/Image/45632>, low resolution, sheet with 2 specimens with different labels, specimen appearing to have seed cones annotated by Cutler as “TYPE ”); isolectotypes GH, NY UC, US (high resolution image, 2 specimens with two labels, one with seed cone has persistent connate leaves, annotated syntype by Ickert-Bond, May 2002); the other with pollen cones, Palmer s.n., 1880, with handwritten annotation, reference to Cutler(?), “duplicate of type,” lacks connate leaves]. Kern Co.: Ft. Tejon & vicinity, Xantus de Vasey 112, collected 1857-58 (US!, syntype for E. nevadensis, annotated E. aspera by Ickert-Bond. CCH-two records: (1) N. Cooper, 19 Apr 1949, from County line west of Kramer along US Hwy.466 (Hwy 58), 2800 ft (n.v.); (2) between Tehachapi and Mojave (Abrams & McGregor, 28 Jun1908, cited by Cutler 1939, with pollen cones, no seed (HUH!). Plants photographed along the Jawbone-Kelso Creek Canyon Road, 12  May 2011, have the persistent cuspidate leaves but the plants only had pollen cones. Ephedra aspera
links: SEINet, Calflora, Calscape.

            Ephedra aspera—as treated by Ickert-Bond (JM2)—includes E. fasciculata previously recognized in Munz & Keck (with reference to Cutler 1939) as a low often prostrate plant with flexuous branches, in sharp contrast to E. aspera they characterized as an erect plant with rigid branchlets. However, Ephedra aspera was not recognized to occur in California in earlier floras (Jepson, McMinn, Abrams), whereas E. fasciculata—that was recognized by Griffin (JM1)—was referred to as a synonym of E. aspera by Ickert-Bond (JM2); but is among 54 species recognized in Ickert-Bond and Renner (2015).  Ephedra fasciculata and the related E. clokeyi were distinguished from E. aspera by elliptical shape of ovulate scales and by the dark brown furrowed seeds in contrast to the orbicular scales and non-furrowed light brown seeds of E. aspera; E. fasciculata was then separated by seed length, 8–13 mm, in contrast to 5–8 mm for E. clokeyi (Cronquist et al. 1972; Munz & Keck 1959;  Shreeve & Wiggins 1964).  Seed of E. fasciculata in most cone specimens on SEINet appear narrower and acutely tapered to apex from below mid region. This is in contrast to a more plump (oval in outline) in E. aspera specimens.  A key feature of E. fasciculata is the furrowed seed (E. fasciculata SEINet image by Anthony Menzoza), which often is often clearly discernible in specimen images due to much of the seed being obscured by cone bracts. Plants without cones may be recognized by persistent cuspidate recurved leaves, in contrast white parts often seen above the collar in E. fasciculata. California plants may be distinguished as E. clokelyi.

            Although the circumscription of E. aspera has varied among different floristic treatments, key features in common are the usually single seed and paired leaves in contrast to the two seeded “cones” of  E. nevadensis and E. viridis, also with paired (opposite) leaves, in further contrast to the whorled leaf arrangement in other species.  Branchlets of E. aspera are often numerous—in fascicles—growing erect, closely parallel, and rigid.  Ephedra viridis is most similar in habit, and difficult to distinguish in sterile leafless plants, except perhaps by its yellowish green stems.

            The type specimen of E. aspera (high resolution image at Smithsonian Online Herbarium Collections) exhibits two kinds of leaves, one appears associated with vegetative growth as seen near base of specimen where persistent leaves clasp and recurve from their tips; the other more sheathlike and connate to ~7/8 of their length, which can be observed at base of seed shoots and on young shoots, in the axils of old leaves on older shoots as well as at apex of shoots. This latter type of scale-leaf is distinctive but rarely seen in herbarium specimens identified E. aspera; but see Correll & Johnston 18332, reported to have been collected 22 July 1957 (LL, TEX, on SEINet).  See also Ickert-Bond and Renner (2016, Fig. 4)

            Ephedra peninsularis I. M. Johnston 1922, described from Magdalena Island, was interpreted to be widely distributed in Baja California; however, it has been generally treated as a synonym of E. aspera. Its type (isotype, GH, HUH & Libraries, image) shows less tapered pollen cones with well exserted anthers.  An Ephedra I collected on the Vizcaíno Peninsula near Puerto Nuevo was noted to be unusual for the blood red sap.

            Not surprisingly, the geographical ranges of these species have also varied according to different authorities. Stevenson  (FNA 1993) showed E. aspera to reach its northernmost distribution along the US/Mexico boundary—from California to southwestern Texas; hence, the common name boundary ephedra. Griffin (JM1 1993) indicated a much wider range for E. aspera—a more northern occurrence, extending into the Mojave Desert as previously recognized by Munz & Keck (1959). The treatment by Ickert-Bond (JM2) extends the range further north—into the Great Basin Desert of the California flora for which there is a specimen in CCH (Ickert-Bond annotation)—that reported its collection site 25 miles east of Lee Vining, 2296 m.  Ephedra aspera also occurs southwards into mainland Mexico and along the Pacific Coast in Baja California Sur (Turner et al. 1995; see also CIRH).

Ephedra californica S. Watson 1879. Desert tea, California joint fir. Shrub with numerous densely tufted erect green stems (caespitose) from a short woody base, or small tree with a definite single trunk; photosynthetic stems (branchlets) green to yellowish green, most branchlets spreading ~45°, often not closely parallel to one another; scale-leaves in 3's (rarely 2, or 4), united at first, with a greenish or reddish brown medial thicker part, soon splitting with the tips curving backwards (recurved), the basal collar bulging (typical form), darkening with age; cones usually 3 per node; seed solitary, ovoid (type), globose, ellipsoid, or obovoid (Kern Co.), 1–1.5 (-2)× as long as wide.
Scattered occurrences in diverse habitats below 4,000 ft; grasslands of Inner South Coast Ranges near Gilroy in California, coastal strand near San Diego, Peninsular Ranges in California and then south in various habitats to near Bahía Tortugas, Vizcaíno Peninsula (CIRH, Turner et al. 1995).  Also Panamint Range, eastern ranges of Death Valley near Nevada state line, sand dunes and washes in the Mojave Desert, southern Sierra Nevada in Kern Co., Tehachapi Mts., and then south to the Sonoran and Chihuahua deserts of southeastern Arizona. “Californica joint fir scrub” (alliance) recognized in MCV2 when ≥ 2% absolute cover in the shrub layer, noting also that the ecology of the species is poorly known. Type from San Diego Co., Jamul Valley. Original material includes two specimens with leaves collected by Edward Palmer in 1875, Palmer 864 & Palmer 865 mounted on one sheet (
HUH-GH!), without cones except for two seed (Palmer 864) and two immature pollen cones (Palmer 865) mounted below each specimen reported to have come from attached specimen packets, scales and seed for one cone separated, its associated specimen (Palmer 864) has leaves recurved near apex, designated "Type" by Cutler 1939, and lectotype by Ickert-Bond, May 2002, the other with leaves recurved to near mid region and its associated two pollen cones below annotated syntype by Ickert-Bond May 2002, the specimen reportedly collected from San Diego, San Diego Co.  Kern Co.: “Occasional in the California juniper belts from Recruit Canyon and the Panorama Hills in the southern Temblor Range to the east end of Cuyama Valley (where unusually large plants grow on the sand flats), and along the sandy foothills on the east side of the valley from the mouth of Kern Canyon south to Comanche Canyon in the Tejon Hills,” 73–1,300 m (CCH excluding suspected misidentifications).  Also common on steep loose sedimentary rocky banks in Caliente Creek Canyon (pers. obs.). Plants from near the Death Region shown in CCH2 under E. californica may be identified var. funerea; e.g., Annable 557 collected in the Funeral Mountains.   Ethnobotanical (Moerman): Diegueño Infusion of branches to purify blood, improve appetite, or to relieve stomachaches from eating too much, and for kidneys.

            Leaves of plants in Kern County plants such as shown here from bluffs along Caliente Creek, were not found with cones during March but in late July after a summer rain.
E. californica links to SEINet, Calflora, Calscape.

Ephedra coryi  E. L. Reed 1936. Shrub forming clumps from rhizomes, photosynthetic shoots often with whorls of long shoots, olive green; pollen and seed cones stalked; New Mexico and Texas.  Type from Brownfield, Texas, Reed. 4147, 29 Apr 1935 (US).  Lectotype designated by Cutler, Ann. Mo. Bot Gard. 26: 413, annotated by S. Ickert-Bond, May 2002.

Ephedra cutleri Peebles 1940 (new name for Ephedra coryi var. viscida Cutler 1939). Navajo ephedra. Shrub forming clumps from rhizomes, photosynthetic shoots often with whorls of short shoots, bright green, yellowing with age, viscid due to tiny transparent resin droplets; leaves opposite, setaceous, persistent, connate to 1/2, thickened at base, long tapered to apex, drawn out part 5–8 mm; pollen cones shortly stalked; seed cones on stalks 5–25 mm, generally longer towards base of shoots, scales with a broad dark greenish mid region, ¾ or more of a scale, and with narrower hyaline margin; seeds 2, elliptic in outline. Sandy and .rocky slopes and flats, Four Corners region. Type from Apache Co., AZ, 10 miles west of Rock Point. Fresh samples collected and extracted by water by the College of Pharmacy at the University of Arizona during the 1960’s showed antitumor activity, one of the entire plant collected Aug 1961 was found active Jul 1971 in Sarcoma 180 (mouse), and another collected Jul 1962, divided into root—active (Sep 1965) in Friend Virus Leukemia (mouse)—and stem-leaf—active (Apr 1965 ) in Dunning leukemia ascites (rat). Active agents unknown.

Ephedra fasciculata A. Nelson 1934.  Fasciculate ephedra. Shrubs generally broader than high, up to 1 m; the erect branches arising in whorls or as singles from horizontal branches, generally wide, spreading 30° to 45°, pale green, aging (or drying) yellow, smooth except for being grooved; leaves opposite, 1–3 mm, connate 1/2–3/4; membranous, brown, shredding and graying with age, ± persistent, obtusely tapered to apex; pollen cones 2–several per node, 4–8 mm, sessile; bracts 4–8 pairs, light yellow, obovate, 2–3 × 2 mm, membranous, bracteoles exceeding bracts; sporangiophores 3–9 mm, ¼– ¾  exserted, anthers 6–10, sessile to short-stalked (microsporangia less than 1 mm); seed cones 2 or more per node, 6–13 mm, sessile or shortly stalked; bracts 4–7 pairs, elliptic, 3–7 × 2–4 mm, membranous, light brown to green, thickened along center and at base, margins entire. Seeds 1(–2), conical, tapering from below mid region, 5–12 × 3–5 mm, light or dark brown, acutely tapered to apex, longitudinally furrowed, . Dry rocky slopes, washes, and sandy soils; 300–1200 m; AZ, CA, NV, UT.  Included under E. aspera in JM2, recognized in previous California floras and in FNA. Type from "hot dry banks of a sandy wash in low hills near Phoenix"; type specimen lacks cones, thus, the taxonomy of the species is questionable.  California specimens identified E. aspera may actually belong to E. clokeyi. This includes Abrams and McGregor specimen at HUH, collected between Tehachapi and Mojave in Kern County, which has branches spreading at wide angles and pollen cones.
(Moerman). Pima: Powdered roots applied to sores or syphilis.
Links: SEINet

Ephedra foliata Boiss. ex C.A.Meyer 1846 [Mém. Acad. Imp. Sci. Saint-Pétersbourg, Sér. 6, Sci. Math., Seconde Pt. Sci. Nat. 7(2): 297 (1846), n.v.].  [Versuch einer Monographie der Gattung Ephedra 100. 1846. (Mar 1846)!].  In Kern County a low sprawling subshrub to 15 m or more broad, and up to 1/2 m high, stems rather thin, bright green, somewhat flaccid or vine-like, terminal branching dichotomous at 45–70°, and with opposite branches at 90° below apex; leaves opposite, connate with pale green sheath, similar to some forms of E. viridis but not thickened at base, lacking rusty orange near leaf base, appearing mostly linear, reportedly 3-4 mm long, ciliate along margins (Foster 1972). Male strobili described in literature 1-3 per node; cones in Kern County plant terminal on short to long curved branches. Seed cones described to have white fleshy bracts, seeds reportedly 2, brownish black. Distribution in Kern County: three plants observed and photographed along east side of Erskine Creek, creeping loosely over talus at base of steep rocky slope. Plants not found when searched for in March 2023. Similar plants observed elsewhere in Kern County along the Pacific Crest Trail northeast of Tehachapi and near Caliente. Native geographical distribution: Generally known from dry, sandy to rocky areas and slopes, 100–1000 m; Afghanistan, Algeria, Chad, Djibouti, Egypt, Ethiopia, Gulf States, India, Iran, Iraq, Kuwait, Libya, Mauritania, Morocco, Oman, Pakistan, Palestine, Saudi Arabia, Sinai, Socotra, Somalia, Sudan, Tadzhikistan, Turkmenistan, Uzbekistan, Western Sahara, Yemen. Images showing habit and fruit of plant: Flora of Qatar, and of herbarium specimens, SEINet (Ephedra).  Type from Iran: Islamic Republic of Gilan, Aucher-Eloy, P. M. R. 5338, no date. Isotype: K (K000456219!), P (P00738820 photo); Isolectotype: BM (BM 000884470 photo!). A variable species recognized to include 12 synonyms. Also reported from a geological formation in Petrified Forest National Monument in Utah of relatively Recent geological age (Scott 1960); however, this identification has since been questioned (Norbäch-Iversson 2014).  Ephedra foeminea, a Mediterranean species reported from Santa Barbara County, differs in cones developing on short whorled branches that often coil.   

Cultivation. Ephedra foliata has been reported to be cultivated at San Marcos Growers Nursery in Santa Barbara; however, upon further review this may have been E. tweedieana, which is the only species of Ephedra reported in a search of their database inventory, a South American species that has a similar vine-like habit and branching to E. foliataEphedra tweedieana differs by the cones being sessile. An image of
Ephedra tweedieana is shown on the San Marcos Growers nursery website (accessed Dec 23, 2023). They reportedly received their plant from the Huntington Garden Conservatory in 1997. They further note that it is also cultivated in the Mildred Mathias Botanic Garden, and at the UC Davis Arboretum where it was planted along the slopes of Putah Creek as a large scale shrubby groundcover.  An image of a plant reportedly cultivated at the UC Davis Arboretum shown on Wikipedia Commons, identified Ephedra californica or Ephedra sp. (file name), is regarded here as E. foliata. It compares well with images of the Erskine Creek plants as shown above for both. Foster (1972) reported obtaining "shoots" of E. foliata plants growing at the University of California Botanical Garden Berkeley for his study of leaves; however, the species is not currently listed in their database.

Additional Comment
: The name Ephedra ciliata Fisch and Meyer reportedly published in Bull. Cl. Phys.-Math. Acad. Imp. Sci. Saint-Pétersbourg 5: 36 (1845) (
Kew Plants of the Online) suggests an earlier name for E. foliata. See also Faried et al. 2018 who reported E. ciliata to be an illegitimate name based on Kew Plants of the World Online (2017). The  reference to the Kew Online database may have been subsequently revised. Ephedra ciliata was accepted by Kew citing 1845 as publication date, and E. foliata listed as synonym (accessed 12/13/2023); however, C. J. Earle (web page date Mar 2023 for Ephedra tweedieana) noted that Meyer's dated 1845 edition was not actually published until 1847. Additionally, in Meyer's 1846 monograph of Ephedra, he expressed uncertainty in distinguishing E. foliata from E. ciliata  because he felt more study was needed to evaluate the occurrence of male cones on a female plant specimen [translation of Latin here: Ephedra foliata Boiss, Th. Kotschy plant. Exisicc. south of Persia No. 866. Female specimens with anthers not yet explained near the ruins of Persepolis in u. Kuh Ajub (mountains in Iran)].  Thus, E. foliata may differ in its monoecious habit as reported for the species on some websites, while other species of Ephedra are known to be primarily monoecious such as E. tweedieana, a South American species (Earle 2023).  Cones on one plant in Kern County included male cones both sessile and long pedunculed but not fully developed, which may be female, thus, monoecious.

Ephedra funerea Coville & C.V. Morton 1935 [E. californica var. funerea (Coville & C.V. Morton) L.D. Benson 1943]. Death Valley ephedra, Death Valley joint fir. Similar in habit to E. nevadensis in the intricate nearly right-angle branching, differing by the papery scale-leaves in 3’s, ± continuous whitish lines on ridges that give the stems their whitish green (gray) color, and by the spine-tipped branches. Seeds described as usually 1 (but up to 3), bottle-shaped (illus., Stevenson in FNA 1993) to ellipsoid (FNA). Nodal swellings often present, as also in other ephedras, caused by endophytes or gall midges.
Type from Furnace Creek Canyon, ~3,200 ft. 
This species has been considered endemic to the Death Valley region (Cutler 1939; Munz & Keck 1959); however, the treatment by Ickert-Bond (JM2) expands its range to Anza Borrego State Park and western Arizona. CCH data (specimens annotated by Ickert-Bond) suggest it occurs frequently on limestone, gypsum, margins of alkali dry lakes, and among lava rocks on summits.  Death Valley joint fir scrub provisional shrubland alliance in MCV2, the species thus appearing common in parts of the Kingston, Mesquite, and Nopah ranges. Ephedra funerea is the dominant shrub of the desert slopes surrounding Dante's View.  Kern Co. One collection reported from from Through Canyon near Inyo County line observed by Del Faverno 11/26/2008 (Bristlecone Chapter, iNaturalist, observation, Plantnet id but no images provided, also in Calflora with reference to entry by John Malpas, accessed 12/13/2023). Other collections near county line are from lower slopes of Newberry Mt., just south of Newberry off Interstate Highway 40, from Iron Mt—between Barstow and Kramer Junction—and from the Argus Mtns., and from near Gorman along Hwy 138 about 3 miles south of the Kern County line.  These reports of E. funerea remain questionable.  Ephedra funerea may be expected in the El Paso Mts. where E. nevadensis is known (Twisselmann) and in the Piute Mountains. The two species occur together near Pahrump, NV (Spjut obs. 2012), whereas in Inyo County, E. funerea may occur with E. californica.  Links: SEINet, Calflora, Calscape

.Ephedra nevadensis S. Watson 1879 (Proc. Amer. Acad. Arts 14: 298). Nevada ephedra, Nevada joint fir. Stems “pale” green, glaucous, aging gray green, rigid, densely and intricately branched, many branches in whorls spreading nearly at 90, but generally described at 45 (Cutler 1939);  leaves 2 per node, generally scarious, connate ~1/2, free portion triangular (deltoid) to pointed apex, soon deciduous, leaving a white collar around node; seed cones appearing in the spring, on a short stalk, or not stalked in which both sometimes found in the same population, often clustered among whorled branches, bracts glaucous, largest broadly ovate, concave; seeds usually 2, elliptical in outline, outer side hemispherical, inner flat, tapering obtusely to short rounded to pointed apex, maturing brown. 
Primarily deserts of California to Arizona, Utah, and Oregon.  Nevada joint fir scrub recognized in MCV2 when dominant and ≥ 2% absolute cover in the shrub canopy, ranging in elevation from 1,000–1,800 m.  Type from Smoky Valley, 5500 ft (1676 m), Esmeralda Co. west of Tonopah, NV [Type, S. Watson 1108, July 1868, with pollen cones, designated by Cutler 1939 ("G Type", pro.parte!), GH00022597, designated lectotype by Ickert Bond, May 2002. Type sheet at GH includes three additional specimens, all determined by E. viridis by I.M. Johnston (1922) and annotated E. viridis by Ickert-Bond (May 2002;
; they were collected from (1) “Pah Ute Mountains,” 5,000 ft, with mature seed, (2) Eagle Lake, Lassen Co., CA, w/o cones, and (3) location undetermined, w/o cones.  Kern Co.:  Frequent in the Mojave Desert region of Kern Co on rocky slopes of the Eastern Sierra Nevada, especially along the Pacific Crest Trail (PCT) northeast of Cameron where may be recognized as an alliance where alliances of California juniper and sagebrush are less distinct. Also frequent in broad sandy-gravels wash creosote scrub east of California City near the San Bernardino County line. Twisselmann reported Nevada ephedra as occasional to scarce on rocky or gravelly slopes in the Rand and El Paso mountains, and on sand dunes around the Muroc dry lakes. 609–1524 m (CCH) . Image above from PCT near Cameron, ~1,500 m. Forma rosea Cutler (1939) distinguished by the “roseate bracts” and seeds < 5mm; type from Pyramid Lake, NV; and one other specimen reported from Willow Springs in Kern County (Cutler 1939). A photo of E. cf. nevadensis shown above taken in May 2016 near Mojave seems to differ by the bracts curving inwards over seeds, the cone fruit appearing much like a capsule, and the seeds also differ in their shape by appearing almost as wide as high, while other plants in Kern County often differ from the typical form in appearing green instead of glaucous and having longer recurved branches not always developing from whorls of branches.  A photo taken April 2005 of a plant on Piute Mt., identified Ephedra sp., appears to be a hybrid, E. viridis x E. nevadensis.  Links: SEINet, Calflora, Calscape.
Ethnobotanical (Moerman). Apache, White Mountain: Infusion of stems-leaves for gonorrhea or first stages of syphilis.Cahuilla: Decoction of twigs to 'clear the system,.' Coahuilla: unspecified. Navajo: Stem-leaf infusion for kidney and venereal troubles. Paiute: Twigs for lessening disagreeable flavors; compound decoction as a salve for burns; stem decoction for venereal disease. Shoshoni: powdered branches and twigs applied to sores; decoction to stimulate urination, and for venereal diseases. Zuni: Infusion of stems-leaves for venereal diseases, especially syphilis.    

.    Ephedra viridis Coville 1893. Green ephedra. Stems green to bright yellow-green, numerous and erect, all mostly parallel to one another, spreading at an angle of ~30°; leaves 2 per node, pale orange in lower half, or with pale orange median strip,  hyaline above, free to  near base or united  to 4/5 their length, clasping to adpressed, not spreading, deciduous except for basal thickened ring which turns reddish brown to blackish resinous. Male cones spreading, with pale yellow or yellowish green to flesh-colored bracts; female cones erect, sessile or on short to long stalks to 5 cm or more; seeds two.
Widely distributed in California chaparral and desert, 1,000–7,000 ft, to Utah, New Mexico, Colorado, Oregon. Green ephedra ("Mormon tea") scrub recognized in MCV2 when  ≥1% absolute cover and >30% relative cover with other species as seen on hillsides above Sand Canyon Road. Type from
Inyo Co., Coso Mtns. above Crystal Spring, CA. Colville & Funston 923,  (June 12, 1891), annotated by Ickert-Bond May 2002, holotype (US!)  [stems dark green, leaves on young shoots long connate, shallowly notched near apex, lobes rounded obtuse, occasional pollen cone at nodes]. Coville reported he could not find plants with seed. Ft. Tejon & vicinity, L.J. Xantus de Vasey (1857-58), annotation Ickert-Bond, May 2007; syntype for E. nevadensis det. by Ickert-Bond, Nov 2007 (US!). Kern Co.: Occurs most often in pinyon-juniper forest regions of the Sierra Nevada, 360–2,028 m (CCH); also common in gray pine-interior oak woodland along the Kern River near Lake Isabella. Noted by Twisselmann to be especially common in the pinyon woodland around Frazier Park.  CCH records also from the Temblor Range and Inner Coast Ranges.

Ethnobotanical (Moerman). Havasupai: Used as an emetic for bowel complaints. Hopi: Tonic from stems with flowers for syphilis. Kawaiisu: Stem infusion for anemia and backaches. Navajo: Decoction new growth used as cough medicine; infusion of stems for syphilis. Paiute: infusion of stems for diarrhea (for children), or for rheumatism, colds, or to purify blood, stomach ulcers, colds, regulating kidneys, or applied as  powder to sores (also Northern Paiute), to treat stomach ulcers. Shoshone: Compound infusion to children for diarrhea; twigs as blood purifier; powdered stems for burns; decoction of root or salted decoction of stems as a physic, cold remedy Tewa: dried flowers and stems as tonic for syphilis. Tubatulabal: Decoction of stems for syphilis. Washo: Decoction of twigs or branch for delayed or difficult menstruation.  

            Ephedra viridis is generally recognized by the erect closely parallel stems. Variation in the shape and the degree to which scale leaves are united suggest distinct varieties or species.  Forms with long stalked cones as shown in the adjacent image from Frazier Park (May 2012), which might be interpreted by some as belonging to E. coryi, even though cones of E. viridis are described as not stalked to shortly stalked, also occur elsewhere such as in New Mexico; they are considered a variable feature of E. viridis (Sivinski 2010). However, other variation such as seen in the Piute Mountains include two tentatively undetermined species.  One has erect branches, young brownish scale-leaves developing at shore intervals, and short whorls of branches that appear deciduous (Ephedra sp. B), and  a second has divaricate branches that are 4-angled (Ephedra sp. A).  Images of both are presented above.   Plants on rocky benches above the Kern River just south of Lake Isabella have longer transparent scale-leaves and also longer photosynthetic leaves.
Links: SEINet, Calflora, Calscape









Ephedra viridis, seed and pollen cones










USDA ARS historical collection data for cancer research

Ephedra cutleri
Ephedra viridis:


Bolinder K, AM Humphreys, J Ehrlén, R Alexandersson, SM Ickert-Bond, C Rydin. 2016. From near extinction to diversification by means of a shift in pollination mechanism in the gymnosperm relict Ephedra (Ephedraceae, Gnetales). Bot J Linn Soc 180 (4): 461-477.

Calflora. My Observations (Richard Spjut) accessed 11/23/2023.

Caveney, S., D. A. Charlet, H. Freitag, M. Maier-Stolte and A. N. Starratt. 2001. New observations on the secondary chemistry of world Ephedra (Ephedraceae). Am. J. Bot. 88:1199-1208.

CCH2 (Consortium of California Herbaria) accessed 11/13/2023, Ephedra foeminea, det. SM Ickert-Bond 2007, originally identified E. distachya). Coll. R. N. Philbrick, Santa Barbara County, Santa Ynez Mtns: Trout Club, upper San Jose Crk below San Marcos Pass 1996-5-15 (2 specimens, 1 annotated).

Cutler, HC. 1939.  Monograph of the North American species of the genus Ephedra. Ann. Missouri Bot. Gard. 26: 373–427.

Dhote P [Poonam]. Undated. Ephedra: Meaning, Reproduction and Economic Importance | Gnetales. Accessed 12/13/23 (Note: website loaded with ads; however, nice illustrations of Ephedra from various sources, E. foliata cited frequently, indicated to be monoecious..  https://www.biologydiscussion.com/gymnosperm/ephedra-meaning-reproduction-and-economic-importance-gnetales/22572.

Earle CJ. 2023. The Gymnosperm database, Ephedra tweedieana (last modified 03-03-2023). https://www.conifers.org/ep/Ephedra_tweedieana.php, last accessed 12/13/2023. Nomenclatural comments on publication dates for C. A. Meyer's species.

eflora of China. http://www.efloras.org/florataxon.aspx?flora_id=2&taxon_id=111784, accessed 12/13/23. 14 species. Also cited below under Liguo.

eflora of India, photo of plant with fruit of E. foliata. https://efloraofindia.com/2013/12/26/ephedra-foliata/ accessed 11/23/2023.

Elhadef K, S Smaoui, M Fourati, HB Hlima, AC Mtibaa, I Sellem, K Ennouri, Li Mellouli. 2020. A review on worldwide Ephedra history and story: From fossils to natural products mass spectroscopy characterization and biopharmacotherapy potential. Evidence-Based Complementary and Alternative Medicine. Hindawi. Article ID 1540638 | https://doi.org/10.1155/2020/1540638.

Faried A,  A EL-Banhawy, M Elqahtan. 2018. Taxonomic, DNA barcoding and phylogenetic reassessment of the Egyptian Ephedra L. (Ephedraceae). Catrina 17(1): 1-17.

Flora of Qatar, website--E. foliata:  (https://www.floraofqatar.com/ephedra_foliata.htm, 11/21/2023).

Flora of the U.S.S.R. Vol. 1. Archegoniatae and Embryophyta. 1934.  V.L. Komarov, Editor. Israel Program for Scientific Translation, 1968.  9 species.

FNA. See Stevenson.

Foster AS (1972). Venation patterns in the leaves of Ephedra. J. Arn. Arb. 53: 364-78, plates I-VII

Frietag H,  M Maria-Stolte. 1989. The Ephedra species of P. Forsskal. Taxon 38(4):545-556. Designated types for E. aphylla (neotype) and E. foeminea (lectotype) and compared E. foliata with E. foeminea, E. aphylla, and E. fragilis.

Florin CR. 1933. Über einige neue oder wenig bekannte asiatische Ephedra Arten der Sektion Pseudobaccatae Stapf. Kongl. Svenska Vetenskapsakad. Handle, 12, 1–49. [Google Scholar] Among them, E. fedtschenkoae seems unique in being monoecious, but Florin (1933) reported that monoecious individuals are common in Ephedra. The other three species are tetraploids based on previous studies (e.g. Leitch et al. 2001) and our preliminary investigation. Therefore, the exclusion of these species should not greatly affect our inference about the origins of other tetraploid species (mostly allotetraploids).

González-Juárez DE, A Escobedo-Moratilla, J Flores et al. 2020. A review of the Ephedra genus: Distribution, ecology, ethnobotany, phytochemistry and pharmacological properties. Molecules 2020, 25, 3283; doi:10.3390/molecules25143283.

Hollander JL, SB Vander Wall, JG Baguley 2010 Evolution of seed dispersal in North American Ephedra. Evol Ecol 24:333–345.

Huang J, DE Giannasi, RA Price. 2005. Phylogenetic relationships in Ephedra (Ephedraceae) inferred from chloroplast and nuclear DNA sequences. Mol Phylogenet Evol 35:48–59.

Ickert-Bond SM, S Renner 2016. The Gnetales: Recent insights on their morphology, reproductive biology, chromosome numbers, biogeography, and divergence times. J Systemat Evol 54(1): 1-16. https://doi.org/10.1111/jse.12190 54 species listed.

Ickert-Bond SM. 2012. Jepson Flora Project (eds.) 2023, Jepson eFlora, Ephedra. https://ucjeps.berkeley.edu/eflora/, accessed on November 22, 2023. Additional note: "No expert verified images found for Ephedra foeminea".

Icket-Bond SM, C Rydin. 2011. Micromorphology of the seed envelope of Ephedra L. (Gnetales) and its relevance for the timing of evolutionary events. International Journal of Plant Sciences 172: 36–48.

Ickert-Bond SM, C Rydin, SS Renner. 2009. A fossil-calibrated relaxed clock for Ephedra indicates an Oligocene age for the divergence of Asian and New World clades and Miocene dispersal into South America. J Syst Evol 47:444–456.

Ickert-Bond SM, MF Wojciechowski. 2004. Phylogenetic relationships in Ephedra (Gnetales): evidence from nuclear and chloroplast DNA sequence data. Syst Bot 29:834–849.

Ickert-Bond SM, JJ Skvarla, WF Chissoe. 2003. Pollen dimorphism in Ephedra L. (Ephedraceae). Rev Palaeobot Palynol 124:325–334.

JM2. The Jepson Manual, 2nd ed. 2012. Ephedra by SM Ickert Bond, also cited above.

JungleDragon. https://www.jungledragon.com/  Accessed 11/24/23. “JungleDragon is a nature and wildlife community for photographers, travellers and anyone who loves nature. We're genuine, free, ad-free and beautiful.”  Images of Ephedra foliata in fruit credited to Ori Fragman-Sapir.

Kew, Royal Botanical Gardens. 2023. International Plant Names Index and World Checklist of Vascular Plants. http://creativecommons.org/licenses/by/3.0. Ephedra ciliata.

Liguo Fu, F Li-kuo, Y Yongfu, H Riedl. 1999. Ephedraceae. Flora of China 4: 97–101. 14 species

Loera I, V Sosa, SM Ickert-Bond. 2012. Diversification in North American arid lands: Niche conservatism, divergence and expansion of habitat explain speciation in the genus Ephedra. Mol. Phylogenet. Evolut. 65, 437–450. doi: 10.1016/j.ympev.2012.06.025

Mabberley DJ. 1987. The Plant-Book. 2nd Ed. Cambridge Univ. Press

Maynard Moe L. 2016. Kern County Flora. CNPS, Sacramento CA.

Mehra PN 1950. Occurrence of hermaphrodite flowers and the development of the female gametophyte in Ephedra intermedia Schrenk.and Mey. Ann. Bot. 14: 165-180.

Meyer CA von 1846. Versuch einer Monographie der Gattung Ephedra. Mém. Acad. Imp. Sci. Saint-Pétersbourg, Sér. 6, Sci. Math., Seconde Pt. Sci. Nat. 5(2): 225--298.

MCV2. Sawyer JO, T Keeler-Wolf, JM Evens. 2008. A manual of California vegetation, 2nd  ed. [MCV2]. CNPS Publication Committee, Sacramento.

Moerman DE. Native American medicinal plants.  An ethnobotanical dictionary. 1998. The medicinal uses of more than 3000 plants by 218 Native American tribes. Timber Press, Portland.

Norbäck-Ivarsson L. 2014. Pollen morphology in Ephedra (Gnetales) and implications for understanding fossil ephedroid pollen from the Tibetan Plateau, using a phylogenetic approach. Master thesis, Stockholm University, Stockholm, Sweden. Also published by K Bolinder with L Norbäck- Ivarsson, AM Humphreys, SM Ickert-Bond, F Han, C Hoorn, C Rydin. Grana 55: 24-51 (2016)

PALYNODATA - one of the classic 20th century palynological databases. This bibliographic database, based on Gerhard O. W. Kremp's initial research, and compiled since 1974 by Palynodata Inc., under the direction of Ken Piel, indexes 122,422 species from 22,152 documents. The last entry was made in 2006 and copyright was transferred to Canada in 2007. Ephedra data included 96  species names, several of which appear to be variations in the spelling of a species name among the 412 literature citations. Palynodata Inc., White, J M; 10.4095/225704. Last accessed 12/22/2023.

Pant DD, BK Verma. 1974. Taxonomy of the genus Ephedra. Significance of stem and leaf epidermis and cuticle. Bot. J. Linn. Soc. 69: 287-308.

Peebles, RH, LC Wheeler. 1940. Arizona plants: a new variety and new names and combinations. Journal of the Washington Academy of Sciences 30:473.

Powell AM. 1998. Trees and shrubs of the Trans-Pecos and adjacent areas. University of Texas Press, Austin.

Price RA. 1996. Systematics of the Gnetales: a review of morphological and molecular evidence. International Journal of Plant Sciences (Supplement) 157: S40–S49.

Rydin C, R Blokzij, O Thureborn, N Wikström. 2021. Node ages, relationships, and phylogenomic incongruence in an ancient gymnosperm lineage – Phylogeny of Ephedra revisited. Taxon 70, 701–719. doi: 10.1002/tax.12493

Rydin C, P Korall. 2009. Evolutionary relationships in Ephedra (Gnetales), with implications for seed plant phylogeny. Int J Plant Sci 170:1031–1043.

Rydin C, KR Pedersen, EM Friis. 2004. On the evolutionary history of Ephedra: Cretaceous fossils and extant molecules. PNAS 101 (47) 16571-16576, https://doi.org/10.1073/pnas.0407588101.

San Martin JAB, RE Pozner, VS Di Stilio. 2022. Heterochrony and repurposing in the evolution of gymnosperm seed dispersal units. Evodevo. 2022 Feb 16;13(1):7. doi: 10.1186/s13227-022-00191-8. PMID: 35172885; PMCID: PMC8851845.

Scott RA. 1960. Pollen of Ephedra from the Chinle formation (Upper Triassic) and the genus Equisetosporites: Micropaleontology, v. 6, p. 271-276, 1 pI.

SEINet accessed 2023-11-13. Ephedra foliata: 17 specimens, Rodin 8112, collected from cultivation in India.

Spjut RW. 2024 (in press). Ephedra novelties in the Piute Mountains of Kern County: E. foliata, a nonnative Mediterranean species found along Erskine Creek, and E. cf. viridis in Squirrel Canyon with a bisexual cone. Mimulus Memo at CNPS Kern Chapter Website Mimulus Memo full issues at Biodiversity Heritage Library (BHL).

Spjut RW. 2015. Status on trees and shrubs of Kern County. CNPS Kern Chapter, Mimulus Memo September, p 3-4 (President's Message). Full issue at Biodiversity Heritage Library (BHL).

Spjut RW 2014. Some botanical oddities in the Kern County Flora. September, p 3-4 (President's Message). Full issue at Biodiversity Heritage Library (BHL).

Spjut RW. 2014.  A revised key to Gymnosperm fruit types (pdf, 22 April 2014, not formally published, many new terms proposed).

Spjut RW 2007. A phytogeographical analysis of Taxus (Taxaceae) based on leaf anatomical characters. J. Bot. Res. Inst. Texas 1(1): 291–332.

Spjut, R.W. 1994: A systematic treatment of fruit types. Mem. New York Bot. Gard. 70:1-182.

Stevenson DW. 1993. Flora North America (north of Mexico). Vol. 2: 428-434, 12 species. eflora http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=10313, accessed 12/13/2023

Thoday (Sykes) MG, EM Berridge. 1912. The anatomy and morphology of the inflorescences and flowers of Ephedra. Annals of Botany XX VL No. CIV.

Tropicos. Ephedra. Tropicos.org. Missouri Botanical Garden. 26 Jan 2024 <https://tropicos.org> © 2024 Missouri Botanical Garden - 4344 Shaw Boulevard - Saint Louis, Missouri 63110.

Van Gelderen DM,  JRP van Hoey Smith. 1996. Conifers, 2 Vols. Timber Press Vol. 1. Image of E. foliata.

Villanueva-Almanza L, RM Fonseca. 2011. Revisión taxonómica y distribución geográfica de Ephedra (Ephedraceae) en México. Acta Botanica Mexicana 96: 79-116. 9 species, one endemic to Mexico.

Wikimedia Commons. Ephedra. Ephedra californica images, "At the University of California Davis Arboretum, in Davis, Northern California."  Last accessed Jan 22. 2024.

Wikipedia. Ephedra. Accessed 11/23/2023. Map showing geographical distribution of the genus Ephedra.

Wu H, Ma Z, Wang MM, Qin AL, Ran JH, Wang XQ. A high frequency of allopolyploid speciation in the gymnospermous genus Ephedra and its possible association with some biological and ecological features. Mol Ecol. 2016 Mar;25(5):1192-210. doi: 10.1111/mec.13538. Epub 2016 Feb 16. PMID: 26800145; PMCID: PMC7168403.

Yang Y, Wang Q (2013) The earliest fleshy cone of Ephedra from the Early Cretaceous Yixian Formation of Northeast China. PLoS ONE 8(1): e53652. doi:10.1371/journal.pone.0053652

Yu Q, Yang F-S, Chen Y-X, Wu H, Ickert-Bond SM, Wang XQ. 2023. Diploid species phylogeny and evolutionary reticulation of Ephedra in the Tethys Coast. JIPB Oct 13.  https://doi.org/10.1111/jipb.13573

Note: Additional references cited under main page for Trees and Shrubs of Kern County.