Table Of ContentMadrono, Vol. 48, No. 3, pp. 198-204, 2001
SALTUGILIA LATIMERI: A NEW SPECIES OF POLEMONIACEAE
Terri L. Weese and Leigh A. Johnson
Department of Botany and Range Science, Brigham Young University,
Provo, UT 84602-5181
Abstract
Saltugilia latimeri, a new species of Polemoniaceae, is endemic to dry, rocky to sandy slopes and
foothills, primarily in San Bernardino and Riverside Counties, California. Saltugilia latimerisuperficially
resembles S. australis and has, until now, been considered conspecific with this taxon based on the small
corolla size shared by both species. The two species differ in several morphological characters including
stature, corolla throat coloration and exertion, and degree of glandularity.
Saltugilia latimeri, species, Polemoniaceae, A fifth taxon in Saltugilia has been identified.
taxonomy Our observations of greenhouse-grown plants and
herbarium specimens indicate that two distinct
Gilia is the historical catchall genus within Po- forms have been referred to S. australis (as Gilia
lafefmionintiyac(eWaheerforrysp1e9c4i0e,s MofasuonnceratnadinGprhaynltoge1n9e4t8i)c. australis). Herbarium collections at RSA indicate
that Latimer (1958) recognized, in his unpublished
Within this polyphyletic genus. Mason and Grant dissertation, these two forms as distinct subspecies
(1948) described a new taxon, Gilia spleiideiis ssp.
australis H. Mason & A. D. Grant, with stamen within G. australis: a typical subspecies australis,
and a geographically restricted subspecies deserto-
insertion and corolla throat and tube proportions
similar to those of Gilia spleiuiens H. Mason & A. gruuims.hiBnagsecdharoancteorbss,ervwaetiohnesreofdeasdcdriitbieonatlhisdisgteion--
D. Grant, but with a smaller corolla and a propor- graphically restricted taxon as a new species in the
tionally longer limb, similar in these respects to Gi-
genus Saltugilia.
lia caruifolia Abrams.
As characters delimiting putative natural groups Saltugilia latimeri T. L. Weese & L. A. Johnson,
within Gilia were identified, such groups have been sp. nov. (Fig. 1)—TYPE HERE DESIGNATED:
recognized with sectional or generic status. In one USA, California, Riverside County, canyon in
such case. Grant and Grant (1954) constructed Gi- pass between Whitewater and Morongo Valley,
lia section Saltugilia and included therein five spe- 0.4 miles south of the San Bernardino county
cies divided into two species groups. The section line, 9 April 1950, Verne & Alva Grant 8840
was considered to be natural from observations of (holotype, RSA!).
corolla morphology and ecology of the members.
Within their newly erected section Saltugilia, Grant Species nova ab Saltugilia australis (H. Mason
and Grant (1954) elevated G. splendeiis ssp. aus- & A.D. Grant) L.A. Johnson differt tubo corollae
tralis to the species level, as Gilia australis (H. violaceo et exserto (nec albo et incluso), lobi cor-
Mason & A. D. Grant) V. E. Grant & A. D. Grant. ollae acutis (nec cuspidatis), et calyce dense glan-
Grant and Grant (1954) distinguished G. australis duloso (nec glabro, glabrescenti, vel sparse glan-
from G. splendens and G. caruifolia by its smaller duloso).
stature, whitish flowers (pale violet to white), and Annual herbs, to 30 cm in height, scapiform with
simpler leaves. The authors also noted minimal a central stem and several basally branching axes
geographic overlap, partial to full breeding barriers, subequal to equal with the main axis. Lower stems
and apparent ecological isolation between G. aus- with long, uniserriate transparent trichomes com-
tralis and these other species. monly terminating in transparent glands (Fig. II;
Johnson (in Porter and Johnson 2000) elevated stalk cells becoming flattened and chain-like upon
Saltugilia to generic status within Polemoniaceae. drying). Mid to upper stems densely glandular with
Following rules ofpriority. Porter and Johnson also the glands multicellular, flat-topped, and translucent
restored Brand's (1907) epithet, grinnellii, in place (Fig. 1J; chloroplasts present in glands, but thesenei-
ofthe more recently used epithet, splendens, forthe ther darken beyond amber nor appear anthocyanic,
largest-flowered species. As circumscribed by Por- as is common in Gilia). Lower leaves persistent, 5-
ter and Johnson (2000), the genus Saltugilia com- many, forming a loose to dense basal rosette. Basal
prises four closely related taxa: Saltugilia australis and lower cauline leaves generally 2X-3X pinnately
(H. Mason & A. D. Grant) L. A. Johnson, S. ca- divided with 3-10 sub-opposite lobes; leafblade 20-
ruifolia (Abrams) L. A. Johnson, S. grinnellii 45 mm long (-70 mm on greenhouse grown plants),
(Brand) L. A. Johnson subspecies grinnellii and S. 6-16 mm wide (-40 mm on greenhouse grown
grinnellii subspecies grantii (Brand) L. A. Johnson. plants; Fig. IC). Rosette and lower cauline leaves
2001] WEESE AND JOHNSON: SALTUGILIA LATIMERI 199
with transparent, glandular trichomes of varying ite Mountains, Mojave Desert, 13 May 1939, Jae-
lengths (Fig. IH). Upper cauline leaves glandular gers.n. (CAS, POM); About 3 miles East ofJoshua
(Fig. IK), reduced, bract-like, entire or occasionally Tree off road to Twentynine Palms, 3 May 1964,
pinnately divided, 1.5-14 mm long (-30 mm on Thome 33975 (RSA).
mm
greenhouse grown plants) and 0.2-6 wide (-17
mm on greenhouse grown plants; Fig. ID). Inflores- Comparative morphology and relationships. Un-
cence cymose, with flowers borne singly, or paired til now, Saltugilia latimeri has been regarded as
on ultimate stemextensions. Pedicels 2-16 mm long, conspecific with S. australis. Both species possess
occasionally longer, densely glandular (cf. Fig. IL). small flowers, tend toward a smaller stature, and
When paired, pedicel of terminal (first maturing) are ecologically disposed to drier habitats relative
flower usually shorter than pedicel oflateral (second to S. grinnellii and S. caruifolia. Because conver-
maturing) flower (Fig. IE). Calyx 2.6-3.5 mm long gence in these features may result from selection
in flower, densely glandular (Fig. IL-N), with an imposed by similar habitat types and available pol-
average of 20 glands per calyx lobe on herbarium linators, they are not necessarily useful indicators
sheets (range from 6 to 35 glands per calyx lobe). of phylogenetic affinities or taxonomic boundaries.
Calyx lobes dull green, sometimes purple spotted, Flower size, in particular, has received dispropor-
0.8 mm wide, united 3/5 length ofcalyx by an byline tionate emphasis as an identification tool in Saltu-
membrane. Corolla 7.5-10 mm long, the fused por- gilia (Day 1993). As a result, small flowered rep-
tion 3-6.5 mm long, tube (=unexpanded portion of resentatives of S. caruifolia and S. grinnellii have
the fused corolla) exserted from the calyx. Tube dark been misidentified as S. australis (e.g., CAS
POM
lavender-purple, fading to blue (or yellow with ex- 627511, 48851). Similarly, it seems likely
treme age) when dried. Lower throat (=expanded that S. latimeri has escaped previous delimitation
portion offusedcorolla) yellow spotted; upperthroat by over reliance on flower size, to the exclusion of
and lobes pinkish-lavender, but also fading to pale other characters that clearly distinguish S. latimeri
blue or yellowish on herbarium sheets. Corolla lobes from S. australis.
2.0-3.5 mm long and 1-3.5 mm wide, acute. Sta- Beyond flower size, S. latimeri differs from S.
mens inserted equally in the sinuses of the corolla australis in other floral features (Table 1). The co-
lobes. Filaments 0.5-1.1 mm long to point of inser- rolla of S. australis is whitish overall. The lobes
tion, anthers 0.65-0.85 mm long and 0.35-0.50 mm are cusped at the apex and may be suffused with
wide. One stamen frequently extends at right angle pink or bluish-lavender, but the tube and throat (ex-
to the throat. Style 7-8 mm long, extending slightly clusive of the yellow spots, characteristic of Sal-
beyond the orifice. The three lobes of the stigma tugilia) remain white. Furthermore, the tube is more
0.8-1.0 mm long with tips curling downward. Cap- or less included within the calyx, the throat flares
sule 3.5-5.0 mm long and 2.0-3.5 mm wide, typi- widely, and the calyx is mostly glabrescent at ma-
cally 1.4-1.8 times as long as wide, more or less turity. These features are consistent with the type
equaling the length of the calyx. Capsule dehiscent (UC!) and populations examined from throughout
in three sections from the top to the base, 6-9 ovules S. australis' range. In contrast, S. latimeri has sat-
per locule. Seeds ± 0.85 mm long, honey gold in urated pink to lavender, tapering acute corolla
color. Testa verrucate with anticlinal epidermal cell lobes, a colored and more narrowly funnelform
walls defined as low ridges, the ridges lost and cell throat, a purple, exserted tube, and a more intensely
wall boundaries obscure when this outer epidermal glandular calyx, with the glands large and as abun-
layer is shed upon wetting. Seeds mucilaginous via dant as on the pedicel.
expansion of spiricals when wet. Pollen grains blue, Although corolla coloration provides a definitive
±40 |jLm in diameter, 5-6 zonocolpate, with lirae means of distinguishing S. latimeri from S. aus-
radiating from the apertures in magnetic-field fash- tralis on fresh and most herbarium specimens, color
ion, n = 9. may fade from Saltugilia flowers over time and can
be a less reliable diagnostic character on some her-
Paratypes. U.S.A. California, Inyo County: Han- barium specimens ofextreme age. On living plants,
aupah Canon, Panamint Mountains, 16 May 1917, flowers senesce quickly (1-2 days after opening,
Jepson 7091 (JEPS). Riverside County: Box Can- personal observation), and there is insufficient time
yon, 16 Mar 1937, Winblad s.n. (CAS); Palms to for the distinctive coloration to fade while the flow-
Pines Hwy, Mount San Jacinto, 23 Apr 1937, Win- ers remain on the plant, thus identification of S.
blads.n. (CAS); Palm Springs, 11 Apr. 1920, Spen- latimeri relative to S. australis is straightforward.
cer 1569 (POM); Pinyon Flat, 1/2 way between When color has faded on herbarium specimens, the
Palm Canyon Drive and Jeraboa Road, 15 May exserted tube, narrower throat, and glandular calyx
1992, Johnson 92-021 (BRY, RSA, WS). San Ber- are useful secondary characteristics for distinguish-
nardino County: Sandy-rocky place at base offoot- ing between S. latimeri and S. australis.
hills of Little San Bernardino Mountains, south of Relative to other Saltugilia, S. latimeri is readily
Yucca Valley, 6 May 1957, Grant & Latimer 9986 distinguished from S. caruifolia in stamen features
(RSA); Cactus Flat, San Bernardino Mountains, 25 (short filaments inserted in the sinus of the corolla
Jun 1926, Munz 10514 (POM 96484 in part); Gran- lobes vs. long exserted stamens inserted mid-
200 [Vol. 48
I
WEESE AND JOHNSON: SALTUGILIA LATIMERI 201
Table 1. Morphological Characters of Saltugiua Species.
Saltugilia australis Saltugilia cariiifolia Saltugilia grinneltii Saltugilia latimeri
corolla lobe color white/pale pink lavender/blue lavender/bright pink/lavender
pink
throat color white white/lavender lavender/pink pink/lavender
tube color white purple purple/pink purple
petal lobe shape cusped obtuse/rounded obtuse acute, tapering
tube position relative to included exserted exserted exserted
calyx
expansion ofthroat out- flares widely wide flare/cam- narrow flare narrow flare
side calyx panulate
SianiCll lllsciLlUll sinus ofcorolla mid throat sinus ofcorolla sinus ofcorolla lobes
lobes lobes
average number ofglands 3 (0-7) 4 (0-11) 7 (0-27) 21 (6-35)
per calyx lobe (min.-
max. observed)
habit one main central one main central one main central branches near base
leader leader leader
throat), and from S. grinnellii that tends to have semi-lethal gene combinations (see also Latimer
rounder corolla lobe apices and a minutely glan- 1958). Of relevance here, a cross between S. lati-
dular exterior corolla (Table 1). Saltugilia latimeri meri and S. australis produced viable Fl with pol-
corollas most nearly approach the relative propor- len viability of 11% (Latimer 1958), compared to
tions of S. grinnellii corollas, although without the 58% viable pollen in Fl progeny ofacross between
minute glands mentioned above. two populations ofS. latimeri. This reduced fertility
As implied by morphology, Saltugilia are closely in interspecific crosses ofS. latimeri X S. australis
allied species differing primarily in corollafeatures. parallels our own results. A cross of S. latimeri
In addition to these floral characters, the capsule in {Johnson 92-021) with S. australis {Johnson 92-
S. latimeri is more orless equal to the calyx, where- 011, BRY) produced vigorous Fl offspring, but
as it frequently (but not always) greatly exceeds the even hand pollination of the Fl produced only 1 to
calyx in S. australis. Differing from all other Sal- 7 seeds percapsule (average = 2.7; average number
tugilia, S. latimeri also typically branches near the ofovules percapsule = 25) compared to an average
base so that several sub-equal lateral branches soon of 23 seeds (24 ovules) and 19 seeds (21 ovules)
approach the central stem in length and diameter. in selfpollinated capsules ofthe parental S. latimeri
In contrast, other Saltugilia usually have dominant and S. australis, respectively. Pollen viability in hy-
central axes. Although the basal leaves of S. lati- brid progeny from other crosses reported by Grant
meri have broader, and fewer, lobe segments than and Grant (1954) averaged 14% between S. carui-
typical S. australis, the leaves of S. australis show folia and both subspecies of S. grinnellii, and 12%
considerable variation across its range with some between extreme forms of the S. grinnellii. The
specimens possessing broader lobes. ability of S. latimeri and S. australis to cross with
Close relationships among Saltugilia species are S. grinnellii differed substantially (Latimer 1958).
also evidenced by hybridization studies. Grant and These hybridization studies provide inconclusive
Grant (1954) concluded that species now placed in evidence regarding species boundaries and sisterre-
Saltugilia are interfertile, although these studies re- lationships within Saltugilia, but they do indicate
vealed that sterility to partial sterility exists be- levels of differentiation between S. latimeri and S.
tween some populations (from 5 to 28% inviable australis at least as great as between pair-wise com-
individuals following interspecific crosses), leading binations of other Saltugilia species.
Grant and Grant to hypothesize the presence of Were Saltugilia latimeri itself the product ofhy-
FiG. 1. Features of Saltugilia latimeri alone and in comparison with selected features ofother Saltugilia. A. habit in
early flower. B. inflorescencebranchin midflower. C. basal leaf. D.caulineleaf. E,F flower. G.capsule. H-L.trichomes
from living plant. H. basal leaf. I. Lower stem. J. mid stem (gland on right with secretion present). K. flower bract
(secretion present). L. calyx (secretion present). M. calyx gland cleared in choral hydrate. N. paraffin section ofcalyx
gland. O. scanning electron micrograph ofunhydrated seed. P. scanning electron micrograph ofunhydrated seed surface
showing verrucae and ridges formed from anticlinal walls. Q. scanning electron micrograph of untreated pollen grain.
R. capsule of S. australis. S-W. relative size and shape of Saltugilia flowers. S. S. latimeri. T. S. australis {Johnson
97-049, BRY). U. S. caruifolia {Johnson 97-099, BRY). V. S. grinnellii subspecies grinnellii (Johnson 93-098, BRY).
W. S. grinnellii subspecies grantii (Johnson 94-057, BRY). A, Q. Grant & Grant 8840, RSA. B-P, S. progeny of
Johnson 92-021, BRY.
202 MADRONO [Vol. 48
bridization, progeny ofself-pollinated plants should tions ofS. latimeri exist. These were obtained from
demonstrate character segregation. In each genera- the Joshua Tree Wilderness Area at the border be-
tion, less intermediacy and more individuals with tween San Bernardino and Riverside Counties, in
parental phenotypes would be observed. Four gen- the foothills of the Little San Bernardino, San Ber-i
erations of greenhouse grown progeny from a sin- nardino, and Santa Rosa Mountains, in the Granite
gle population have produced no segregation of Mountains of central San Bernardino County, and
morphological characters, supporting the autonomy one disjunct collection in the Panamint Mountains
of this species. Additionally, greenhouse grown of Inyo County (Fig. 2). This latter population is
plants readily set full capsules of seeds upon self- approximately 120 miles from other known S. la-
ing, suggesting that Saltugilia latimeri is autoga- timeri populations.
mous. Further evidence for autogamy is provided Several collections of S. latimeri from the Gran-
by allozyme analyses (26 individuals; Johnson 92- ite Mountain range pose some taxonomic difficulty
021) that reveal complete homozygosity at 28 sur- because they posses—s a central leaderhabit and cus-
veyed putative loci representing 19 enzyme sys- pidate corolla lobes two features typical ofS. aus-
tems: AAT, ACN, ACPH, ALD, BGAL, CAT, EST, tralis, but not characteristic of the holotype and
GDH, G3PDH-1, G3PDH-2, IDH, MDH-1, MDH- paratypes listed above for S. latimeri. However,
2, MDH-3, ME-1, ME-2, MNR-1, MNR-2, 6PGD- these collections fall within the limits ofS. latimeri
1, 6PGD-2, PGI-1, PGM-1, PGM-2, PGI-2, and can be identified based on theirexserted, purple
SKDH, SOD, TPI-1, TPI-2 (Following methods of corolla tube, narrow corolla throat, capsule that is
Soltis et al. 1983 with or without minor modifica- subequal to the calyx, and propensity for glandular
tions). Six of these 28 loci showed polymorphism calyces. The Granite and adjacent Old Dad Moun-
with at most two alleles. tains (Fig. 2) represent a locality for Saltugilia sep-
Sequences of the chloroplast matK region from arated by 50 miles from the nearest portion ofSal-
S. latimeri {Johnson 92-021) have been included in tugilia's range in the desert slopes of the San Ber-
molecular studies (as Gilia australis; Johnson and nardino and Santa Rosa Mountains. Specimens re-
Soltis 1995, Johnson et al. 1996) and analyses of ferred to as S. grinnellii, S. australis, andS. latimeri
these data provide strong support for placing S. la- have all been collected in the Granite Mountains,
timeri with other sampled representatives of Saltu- but only infrequently. Further study ofthis material,
gilia. Additional sampling of multiple populations including new accessions, is warranted. The pres-
of all Saltugilia taxa (Weese and Johnson unpub- ence of these minimally intermediate S. latimeri
lished data) with sequences of the chloroplast trnL collections in a geographically restricted location in
intron-trnFspacer and nuclearITS regions strongly no way interferes with the ability to differentiate S.
support the monophyly of Saltugilia, but provide latimeri from S. australis in other portions ofthese
insufficient resolution of taxa within the genus to species' ranges. In addition to the GraniteMountain
use these markers alone as the basis of species rec- area, S. latimeri is in close proximity to S. australis
ognition. in the Santa Ana and Little San Bernardino Moun-
Grant and Grant (1954) proposed that species tains, and with S. grinnellii in the Morongo Canyon
now included in Saltugilia evolved from an ances- area of the Little San Bernardino Mountains (Fig.
tral type, represented by S. grinnellii or a similar 2). Mixed collections ofS. latimeri with other Sal-
large flowered ancestor, occupying mild climatic tugilia on herbarium sheets have not been ob-
conditions similar to those found along the Cali- served.
fornia coast. Arising from this ancestral type were Inferred from the small number of specimens
taxa adapted to extreme conditions, including the present among Saltugilia collections acquired from
dry, arid habitats occupied by S. australis. This im- seven herbaria (CAS, JEPS, POM, RSA, SD, UCR,
plies that S. australis represents an autogamous de- UC), S. latimeri may merit conservation concern.
rivative from S. grinnellii or another similar large Though almost certainly restricted in distribution,
flowered ancestor. This hypothesis applies equally it is not known whether this species is truly rare or
well to S. latimeri. We suggest S. latimeri repre- simply overlooked by collectors because of its
sents a second, independent lineage of small-flow- small stature, inconspicuous habit, and short flow-
ered, autogamous desert annuals derived from an ering duration. The habitat of S. latimeri includes
ancestral Saltugilia species with features generally desert scrub and chaparral communities that can be
approaching those of S. grinnellii. dense and difficult to navigate, perhaps leading to
under-representation of this species in herbaria.
Ecology and distribution. Saltugilia latimeri However, more extensive collections ofS. australis
grows in dry, desert slopes ofcoarse sandy to rocky are available in several of these same areas, sug-
soils at elevations of400-1900 m. Flowering times gesting that S. latimeri is less abundant in nature.
typically range from mid April to early June, al- With human development encroaching in some of
though collections have been made as early as mid the known locations for this species, further study
March and as late as the end of June. Our survey is warranted to establish the rarity of S. latimeri.
of herbaria (CAS, JEPS, POM, RSA, SD, UCR, Whereas S. australis is frequently collected on
UC) reveals that fewer than 20 independent collec- first year burns within its range and its seeds re-
WEESE AND JOHNSON: SALTUGILIA LATIMERI 203
2001]
Fig. 2. Distribution of Saltiigilia species in California, USA. The ranges of Saltiigilia australis and S. caniifolia
extend into Baja Mexico, but these populations are not shown. Populations are represented by symbols: # Saltugilia
latimeri. + Saltugilia australis. A Saltugilia grinnellii. Saltugilia caruifolia. Locations mentioned in the text are
indicated by numbers: 1. Little San Bernardino Mountains. 2. San Bernardino Mountains. 3. Santa Rosa Mountains.
4. Old Dad/Granite Mountains. 5. Panamint Mountains. 6. Santa Ana Mountains. 7. Cushenbury Grade.
spond positively to charcoal as a germination cue grantii in Johnson et al. 1996), yet forms a well-
(Keeley and Keeley 1987), no mention of fire is supported clade distinct from Gilia.
made on any specimen labels for S. latimeri. We In their initial circumscription of Gilia section
routinely add charcoal when germinating seeds of Saltugilia, Grant and Grant (1954) performed chro-
all Saltugilia with great success, but have not rig- mosome counts as well as extensive crossing stud-
orously determined whether S. latimeri responds ies within and among species of Saltugilia. The
more positively to this treatment than it would population of G. australis collected by Grant from
without the addition of charcoal. Morongo Canyon in the San Bernardino Mountains
Taxonomic evaluation ofearlierstudies. Because (Fig. 2) used for these studies, and cited by Grant
Saltugilia latimeri previously has not been distin- (1956), is S. latimeri. Additionally, specimens cited
guished from S. australis, earlier literature may in- under the description of G. australis by Grant and
clude either ofthese species under the name "'Gilia Grant (1954) include both S. australis and S. lati-
australis''. Johnson and Soltis (1995) and Johnson meri. This is significant because the "desert race"
et al. (1996) both include, as Gilia australis {John- of Gilia splendens discussed by Grant and Grant
son 92-021), a population ofS. latimeri in compar- (1965) could be misconstrued to be our S. latimeri.
ative DNA sequencing studies of the chloroplast This clearly is not the case, however, because this
matK gene. In both studies, Saltugilia is incom- desert race of G. splendens is also discussed in
pletely sampled (represented by S. latimeri, S. ca- Grant and Grant (1954), Latimer (1958) and Grant
ruifolia and S. grinnellii ssp. grantii in Johnson and (1966).
Soltis 1995 and S. latimeri and S. grinnellii ssp. Latimer's (1958) unpublished thesis recognized
MADRONO
204 [Vol. 48
S. latimeri at the subspecies level (as G. australis 5/ Corolla tube 7-18 mm long, 2-5 X calyx
ssp. desertorum). In Grant's (1966) citation ofLat- subspecies grantii
imer's work, the populations ofGilia australis con-
sidered sympatric with G. splendens (collected in Acknowledgements
the Morongo Valley and Cushenbury Grade of the We thank the herbarium staff at CAS, JEPS, POM,
San Bernardino Mountains) are in fact S. latimeri, RSA, SD, UCR, and UC for loans ofherbarium material,
while those populations considered allopatric with J.M. Porter for discussion, G. Baird for proofreading and
G. splendens (collected inthe Santa Ana Mountains improving the Latin diagnosis, the Botany Department at
andBaCsaejdononPatshse)searoebsS.eravuasttiroanlsi,s t(hFeig.re2c)o.gnition of NRoarntghe CSacrioelnicnea DSetpaatertmUennitverastitByYaUndforthesupBpootratniyngarned-
Saltugilia latimeri as a new species in Polemoni- rseeavricehwetrhsatfoaridtheedirinhetlhpifsuclonctormimbeutnitosn., Tanhdetpowpoulaantoinoynmuosuesd
aceae is warranted. This recognition is a starting in our greenhouse studies {Johnson 92-021) was obtained
point for further species level studies of Saltugilia from a collecting trip supported by a grant from the Cal-
latimeri and larger scale investigations both within ifornia Native Plant Society, and the enzyme electropho-
Saltugilia and Polemoniaceae. To facilitate the cor- retic survey funded by NSF grant DEB 9321788 to D. E.
rect identification and incorporation of S. latimeri Soltis and LAJ.
in future works, the following key is provided.
Literature Cited
Key to Genus Saltugilia Brand, A. 1907. Polemoniaceae. Pp. 1-203 in A. Engler
(beginning with couplet 55 OE Porter and John- (ed.). Daspflanzenreich IV. 250. Engelmann,Leipzig.
son 2000) Day, a. G. 1993. Gilia. Pp. 828-836 in J. Hickman, (ed.).
The Jepson manual: higher plants ofCalifornia. Uni-
1. Trichomes on basal leaves eglandular white arach- versity ofCalifornia Press, Berkeley, CA.
niod (present at least in axils), orwhite geniculate, Grant, V. 1956. Thegenetic structureofracesandspecies
or translucent; if translucent, inflorescence glands in Gilia. Advances in Genetics 8;55-87.
long stalked, diameter ofgland less than length of . 1966. The selective origin ofincompatibility bar-
stalk and generally equally abundant on pedicel riers in the plant genus Gilia. American Naturalist
and calyx, or inflorescence trichomes including 100;99-118.
long hairs Gilia and a. Grant. 1954. Genetic and taxonomic
1.' Trichomes on basal leaves translucent, minutely studies in Gilia. VII. The woodland gilias. Aliso 3:
gland tipped; inflorescence glands subsessile, di- 59-91.
ameter of gland > length of stalk, more abundant AND K. Grant. 1965. Flower pollination in the
on pedicel than calyx; pedicels and calyx without phlox family. Columbia University Press, New York.
long stalked hairs 2 (Saltugilia) Johnson, L. A. and D. E. Soltis. 1995. Phylogenetic in-
2. Corolla tube and throat white (throat yellow spot- ference in Saxifragaceae sensu stricto and Gilia (Po-
ted), adaxial lobe surface white to white suffused lemoniaceae) using niatK sequences. Annals of the
with blue, lavender, or pink; corolla tube included Missouri Botanical Gardens 82:149-175.
in calyx, corolla lobes cuspidate S. australis Johnson, L. A. J. L. Schultz, D. E. Soltis, and P. S.
2.' Corolla tube and throat pigmented (magenta to Soltis. 1996. Monophylyandgenericrelationshipsof
pink to purple and throat yellow spotted), adaxial Polemoniaceae based on matK sequences. American
lobe surface saturated blue, lavender, or pink; co- Journal of Botany 83:1207-1224.
rollatube exsertedfromcalyx, corollalobesround- Keeley, J. E. and S. C. Keeley. 1987. Role offire in the
ed to acute (cuspidate only in Granite Mountains germination of chaparral herbs and suffrutescents.
of San Bernardino County, California, but then Madrono 34:240-249.
possessing pigmented and exserted corolla tube) Latimer, H. L. 1958. A study of the breeding barriers
3 between Gilia australis and Gilia splendens. Unpub-
3. Stamens exserted well beyond the corolla lobes, lishedPh.D. dissertation,ClaremontGraduateSchool,
inserted equally mid throat; corolla lobes often re- Claremont, CA, USA.
flexed, throat widely flaring, nearly campanulate, Mason, H. L. and A. D. Grant. 1948. Some problems in
with purple marks visible on the interior of the the genus Gilia. Madrono 9:201-220.
throat at the base ofeach corolla lobe S. caruifoUa Porter, J. M. and L. A. Johnson. 2000. A phylogenetic
3.' Stamens less than orequal to corolla lobes, insert- classification of Polemoniaceae. Aliso 19: 55-91.
ed equally in the sinus ofthe corolla lobes; corolla Soltis, D. E., C. H. Haufler, G. J. Gastony, and D. C.
lobes not reflexed, throat narrowly flaring, lacking Darrow. 1983. Starch-gel electrophoresis offerns: a
purple marks on interior ofthroat 4 compilation of grinding buffers, gel and electrode
4. Corolla less than 1 1 mm long, corolla lobes pale buffers, and staining schedules. American Fern Jour-
pink to lavendar, tube purple; corolla tube eglan- nal 73:9-27.
dular and calyx glandular S. latimeri Weese, T. L. and L. a. Johnson. 2000. Taxonomic limits
4.' Corolla generally more than 1 1 mm long, corolla and phylogenetic affinities of Saltugilia: molecular
lobes pale to bright pink, tube pink to magenta; and morphological support for generic recognition
corolla tube minutely glandular, calyx eglandular apart from Gilia. American Journal of Botany, 87
to glabrescent 5 (S. grinnellii) (supplement):166. [Abstract.]
5. Corolla tube 4-10 mm long, 1-2 X calyx Wherry, E. T. 1940. A provisional key to the Polemon-
subspecies griiviellii iaceae. Bartonia 20:14-17.
