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Two new desert Eschscholzia (Papaveraceae) from southwestern North America

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Two new species of Eschscholzia are described. Both are found in the deserts of California and one extends outside the state boundary into Arizona. Eschscholzia androuxii Still, sp. nov. is found mainly in and around Joshua Tree National Park in Riverside and San Bernardino counties. Eschscholzia papastillii Still, sp. nov. is found from the northern Mojave south through Joshua Tree National Park to central Imperial County. Both are annuals found in coarse, sandy soil and have yellow flowers typical of desert Eschscholzia. Eschscholzia papastillii has an expanded receptacular rim similar to that of Eschscholzia californica. Eschscholzia androuxii has anthocyanin bands around the stamen filaments.
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Two new desert Eschscholzia (Papaveraceae) from southwestern North America 45
Two new desert Eschscholzia (Papaveraceae)
from southwestern North America
Shannon M. Still 1
1 Department of Plant Science and Conservation, Chicago Botanic Garden, 1000 Lake Cook Road, Glencoe,
IL 60022
Corresponding author: Shannon M. Still ( still.shannon@gmail.com )
Academic editor : D. Soltis |Received 4 December 2013 |Accepted 24 February 2014 |Published 11 March2014
Citation: Still SM (2014) Two new desert Eschscholzia (Papaveraceae) from southwestern North America. PhytoKeys
35 : 45 – 56 . doi: 10.3897/phytokeys.35.6751
Abstract
Two new species of Eschscholzia are described. Both are found in the deserts of California and one extends
outside the state boundary into Arizona. Eschscholzia androuxii Still, sp.nov. is found mainly in and
around Joshua Tree National Park in Riverside and San Bernardino counties. Eschscholzia papastillii Still,
sp. nov. is found from the northern Mojave south through Joshua Tree National Park to central Imperial
County. Both are annuals found in coarse, sandy soil and have yellow  owers typical of desert Eschschol-
zia. Eschscholzia papastillii has an expanded receptacular rim similar to that of E. californica. Eschscholzia
androuxii has anthocyanin bands around the stamen  laments.
Keywords
Arizona, California, Eschscholzia, Papaveraceae, new species, Eschscholzia androuxii, Joshua Tree poppy,
Eschscholzia papastillii, cryptic desert poppy, Mojave Desert, Colorado Desert, Sonoran Desert
Introduction
Eschscholzia Cham. (1820) is a genus in the Papaveraceae tribe Eschscholtzieae along
with the genera Hunnemannia Sweet and Dendromecon Benth.  e type genus is na-
tive to the mainland and islands of western North America in both the United States
and Mexico, but the type species, Eschscholzia californica Cham. has invaded Mediter-
ranean regions around the world.  e taxa are native to mesic and xeric landscapes.
Recent treatments for Eschscholzia (Clark 1997, Roberts 1989) recognize 12 species
PhytoKeys 35: 45–56 (2014)
doi: 10.3897/phytokeys.35.6751
www.phytokeys.com
Copyright Shannon M. Still . This is an open access article distributed under the terms of the Creative Commons Attribution International License
(CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
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Shannon M. Still / PhytoKeys 35: 45–56 (2014)
46
and several subspecies for a total of 16 taxa. Recent phylogenetic work (Still and Pot-
ter 2013) indicates 18 distinct taxa. While the genus is fairly small, there are nearly
198 taxon names and 168 type specimens, with the majority described by EL Greene
(1905).  e majority of described taxa are synonymous with E. californica.
All taxa are herbaceous annuals or perennials with taproots and basal rosettes.  e
leaves are ternately-dissected 2–many times and range from bright green, dark green
to glaucous grey-green. Flowers are bisexual, have two sepals fused into a single cap
structure, four petals and many stamens.  e sepals fall from the  ower upon opening.
e owers are yellow, orange or can be yellow with an orange basipetal spot on each
petal.  e desert taxa of the genus can be di cult to identify (personal experience) and
this resulted in further morphological and molecular examination of Eschscholzia (Still
2011, Still and Potter 2013). In the course of study, two new taxa were discovered
among Eschscholzia native to desert regions (Still 2011).  ese two taxa exhibited both
morphological traits and mutations in nucleotide sequences from nuclear and plastid
regions that distinguish them from one another and from all previously described spe-
cies of Eschscholzia.  e two new taxa are here described as new species.
Taxonomy
Eschscholzia androuxii Still, sp. nov.
urn:lsid:ipni.org:names:77136479-1
http://species-id.net/wiki/Eschscholzia_androuxii
Figs 1–3
Type. UNITED STATES, California: Riverside County, just south of entrance sign
to Joshua Tree National Park heading north on Cottonwood Springs Road from US
Interstate-10, [33°41.188'N, 115°48.103'W], 610 m alt., 14 Feb 2008, Shannon M.
Still 258A with Jennifer Still and Charles Still (holotype: DAV!).
Diagnosis. Eschscholzia androuxii is similar to E. minuti ora subsp. twisselmannii
C. Clark & M. Faull but with ultimate lobes of the dissected leaves more numerous and
narrower than E. minuti ora subsp. twisselmannii. Eschscholzia androuxii is similar to E.
minuti ora subsp. minuti ora S. Watson and E. minuti ora subsp. covillei (E. Greene)
C. Clark but with larger  owers and consistently appearing, pronounced black-blue or
darkened anthocyanin area or spot basipetally located on the fused  lament bases of the
stamens. Eschscholzia androuxii di ers from E. papastillii and E. parishii with the afore-
mentioned stamen spot and basal foliage that appears more compact in habit.
Description. Annual herb, erect or spreading with a basal rosette of leaves from a
taproot. Leaves highly ternately-dissected into a great number of ultimate lobes, which
may number to 100 on larger specimens. Leaves glaucous-green with ultimate lobes
more rounded than pointed. Basal leaves are 3–11 cm long and 0.8–3.2 cm wide and
held on a petiole comprising 2/3 the entire leaf length. Younger plants will have fewer
ultimate lobes and shorter, narrower leaves. In orescence with few  owers held above
Two new desert Eschscholzia (Papaveraceae) from southwestern North America 47
Figure 1. Illustrations of leaves, buds and  owers of Eschscholzia androuxii. A Basal leaf B Cauline leaf
CBud D Petal E Stamens showing diagnostic anthocyanin spot at base of the fused  laments.
the foliage and to 4 dm above the ground. Leaves on the in orescence are 2–20 mm
long and are divided into 2–23 ultimate lobes. Buds nodding and 4.5–11.5 mm long
with an apiculate bud tip less than 25% of the total bud length. Less mature buds may
Shannon M. Still / PhytoKeys 35: 45–56 (2014)
48
be shorter than average with a longer bud tip by percentage. Flowers held upright and
are yellow with four petals 10.5–23 mm long. Each  ower has 20–36 stamens fused at
the base.  ere is a darkened area or patch, often black-blue, located at the fused  la-
ment bases of the stamens. Receptacles obconic and 2.5–5.5 mm long and 1.1–3 mm
wide and often have a scarious inner hyaline rim. Fruit 3.5–6.5 cm long with 10–12
nerves, dehiscing at maturity. Seeds with reticulate ridges.
Distribution (Fig. 3). Found in and around Joshua Tree National Park in both
Riverside and San Bernardino counties of California.
Habitat and ecology. Desert washes,  ats, and slopes in coarse, sandy soil.
Phenology. Eschscholzia androuxii typically  owers between late-February and
early-May but may  ower earlier in the season, including in the fall, during years with
a summer rain and cool fall temperatures.
Etymology. e species is named for James André and Tasha La Doux, two de-
sert botanists and friends that helped point to the problems with desert Eschscholzia
identi cation.
Suggested common name. Joshua Tree poppy.
Conservation status. As this is a new taxon it has yet to be considered for conser-
vation status. Due to the limited range and low number of occurrences, the author sug-
gests the California Native Plant Society consider this taxon for listing as a rare plant.
Specimens examined. U.S.A. California: Riverside Co.: White Water, Apr 1907,
S.B. Parish 6103 (DS!); slope of hill at west side of mouth of Whitewater Canyon, 18
Mar 1962, D.W. Kyhos 62-43 (DS!); Cottonwood Pass, Joshua Tree National Monu-
ment, 19 Mar 1949, C. Francis Shutts 58 (ASU!, DES!); Coachella Valley, Desert
Hot Springs, N of intersection of Pierson Blvd. and Atlantic, north of  ood control
ditch, 6 Apr 2001, A.C. Sanders, Mitch Provance & T.B. Salvato 23939 (DES!); Joshua
Tree National Park, [33°41.18333'N, 115°48.1'W], 15 Feb 2008, Shannon M. Still,
Jennifer R. Still, & Charles M. Still 258B (DAV!); id., Cottonwood Wash on west
side of road, [33°41.21299'N, 115°48.15100'W], 22 Feb 2009, Shannon M. Still 444
(DAV!); id., [33°41.833'N, 115°48.17598'W], 3 Mar 2009, Shannon M. Still & Rob-
ert Lee 457 (DAV!); id., [33°50.22799'N, 115°45.174'W], 28 Mar 2009, Shannon M.
Still, Steven M. Still & Carolyn M. Still 512 (DAV!); San Bernardino Co.: 1.4 mi N
of Yucca Valley on road to Lucerne Valley. About 19 mi west of town of Twentynine
Palms, 6 Apr 1957, John H.  omas 6627 (DS!).
Discussion. is new taxon has a darkened area basipetally located on the stamen
laments (Fig. 1e), which are fused at the base. Eschscholzia minuti ora subsp. twis-
selmannii also has regularly occurring stamen spots, but only on approximately 70% of
specimens examined. No other closely related taxa have these stamen spots.  e ower
size for this new species is similar to that of the diploid E. minuti ora subsp. twis-
selmannii but larger than both the hexaploid E. minuti ora subsp. minuti ora and the
tetraploid E. minuti ora subsp. covillei.  e petal size for E. minuti ora subsp. covillei,
described in Flora of North America as 6–18 mm long, does overlap with the petal size
of E. androuxii, with petals 10–23 mm long. But more recent morphological study of
the genus (Still 2011, Still in preparation) indicates that the petals in E. minuti ora
Two new desert Eschscholzia (Papaveraceae) from southwestern North America 49
subsp. covillei range from 4.5–12.5 mm.  e reason for this discrepancy may be that
some of the larger- owered E. minuti ora subsp. covillei specimens are actually the
new taxon, E. androuxii.  e Joshua Tree poppy has an overlapping range with several
species but is found only in Riverside Co. and the southern part of San Bernardino Co.
Figure 2. Photographs of Eschscholzia androuxii. A Species pro le shot B Species pro le shot CEschscholzia
androuxii in the type area in a heavy- owering year.
Shannon M. Still / PhytoKeys 35: 45–56 (2014)
50
and not much further north or south of Joshua Tree National Park.  e range does not
overlap, and there are more basal leaf ultimate lobes, than with E. minuti ora subsp.
twisselmannii.  e tips of the basal leaf ultimate lobes are more rounded (Fig. 1a) than
what is found in either E. parishii or E. papastillii (Fig. 4a), and E. androuxii has three
times the number of cauline leaf ultimate lobes (Fig. 1b) as these two taxa.
Eschscholzia papastillii Still, sp. nov.
urn:lsid:ipni.org:names:77136480-1
http://species-id.net/wiki/Eschscholzia_papastillii
Figs 3–5
Type. UNITED STATES, California: Riverside County, Joshua Tree National Park
next to stone outcropping o Old Dale Road. [33°50.232'N, 115°45.12'W], 724 m
alt., 19 Apr 2009, Shannon M. Still 546A (holotype: DAV!).
Figure 3. Distribution map showing the range for the two new Eschscholzia. Eschscholzia androuxii is repre-
sented by right-slanting cross-hatching. Eschscholzia papastillii is represented by left-slanting cross-hatching.
Two new desert Eschscholzia (Papaveraceae) from southwestern North America 51
Diagnosis. Eschscholzia papastillii is similar to E. parishii Greene but with more
basal leaf ultimate lobes and more broadly spreading leaves. Eschscholzia papastillii has
an enlarged receptacle (Fig. 4d) that is widely-obconic or bell-shaped, and wider at
the midpoint of the receptacle than E. parishii, E. androuxii or any of the subspecies
of E. minuti ora, which are usually more obconic or funnel-shaped.  e expanded
receptacular rim of E. papastillii is similar, but typically smaller, than the expanded
receptacular rim of E. californica. Eschscholzia papastillii di ers from E. androuxii and
E. minuti ora with basal foliage that appears less compact in habit.
Description. Annual herb, erect or spreading with a basal rosette of leaves from
a taproot. Leaves highly ternately-dissected into 17–70 ultimate lobes with the higher
number on larger specimens. Leaves glaucous-green to green with ultimate lobes more
pointed than rounded. Basal leaves are 2.7–16 cm long and 0.9–7 cm wide and held
on a petiole comprising 2/3 the entire leaf length. Younger plants have few basal leaf
ultimate lobes and shorter, narrower leaves. In orescence with few  owers held above
the foliage and to 5 dm above the ground. Leaves on the in orescence are 3–50 mm
long and are divided into 1–13 ultimate lobes. Buds nodding to erect and 2.5–16 mm
long with an apiculate bud tip greater than 30% of the total bud length. Less mature
buds may be shorter than average with a longer bud tip by percentage. Flowers held
upright and are yellow with four petals 5–24 mm long. Each  ower has 12–32 stamens
fused at the base. Receptacles widely-obconic or funnel-shaped to nearly bell-shaped,
3–9 mm long and 1.5–4.7 mm wide. Receptacular rim typically noticeable and often
thick but can be scarious, expanded laterally up to 1.2 mm from the top of the recep-
tacle.  e receptacle often has a scarious inner hyaline rim in addition to the outer rim
diagnostic of the species. Fruit 4 –8 cm long with 10–12 nerves, dehiscing at maturity.
Seeds with reticulate ridges.
Distribution (Fig. 3). Found north to the northern Mojave Desert; south into
northern Colorado Desert of San Diego Co., and possibly south along the east side of
the Sea of Cortez in Mexico; east to the California-Arizona border (Whipple Moun-
tains); west to the western end of Joshua Tree National Park.
Habitat and ecology. Desert washes,  ats, and gentle slopes in coarse, sandy soil.
Phenology. Eschscholzia papastillii typically  owers between late-February and
early-May but may  ower earlier in the season, and in the fall, during years with a
summer rain and cool fall temperatures.
Etymology. e species is named in honor of Dr. Steven Still, my father and men-
tor and the reason for which I study plants.
Suggested common name. Cryptic desert poppy.
Conservation status. As this is a new taxon it has yet to be considered for conser-
vation status. Due to the range and number of occurrences the author does not sug-
gests this taxon be considered for conservation status.
Specimens examined. U.S.A. California: Kern Co.: Hidden Springs Rd., 6
May 1930, Lester Rowntree s.n. (CAS!); Riverside Co.: Painted Canyon, 4 Mar 1922,
Edmund C. Jaeger s.n. (DS!); Painted Canyon, 12 Apr 1927, Frank W. Peirson 7167
(CAS!); near Shavers Well, 6 Apr 1930, R.A. Piebles and H.F. Loomis 188 (DS!);
Shannon M. Still / PhytoKeys 35: 45–56 (2014)
52
Figure 4. Illustrations of leaves, buds and  owers of Eschscholzia papastillii. A Basal leaf B Cauline leaf C Bud
showing widened receptacle D Enlarged receptacle with expanded receptacular rim common in the species
EPetal F Stamens lacking the anthocyanin spot at base of the fused  laments common to E. androuxii.
Box Canyon, Coachella Valley, 21 Mar 1937, Ynez Whilton Winblad s.n. (CAS!);
Coachella Valley, 21 Mar 1937, Ynez Whilton Winblad s.n. (CAS!); east slope of
Chocolate Mnts, 22 Mar 1937, Ynez Whilton Winblad s.n. (CAS!); above Cotton-
Two new desert Eschscholzia (Papaveraceae) from southwestern North America 53
wood Springs, west end of Eagle Mnts, 13 Apr 1949, Philip A. Munz 13056 (CAS!);
Road to Morongo Valley, 7.8 mi from junction with Highway 99/60/70, 14 Apr
1952, Richard Snow (DS!); Box Canyon, 5 Apr 1953, Richard Snow 51a (DS!); 0.8
Figure 5. Photographs of Eschscholzia papastillii. A Species pro le shot B Buds with enlarged receptacle
common in the species C Flower with enlarged receptacle common in the species D Species pro le shot.
Shannon M. Still / PhytoKeys 35: 45–56 (2014)
54
mi east of Cactus City, 13 mi east of Coachella, near U.S. Highways 60 and 70.
Colorado Desert, 30 Mar 1957, John H.  omas 6523A (DS!); U.S. Highway 60/70,
Indio to Blythe, 1 mi. W of Cactus City, 29 Apr 1958, P.C. Everett and E.K. Balls
23013 (CAS!, DAV!); East of Indio on Highway 60/70, about 6 mi west of Cac-
tus City, 15 Mar 1960, W.R. Ernst 720 (CAS!); Mecca-Joshua Tree Road, 7 miles
southwest of junction with Interstate Highway 10, 13 Apr 1976, Curtis Clark 527
(DAV!); 2 mi S.E. of Desert Center, 1 Jul 1981, J.C. Roos s.n. (ASU!, CAS!); wash
along Eagle Mountain Rd., north along I-10, 3 Mar 1995, John Wear s.n. (DAV!);
Joshua Tree National Park, [33°43.68999'N, 115°49.317'W], 14 Feb 2008, Shan-
non M. Still 253 with Jennifer R. Still, Charles M. Still (DAV!); id., [33°55.012'N,
115°52.60305'W], 3 Mar 2009, Shannon M. Still 452 with Robert Lee (DAV!); id.,
[33°50.22799'N, 115°45.174'W], 28 Mar 2009, Shannon M. Still 513 with Steven
M. Still, Carolyn M. Still (DAV!); San Bernardino Co.: Sheephole Mnts., 8 Apr
1935, P.A. Munz 13823 (DS!); Cave Spring, Lower Sonoran zone, 16 Apr 1940,
C.L. Hitchcock 6073 (DS!); Bristol Lake Basin 8.8 mi N. of summit of Sheephole
Pass on Amboy Rd., 16 Mar 2001, A.C. Sanders, Mitch Provance & Petra Wester
23753 (CAS!); Sheephole Pass just to south of the top of the pass, [34°13.711'N,
115°43.19599'W], 25 Nov 2007, Shannon M. Still 222A (DAV!); Base of Old Dad
Mountains in wash, [34°44.512'N, 115°45.081'W], 26 Nov 2007, Shannon M.
Still, Jim André & Tasha La Doux 248 (DAV!); Base of Old Dad Mountains in
wash, [34°44.512'N, 115°45.082'W], 6 Apr 2008, Shannon M. Still & Steven M.
Still 377A (DAV!); Clipper Mountains, just o the pipeline road, [34°40.573'N,
115°22.73502'W], 18 Apr 2009, Shannon M. Still, Jim André, Je Galvin & Amy
Toulsen 536 (DAV!).
Discussion. While the buds of all desert Eschscholzia appear similar, those of E.
papastillii most resemble E. parishii, as the bud tip is typically more than 25% of the
total bud length.  e receptacular rim is prominent in this species and E. californica is
the only other species that has a pronounced receptacular rim.  e range of E. papas-
tillii extends from San Bernardino County south to northern Imperial County. Most
collections of E. parishii collected north of San Diego and Imperial Counties are likely
the new E. papastillii. Eschscholzia papastillii extends at least into easternmost San Ber-
nardino County that contains the Whipple Mountains, and likely well into Arizona.
Key to the desert Eschscholzia species
1 Basal leaf ultimate lobes long-linear; leaves ternately-dissected 2–3×;  ower
scapes typically without cauline leaves; seed coats pitted without reticula-
tions ................................................................................... E. glyptosperma
1’ Basal leaf ultimate lobes not long-linear; leaves ternately-dissected 3–7×;
owers typically borne on few- owered racemes with a cauline leaf at each
ower axil; seeds coats reticulate
2 Receptacular rim prominent when in fruit, 0.25 –5 mm
Two new desert Eschscholzia (Papaveraceae) from southwestern North America 55
3 Basal leaf ultimate lobes with length < 3× width, with acute or rounded tips;
leaf blades often deep green with a glaucous patch at the crotch of the leaf
dissections; cauline leaf ultimate lobes many (range 5–30) with rounded to
acute tips; Petals yellow, often with a basipetal orange spot, or petals orange
4 Petals yellow, often with a basipetal orange spot, or petals orange, or rarely white
(Arizona mountains); cotyledons entire; annual; limited to eastern Mojave De-
sert in California and through Arizona ............ E. californica subsp. mexicana
4’ Petals yellow, often with a basipetal orange spot, or petals orange; cotyledons
bi d (2-lobed); annual or perennial; widespread but mostly along highways,
railways, and planted areas ........................ E. californica subsp. californica
3’ Basal leaf ultimate lobes with length 3.5 (2–8)× width, with acute tips; leaf
blades bright-green to yellow-green; cauline leaf ultimate lobes 3 (rarely 5–13)
with acute tips; petals yellow without basipetal orange spot .......E. papastillii
2’ Receptacular rim not prominent in fruit, < 0.25 mm
4 Petal < 1 cm long
5 Buds with tip < 25% total bud length; cauline leaves generally with > 5 (rarely
< 6) ultimate lobes, ± rounded to acute; 2n=24 or 36
6 Basal leaf ultimate lobes ± narrow, length ca. 4.5× the width; petals generally
less than 5.5 (rarely 2–9) mm long, stamens 6–18, typ. 12; 2n=36 ...............
.............................................................. E. minuti ora subsp. minuti ora
6’ Basal leaf ultimate lobes widened, length ca. 2.5× the width; petals generally
greater than (5–) 9 (–12) mm long; stamens 6–18, typ. 14–16; 2n=24 .........
...................................................................... E. minuti ora subsp. covillei
5’ Buds with tip > 25% total bud length; cauline leaves generally with ≤ 3 (rarely
to 8) ultimate lobes, ± acute to acuminate; 2n=12 .......................E. parishii
4’ Petals > 1 cm long
8 Bud tip generally > 30% length of bud; leaves bright-green to yellow-green, ulti-
mate lobes ± acute to acuminate; cauline leaf reduced to one-few ultimate lobes
9 Receptacle 1–2 mm wide, obconic to funnel-shaped .................... E. parishii
9’ Receptacle 1.5–5 mm wide, widely-obconic to bell-shaped, often  aring at
the end of the receptacle ..........................................................E. papastillii
8’ Bud tip generally < 20% length of bud; leaves more glaucous to grey-green,
ultimate lobes ± round to acute; terminal cauline leaf typically with 5+ ulti-
mate lobes
10 Basal leaves generally with 35–40 (rarely 26–60) ultimate lobes, and ultimate
lobes ± widened appearance, length of ultimate lobes less than 2× width, cu-
neiform; (12–) 18-20 (–28) stamens often with anthocyanin spot at basipetal
end of  laments fused at the base; plants of El Paso and Rand Mountains in
Kern Co., California ...........................E. minuti ora subsp. twisselmannii
10’ Basal leaves generally with 45–70 (rarely 26–55) ultimate lobes, length of ulti-
mate lobes more than 2× width; stamens (16–) 22-24 (–32), with anthocyanin
spot at basipetal end of  laments fused at the base; plants of Riverside and San
Bernardino Counties in and around Joshua Tree National Park ...E. androuxii
Shannon M. Still / PhytoKeys 35: 45–56 (2014)
56
Acknowledgements
I would like to thank James André, Tasha La Doux, Dan Potter, Steven Still, and the
curatorial sta at the herbaria DAV, RSA, ND-G, SD, DES, ASU and UC for their
inspiration and help with my research. I would also like to thank a few anonymous
reviewers who helped edit the manuscript, species descriptions, and taxonomic key.
References
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Clark C (1997) Eschscholzia. In: Flora of North America Editorial Committee (Eds) Flora of
North America North of Mexico. New York and Oxford, 3: 308–312.
Greene, EL (1905) Revision of Eschscholzia. Pittonia 5: 205–292.
Roberts NC (1989) Eschscholzia. In: Baja California plant  eld guide. Natural History Publishing
Company, La Jolla, CA, 234.
Still SM (2011) Systematic and Taxonomic Studies of Eschscholtzieae (Papaveraceae). PhD
thesis, University of California Davis, CA.
Still SM, Potter D (2013) California Poppy Conundrums: Insights into relationships
of the tribe Eschscholtzieae (Papaveraceae). Systematic Botany 38: 104–117. doi:
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Article
We examined the phylogenetic relationships and patterns of character evolution among the California poppies and their relatives (Papaveraceae, Eschscholtzieae) using morphological comparisons and parsimony and likelihood-based phylogenetic analyses of two chloroplast and two nuclear DNA regions, including 14—3-3-like, a phylogenetically informative nuclear DNA region developed for this research. Each of the three genera in the tribe (Dendromecon, Eschscholzia, and Hunnemannia) is strongly supported as monophyletic, but not all species of Eschscholzia display monophyly based on molecular data. Examination of biogeographic patterns indicates that dispersal to island habitats occurred more than twice in the tribe. Phylogenetic reconstruction of the evolution of chromosome number suggests that there were several polyploidy events within the tribe and likely one or more aneuploid events within Eschscholzia. Based on the results presented here, the subspecies of both Eschscholzia californica and Eschscholzia lemmonii will be promoted to specific status. Additionally, the study provides evidence supporting recognition of two new species of desert Eschscholzia.
s.n. (ASU!, CAS!); wash along Eagle Mountain Rd., north along I-10
  • Mecca-Joshua Tree
Mecca-Joshua Tree Road, 7 miles southwest of junction with Interstate Highway 10, 13 Apr 1976, Curtis Clark 527 (DAV!); 2 mi S.E. of Desert Center, 1 Jul 1981, J.C. Roos s.n. (ASU!, CAS!); wash along Eagle Mountain Rd., north along I-10, 3 Mar 1995, John Wear s.n. (DAV!);
Jim André & Tasha La Doux 248 (DAV!); Base of Old Dad Mountains in wash
  • M Shannon
Sheephole Pass just to south of the top of the pass, [34°13.711'N, 115°43.19599'W], 25 Nov 2007, Shannon M. Still 222A (DAV!); Base of Old Dad Mountains in wash, [34°44.512'N, 115°45.081'W], 26 Nov 2007, Shannon M. Still, Jim André & Tasha La Doux 248 (DAV!); Base of Old Dad Mountains in wash, [34°44.512'N, 115°45.082'W], 6 Apr 2008, Shannon M. Still & Steven M. Still 377A (DAV!);
Jeff Galvin & Amy Toulsen 536 (DAV!). Discussion. While the buds of all desert Eschscholzia appear similar, those of E. papastillii most resemble E. parishii, as the bud tip is typically more than 25% of the total bud length
  • M Shannon
  • Jim Still
  • André
Clipper Mountains, just off the pipeline road, [34°40.573'N, 115°22.73502'W], 18 Apr 2009, Shannon M. Still, Jim André, Jeff Galvin & Amy Toulsen 536 (DAV!). Discussion. While the buds of all desert Eschscholzia appear similar, those of E. papastillii most resemble E. parishii, as the bud tip is typically more than 25% of the total bud length. Th e receptacular rim is prominent in this species and E. californica is the only other species that has a pronounced receptacular rim. Th e range of E. papastillii extends from San Bernardino County south to northern Imperial County. Most collections of E. parishii collected north of San Diego and Imperial Counties are likely the new E. papastillii. Eschscholzia papastillii extends at least into easternmost San Bernardino County that contains the Whipple Mountains, and likely well into Arizona.
  • E L Greene
Greene, EL (1905) Revision of Eschscholzia. Pittonia 5: 205-292.
DAV!); East of Indio on Highway 60/70, about 6 mi west of Cactus City
U.S. Highway 60/70, Indio to Blythe, 1 mi. W of Cactus City, 29 Apr 1958, P.C. Everett and E.K. Balls 23013 (CAS!, DAV!); East of Indio on Highway 60/70, about 6 mi west of Cactus City, 15 Mar 1960, W.R. Ernst 720 (CAS!);