NOMENCLATURAL INNOVATIONS IN PHLOX (POLEMONIACEAE), WITH UPDATED CIRCUMSCRIPTION OF P. CAESPITOSA, P. DOUGLASII, P. MISSOULENSIS, AND P. RICHARDSONII

2009·22.3 MB·English

by  James H Locklear

NOMENCLATURAL
INNOVATIONS PHLOX (POLEMONIACEAE),
IN
WITH
UPDATED CIRCUMSCRIPTION OF
AND
RICHARDSONII
DOUGLASII, MISSOULENSIS,
CAESPITOSA,
P. P. P. P.
James
Locklear
H.
7431
Briarhurst Circle
USA.
Nebraska
Lincoln, 68506,
Phlox condensata subsp. covillei (E.E. Nelson) Locklear, comb, et stat. nov.; P]
Locklear, comb, et stat. nov.; Phlox villosissima subsp. latisepala (Wherry) L<
Phloxes are components of a remarkably diverse array of plant communities and ecological systems across
common
North America, and are often among the most herbaceous species in the associations in which they
Examples Olympic Mountains,
occur. include Phlox in subalpine grasslands of the muscoides in
diffusa P.
Wyoming
Rocky Mountain and
Basin rock outcrop communities, pulvinata in alpine subulata
fell-fields,
P. P.
and
United
in barrens glades in the eastern
States.
A
Wherry
The most recent monograph on the genus Phlox (Polemoniaceae) that of (1955). forthcoming
is
number
manuscript on nomenclatural changes proposed here
Phlox (Locklear, in prep.) requires for a of taxa,
to make these names available for the work. Four of these are changes in rank from variety to subspecies
who
consistency with concepts of rank in the forthcoming work and those of Wherry, used "subspecies"
for
in preference to "variety." The concept of subspecies used here to delimit population systems that exhibit
is
and
geographic
variation distinctive ecological associations.
new
proposed combination updated
the of condensata subsp. requires
Justification for P. covillei cir-
new
cumscriptions of caespitosa and douglasii, which are provided here. Justification for the proposed
P. P.
updated
combination of richardsonii subsp. borealis requires circumscription of richardsonii, also provided.
P. P.
an updated circumscription provided.
Additionally, of missoulensis
is
P.
NOMENCLATURAL
Journal of the Botanical Research Institute of Texas 3(2)
comb,
ina Coville subsp. lutescens (S.L. Welsh) Locklear, et stat. nov. basiqnym: Phlox aus-
&
- May
Rimrock blackbrush, ash, squawbush, comm., 11 1983, S.L. Welsh, B. Welsh M. Chattzrley 21972 (holotype: BRY;
elev.,
ft
paper on austromontana complex, Welsh (1985) recognized four varieties—the first-described
In his the
P.
and most widespread form austromontana) plus three intergrading expressions (vars. jonesii, lutescens,
(var.
and much more occurring in the canyon country of southern Utah and ad-
limited distribution
prostrata) of
jacent Arizona. These taxa are distinguished at the rank of subspecies because of their discrete distributions
been
and unique The combination austromontana subsp. prostrata has published
ecological associations.
P.
(Wherry
previously 1939).
comb,
Phlox condensata Gray) E.E. Nelson subsp. covillei (E.E. Nelson) Locklear, et stat. nov. Basionym:
(A.
Wherry
Mountains, 08 Jul 1891. E Covilk with F Funston 1801 (holotype: destroyed by fire: CAS; neotype, designated by 1942: US!;
V.
B, GH! K, NY, internet image! PH, digital image!).
isotypes:
from
The taxonomic history of condensata complex. Gray (1870) described caespitosa var. condensata
P.
P. is
Rocky Mountains
Southern
type material collected by Charles Christopher Parry in alpine habitat in the
NY, PH, Nelson (1899a)
of Colorado (Hololtype: GH!; Isotypes: ISC internet image! digital image!). Elias
Wherry
elevated to species rank as condensata. (1941, 1955) reduced to P. caespitosa subsp. condensata,
P. it
it
As
but Cronquist (1959) elevated back to species rank. detailed in the circumscription of P. caespitosa that
it
rank
condensata treated here the of species.
follows, at
is
P.
White Mountains
Nelson described from type material collected in the of eastern
Elias (1899) covillei
P.
West occur
and subsequent exploration of the Intermountain revealed this entity to in alpine/
California,
subalpine situations in a number of other high elevation mountain ranges of the Great Basin as well as in the
Wherry taxon but Cronquist
Sierra Nevada. (1942, 1955) affirmed recognition of this at the species level,
Wherry
reduced synonymy under condensata. (1969) disagreed, defending recognition of
(1959) to P.
it P.
from
on morphological and complete geographic separation condensata
the basis of differences
P.
covillei
am
Rocky Mountains, but Cronquist (1984) was not convinced, noting, unable to distinguish the
of the "I
Wherry
Colorado from more western While morphological differences cited by are not
plants materials."
because
profound (see key below), the taxon covillei warrants recognition as a subspecies of its entirely
pine
separate range, relationship paralleled in the similar bi-centric distribution pattern of the bristlecone
a
and Intermoun-
complex, with the Southern Rocky Mountain expression recognized as Pinus aristata the
with pine
tain expression as Pinus longaeva (Bailey 1970). Phlox condensata is strongly associated bristlecone
New
and Mexico and
subalpine woodland, subsp. condensata with Pinus aristata in the Rockies of Colorado
mountain and Nevada.
subsp. with Pinus longaeva in Great Basin ranges in California
covillei
NELSON
TO THE OF PHLOX CONDENSATA GRAY)
KEY SUBSPECIES (a. E.E.
New
cence herbage glandular-pubescent; Southern Rocky Mountains Colorado and Mexico
in
made
by
confused
and condensata the
>ting ecological literature in relation to is difficult
floristic P.
lature surrounding this species and subspecies. Literature pertaining to subsp. condensata (generally
its
Cox
1933
Baker Boyce 2005; Cockerell 1903;
idensata) includes these: 1983; et (as P. caespitosa);
al.
&
&
Langenheim
n Hartman Rottman 1988 Kiener 1962
Mitchell 1979; 1985, pulvinata); 1967;
(as
P.
647
Nomenclatural innovations Phlox
Locklear, in
&
&
Weber Wittmann Wooton
Shepherd 1975 2001a, 2001b; Standley
(as caespitosa)]
(as caespitosa)] P.
P.
1915 caespitosal Literature pertaining to subsp. covillei (generally as P. covillei) includes these: Billings
(as
P.
& &
&
Thompson Sydoriak 2001 condensata)] Chabot Billings 1972; Clokey 1951; Goodrich
1957; Botti (as
P.
2004 Krantz 1994
Kimball condensata);
1981 Kartesz 1988; condensate)] (as
condensate); et (as P.
(as al. P.
P.
&
&
Mooney
Marchand
Linsdale 1952; Lloyd Mitchell 1973; Major Taylor 1977; 1973; et al. 1962;
et al.
&
&
&
Munz Wilken
Munz Keck Nelson Kennedy 1906 Patterson 1993
1974; 1959; (as P. dejecta)] (as P.
Peinado 2005; Pemble 1970; Sharsmith 1940 (as caespitosa var. condensate); Spira 1991
condensata)] et P.
al.
&
&
Wenk Wherry Mooney
Dawson 2007 Wright
1965.
1942;
condensata);
condensata); (as
(as P.
P.
comb,
Phlox pilosa subsp. longipilosa (Waterf.) Locklear, et stat. nov. Basionym: Phlox longpihsa Waterf., Rhodora
L.
and
from
Waterfall (1971) recognized this entity as related to but distinct pilosa. Taylor Taylor (1981)
P.
endemic Wichita
in the
reduced to a variety of pilosa. Phlox pilosa subsp. longipilosa is to granitic soils
P.
it
main
Mountains region southwestern Oklahoma, remote from the distribution of pilosa to the east.
of P. It
shown
complex
distinguished from the broader pilosa as in the following key.
is P.
many
sometimes
Stems per plant numerous, with 0-1 8 leaf-nodes; uppermost leaves and bracts alternate, as
1
mm
2-4
as 7 to 8 being so; inflorescence stems and calyx densely pilose with long jointed hairs mostly long;
somewhat awn Phlox
pilosa subsp.
twisted
calyx-lobes apically bearing relatively long,
longipilosa Locklear
(Waterf.)
Phlox sensu
pilosa
lato
on
from by John Richardson while serving
Hooker described type material collected
(1838) richardsonii
P.
BM,
Captain John Franklin's Second (1825-1827) Overland Expedition to the Arctic (holotype: K; isotypes:
Cape on
GH! NY, Richardson the type in 1826 near Bathurst the
image! internet image!). collected
digital
northwest Canada.
Arctic seacoast of
up by Linnaeus
The has long been with that of described (1753)
identity of richardsonii tied sibirica,
P.
P.
Hooker
most southern Although (1838)
from original material collected in "Asia boreali," likely in Siberia.
by Lay and
considered the two species he identified specimens of richardsonii collected in 1826
distinct, P.
no from
Kotzebue Sound on the Alaskan coast as noting the plants differed "in respect"
Collie sibirica,
at P.
The from North American continued
treatment phloxes the Arctic as
material collected in Asia. of P. sibirica
well into the twentieth century, as exemplified by the works of such notable students of the Arctic flora as
and Hulten
Porsild (1939a, 1939b) (1948, 1968).
Wherry North American from recognizing
(1955) separated the phloxes of the Arctic sibirica, P. bo-
P.
and Major morphological differences
subsp.
subsp. richardsonii richardsonii.
richardsonii alashensis,
P.
realis, P.
Wherry
by Wherry are reflected in the key below. Not noted by are significant ecological differences,
cited
being with mid- graminoid-dominated steppe communities, primarily in the
associated elevation,
sibirica
P.
(Ermakov Grubov Knyaszev
cold, dry interior plateaus of Middle Asia et al. 2002, 2006; 2001; et al. 2007;
and
Krasheninnikov Mirkin 1985; Peshkova 1972, 2001; Sochava 1945; Yurtsev 1963, 1967, 1972),
1937;
et
al.
with dwarf-shrubland
occurring almost above the Arctic Circle in association alpine
entirely
richardsonii
P.
and herb communities.
alpine
cm
and branched, forming 8-1 from
Plant an upstanding suffrutescent perennial, sparingly diffusely tufts 5 tall
cm
the erect-ascending flowering shoots 5-1 long with 5 leaf-nodes below the inflorescence;
taproot, ca.
a
h of Texas
Institute 3(2)
mm;
m 20-40
ova
pedicel length
North
complex forms western
x richardsonii recognized here as a of intergrading largely limited to the
is
and and occupying
Arctic, differing morphologically in stature size of floral parts distinct floristic
:rican
shown under
)ns of the Arctic. As in the key below, three subspecies are recognized here the first-published
new
combination subsp.
e of richardsonii, necessitating the of P. richardsonii borealis.
P.
PHLOX HOOK.
OF RICHARDSONII
SUBSPECIES
E
mm
Longest (4-)8-25
pedicels Ion
1.
borealis (Wherry) Locklear
coastal areas of the Bering Strait p.
mm
-6
Longest pedicels long; core
1
I .
The distribution of the richardsonii complex ranges from the Bering Sea region of Alaska east to northwest
P.
The
from
Canada, mostly north of the Arctic Circle, with a few highly limited occurrences reported Asia.
known Canada
from few northwest
first-described expression (subsp. richardsonii) relatively stations in
is
Canada, from Cape
and northern Alaska. The core of range along the Arctic seacoast of Bathurst in
its is
mainland promontory on
Mackenzie Northwest
the District of of the Territories, east to Clifton Point, a
Nunavut occurs on Banks Island of the Canadian Arctic Archipelago. Floristic
the coast of Territory. also
It
Aiken
and includes
ecological literature pertaining to subsp. richardsonii (generally as richardsonii) these:
P.
&
MaCoun
Holm
Kuc
Freedman 1990 Johansen
1999; 1922; 1924; 1974, 1996;
et al. et al. (as P. sibirica);
Holm
1921.
and mountains
Phlox richardsonii subsp. alaskensis occurs in the higher interior foothills of Alaska,
Yukon, and Northwest occurs most abundantly in the Brooks Range of northern Alaska,
the Territories.
It
but also in the Alaska Range of central Alaska and in the Mentasta, Nutzotin, and Wrangell mountains in
Yukon
Mountains and Richardson Mountains
south-central Alaska. also occurs in the Ogilvie of the Ter-
It
2006a
ritory. Floristic and ecological literature pertaining to subsp. alaskensis includes these: Carlson et al
&
P Cody 2002 Cook Roland 2002 Cooper 1989
(as alaskensis); et al. (as P. alaskensis); (as P. richardsonii);
&
Drew Shanks 1965 1952 Juday 1989
Jordal (as alaskensis) (as sibirica);
(as P. sibirica); (as P. sibirica) P. P.
; ;
1972 Spetzman 1959 Walker 1989, 1991, 1994
Porsild (as P. alaskensis); (as P. sibirica); et al. (as P. sibirica).
Seward Peninsula northwest
Phlox subsp. distributed along the coast of the of
richardsonii borealis
is
on
Bering has been reported the Asiatic side of the
Alaska, in the vicinity of the Strait. It also (as P. alaskensis)
&
Bering in the southeastern portion of the Chukotka Peninsula of the Russian Far East (Balandin
Strait far
On
American
Razzhivin 1980; Razzhivi 1986, 1994). side of the occurrences of richardsonii subsp.
Strait, P.
known Norton Sound and Kotzebue Sound, and from Cape Krusenstern,
from
are coastal areas of
borealis
Cape Lisburne, Cape Prince of Wales, Cape Sabine, and Cape Thompson. Floristic and ecological literature
2006b Hansen
1951
Carlson
pertaining to subsp. borealis includes these: et al. (as P. alaskana); (as P. sibirica);
Muir
Hutchison 1934 Johnson 1966 Kelso 1989 1917
(as
(as P. sibirica); et al. (P. sibirica); (as P. sibirica); P.
1994 1963
1939b Razzhivin 1986, Shetler
sibirica); Porsild 1939a, (as P. sibirica); (as P. alaskensis); (as P.
subsp.
sibirica borealis).
show genus
The geographic distribution of these subspecies parallels with patterns of speciation in the
Douglasia (Primulaceae) (Kelso 1992; Kelso 1994). also roughly corresponds to the extent of three
et
al. It
—
Nomendatural
innovations Phlox 649
Locklear,
in
by
floristic subdivisions of the Arctic described Yurtsev (1994), each reflecting regional differences in envi-
ronment and
climate. Subspecies richardsonii associated with the Central Canada subprovince, which
is is
markedly continental. Subspecies alaskensis associated with the North Alaska subprovince, which encom-
is
more
passes the continental parts of the Brooks Range and northern the Richardson Mountains,
foothills,
its
and which
Alaska with
the Arctic slope of cold, true Arctic climate, a region experiences colder winter
its
summer
temperatures than coastal areas but also experiences more heat and dryness. Subspecies
borealis
associated with the Beringian Alaska subprovince, which encompasses the Alaskan coast of the Bering
is
notably the Seward Peninsula and coastal areas of Kotzebue Sound, Norton Sound, and the Chukchi
Strait,
warm
Sea. This subprovince has an oceanic climate due to the influence of a sea current. While the sum-
mer
and more
climate along the Bering Strait generally colder, windier, overcast than continental areas
is
of Alaska, the overall climate is less harsh than in the interior mountains due to the proximity of the sea.
on on Chukotka
Species borealis also occurs the Asiatic side of the Bering Strait the coast of the Peninsula
A
number
in the parallel Beringian Chukotka subprovince. of vascular plants are considered "Beringian"
characteristic of or endemic to the Bering Sea region on both the American and Asiatic sides of the Strait
more
(Kelso 1987, Yurtsev 1994), with the floras of these coastal areas being diverse than that toward the
(Kelso
interior 1989).
Phlox
subulata setacea comb,
subsp. Locklear, nov.
L. (L.) et stat. Basionym: Phlox setacea Sp. 1:153. 1753; Phlox
l., PI.
name
on
Linnaeus established the setacea in Species Plantarum (1753) based a figure (table 98, figure
3)
P.
published in Plukenet's Phytographia (1691). The epithet setacea was subsequently misapplied by Curtis
(1798) in the Botanical Magazine for what was an illustration and description of (Loddiges 1823;
nivalis
P.
Sweet 1827) and the two names became entangled until Wherry (1929, 1937) sorted out the confusion and
Wherry
and
reinstated (1929) interpreted setacea as a glabrate expression of subulata
P. nivalis. P. P. later
name synonymy
(1955) placed the Linnaean in under subulata subsp. subulata. Reveal (1982) located
et
P. al.
specimen Herbarium Museum, London which
a pertinent in the Sloane of the Natural History (BM-SL), they
recognized typotype Plukenet Noting specimen matched
as the of the figure. that the clearly the attributes of
Wherry
what (1929) treated as subulata Reveal (1982) demonstrated that the Linnaean
var. australis, et
P. al.
name
epithet setacea has priority over and resurrected subulata a proposed by Brand
australis, P. var. setacea,
This taxon recognized here the rank of subspecies.
(1907).
is at
Phlox
yillosissima (A. Gray) Small subsp. latisepala (Wherry) Locklear, comb. nov.
Basionym: PhloxpiiosaL. subsp.
Gray (1870) described drummondii from type material collected by Charles Wright in 1851
var. villosissima
P.
on Nueces
the "Pebbly bars of the [River]" in southern Texas (holotype: GH!). Small (1903) elevated to
it
species rank, but considered villosissima an annual like drummondii. Whitehouse (1935) relocated
still P. P.
by Nueces where
the likely of Wright's collection searching crossings of the River near Uvalde, Texas
site
she found "numerous plants that closely matched the type" growing in the chalky gravel beds of the river
and determined than an
that the entity a perennial rather annual, providing further merit for recognition
is
Wherry
recognized but
as a species. (1955) the distinctiveness of this taxon, treated as pilosa subsp.
it P.
riparia.
Levin from by
(1968) reinstated villosissima as a species distinct pilosa, reproductively isolated ecol-
P. P.
&
Smith and
karyology (Levin 1966; Levin 1967) experimental crosses (Levin 1966). Phlox a
villosissima
is
tetraploid (2n = 28) and appears to be an allotetraploid. These data, coupled with those from studies of seed
&
&
protein chemistry (Levin Schaal 1970) and plant pigment (flavonoid) chemistry (Levy Levin 1971,
be between
1974) indicate villosissima could the stabilized derivative of past hybridization pilosa subsp.
P. P.
650 Journal of the Botanical Research Institute of Texas 3(2)
Evidence
and drummondii
subsp.
pilosa mcallisteri. froi
P.
more
complicated
this hypothesis but also indicate relati
2002).
Amos
by Arthur
Nelson described aspera from type material collected Heller in central
(1899)
Elias
P.
Texas. Brand (1907) discerned a relationship with villosissima, treating the taxon as P. drummondii subsp.
P.
Wherry
drummondii subvar. aspera. (1955, 1966) also recognized a close relationship be-
var. villosissima
proposing subsp.
tween the two, but, as with P. villosissima, treated P. aspera as a variant of P. pilosa, P. pilosa
homonym
was and Levin
(Wherry unavailable subspecific
noted Nelson's aspera a as a epithet).
latisepala
and occupied
same noting
the entity
(1968) reinstated aspera, citing the justifications as for villosissima
P.
P.
morphology and Both Levin
an between and in terms of ecology.
intermediate position villosissima pilosa
P. P.
on Edwards
and Wherry considered the combined range of this pair of taxa to be centered the Plateau of
common
and
most western portion the plateau
Texas, with villosissima/?. pilosa subsp. riparia in the of P.
P.
more and
occurring the north.
subsp. to east
aspera/?. pilosa latisepala
and with
two with each other
Ferguson (1998) cited possible local integration of these entities P. pilosa
noted by Wherry reason maintaining them the subspecies level pending further study.
[1966]) as for at
(also
While
combined both under
In the most recent treatment of Phlox in Texas, Turner (1998) P. villosissima.
may show morphological and
these taxa intergrade where their ranges overlap, they distinctive differences
The
complex key
ecological associations toward the outer portions of the overall range of the (see below).
and recognized here subspecies of
taxon formerly treated as aspera pilosa subsp. latisepala is as a P. vil-
P. P.
new
combination.
necessitating the
losissima,
SMALL
GRAY)
(A.
ton notably
of species range,
ri>
subsp. latisepala (Wherry) Locklear
range
portion of species
OF PHLOX NUTT.
CIRCUMSCRIPTION CAESPITOSA
Wyeth
on
by 1833
described from type material collected Nathaniel Jarvis in
Nuttall (1834) caespitosa
P.
Oregon Expedition (1832-1833). Nuttall described the type locality
the return (eastbound) leg of his
first
20 th The
on dry adding, "Flowering about the of April." date "April
"Flat-Head the sides of
as river, hills,"
22 nd " written on the holotype specimen label (BM) and on isotypes (GH, PH) of caespitosa. According
P.
is
Wyeth journal, 22 April 1833 was the day he departed "Flathead Post," also called "Flathead House,"
to
's
trading post historians place not on the Flathead River but on the Clark Fork River near the present-day
a
A
number more
town Thompson northwestern Montana (Sanders County) (McKelvey 1955). of
of Falls in
Thompson
made
been
recent collections of caespitosa have in the vicinity of Falls.
P.
open-growing form
Wherry complex
(1941, 1955) interpreted caespitosa as a consisting of a taller,
P.
more
occurring middle elevations in the Rocky Mountains, plus three shorter, compact
(subsp. caespitosa) at
under
and occurring higher elevations alpine/
expressions (subspp. condensata, platyphylla, pulvinata) at
subalpine conditions. Cronquist (1959) revised this complex, raising the high elevation forms to the rank
and
of species as condensata and pulvinata (incorporating subsp. platyphylla) interpreting P. caespitosa
P. P.
woodland and
savanna.
open-growing lower primarily in Pinus ponderosa
species of elevations,
as a
taller,
and
Wherry with considering condensata
1965a, 1965b, 1969) took exception these revisions,
(1962, P.
pulvinata mere high altitude expressions or "ecads" of caespitosa. However, the ecological divergence of
P.
P.
and
from pronounced, and, following Cronquist, condensata
these alpine/subalpine entities caespitosa is P.
P.
pulvinata are recognized here as separate species.
P.
—
Nomendatural innovations Phlox
Locklear, in
cm
and branched, forming open 5-15 the erect-ascending
Plant a subshrub, sparingly diffusely tufts
tall,
1.
cm
montane
woodland,
flowering branches 2.5-7.7 long, with internodes typically apparent; associated with
Phlox
savanna and grassland
<
more compact
growth forming mats or cushions
Plant a caespitose perennial herb, habit pulvinate, or less
1,
cm
cm
upper
2-7.5 the flowering shoots 1-3(-5) long; associated with alpine or subalpine ecological
tall,
mm
5-10
packed
sely together; (eaves appressed, erect,
Rocky Mountain Northern
As circumscribed here, caespitosa a species of the System, chiefly in the
P. is
Rocky Mountains southeastern Columbia, northern Idaho, and western Montana. occurs most
in British
It
on Mountains and intermoun-
abundantly in Montana, particularly the eastern slope of the Bitterroot in the
m
800-1500
tain valleys of the Rocky Mountain Trench. occurs at lower to middle elevations (mostly but
It
}
up 2040 m) piedmont and mountains, and with morainal
in the in association features of river valleys.
to
common
woodland and
Phlox strongly associated with Pinus ponderosa savanna, but also in
caespitosa is
is
where and
montane, and valley grasslands in the region Pseudoroegneria spicata, Festuca idahoensis,
foothill,
graminoids.
Festuca campestris are the chief
made
and by con-
the
Interpreting ecological literature in relation to caespitosa difficult
floristic P. is
name
fused nomenclature surrounding this plant, particularly in connection with the douglasii (discussed
P.
name
below), but with added problems coming from the literature of alpine ecology where the caespitosa
P.
has been misapplied condensata and pulvinata. Literature pertaining to caespitosa includes these:
to P. P. P.
& &
On
Shaw
Habeck
Lackschewitz 1991; Lesica 2002; Root 1972 (as pulvinata); 1979.
P.
CIRCUMSCRIPTION OF PHLOX D
Hooker (1838) described douglasii in his Flora Boreali-Americana. As one of the first of the small-leaved
P.
name
cushion-forming phloxes described from western North America, the subsequently served
douglasii
P.
many number which were
as catch-all for later-discovered, superficially similar species, a of treated as va-
some nomendatural
including
or subspecies of point in their histories, andicola,
rieties douglasii at P. P.
P.
and Further confusion was caused by the emergence of the term "Douglasii Hybrids"
austromontana,
P. diffusa.
which was wide cushion phlox
in horticultural literature in the early 1900s, applied to a variety of cultivars
much
Wherry on
of probable hybrid origin. brought needed clarity to this situation through his papers the
and 1955 monograph which he recognized
western phloxes (1938, 1941) his in douglasii as a distinct
P.
on Columbia Northwest.
species with a distribution centered the Plateau in the Pacific
synonymy
under under
Cronquist (1959) reduced douglasii to caespitosa, justifying this treatment his
P. P.
discussion of pulvinata which he separated from caespitosa. Cronquist described the type of caespitosa
P. P.
P.
known name
was
as "a compact plant of the taxon usually as P. douglasii." Since the P. caespitosa published
by Nuttall (1834) four years before Hooker (1838) published douglasii, Cronquist reasoned that "the latter
P.
Wherry and
(1962, 1965a) considered this "a serious misunderstanding," calling douglasii caespitosa
P. P.
among Wherry
two
"about as distinct species as can exist the Microphloxes." Indeed, (1955) placed these
species in independent subsections of the genus Douglasianae and Caespitosae. Despite Wherry's protests,
most subsequent and ecological workers in the Pacific Northwest have followed Cronquist. Based on
floristic
BM,
examination of the type material of both douglasii (holotype: K, digital image!; isotypes: digital image!
P.
BM, GH!
GH! NY, internet image! PH, digital image!) and caespitosa (holotype: digital image!; isotypes:
P.
Washington,
NY, internet image! PH, digital image!), plus study of these entities in the field douglasii in
(P.
652 Journal of the Botanical Research Institute of Texas
3(2)
As by
Oregon, and California; caespitosa in Montana), they are recognized here as separate species. noted
P.
Wherry and
dark needle-shaped
(1962, 1965a, 1969), the leaves of douglasii are green, slender (acicular
P.
or acerose), and covered by long gland-tipped hairs, while the leaves of caespitosa are pale green, relatively
P.
broad and and bear coarse glandless along the margins. The distribution
(linear-oblong), thickish, cilia
flat
on Columbia while centered in the Northern
of douglasii centered the Plateau, that of caespitosa is
P. is P.
Rocky
Mountains.
made
and by
the con-
Interpreting floristic ecological literature in relation to P. douglasii is difficult
was
fused nomenclature surrounding this species. Prior to Cronquist (1959), this taxon treated as
P.
Wherry
or the recognized by (1938, 1955) as a reduced expression of the former
latter
douglasii rigida,
P.
under the name subsp. Following Cronquist's revisions, most workers abandoned these
douglasii rigida.
P.
names
Applegate Culver
in favor of caespitosa. Literature pertaining to douglasii includes these: 1938;
P. P.
&
1964 Daubenmire Daubenmire 1968 Driscoll 1964a, 1964b; Eckert
(as (as caespitosa);
P. diffusa); P.
&
Munz
Keck Peck
1957 (as Hall 1967; Howell 1903; 1959 (as douglasii subsp. rigida); 1941;
P. diffusa); P.
&
The
Piper 1906 (as rigida); Piper Beattie 1914 (as P. rigida); St. John 1937 (as P. douglasii var. rigida).
P.
following key separates from and, additionally, from a similar-appearing
douglasii caespitosa
P. P. P. diffusa,
species that occurs in range.
and Northern and
Dng) with coarse glandless
thickish,
cilia;
flat,
w
mostly
lacking or
fine;
ia
OF PHLOX
CIRCUMSCRIPTION
Wherry from by Merton Reed 1939 on Waterworks
(1944) described missoulensis the type collected in
P.
J.
Montana PH, Cronquist reduced
Hill near Missoula, (holotype: digital image!). (1959) this entity to kelseyi
P.
Wherry
which
relationship and reduction with (1962, 1965b) strongly disagreed, calling
var. missoulensis, a
on
missoulensis "a striking endemic...wholly unrelated" to kelseyi. In her thesis the P. kelseyi complex,
P.
P.
Campbell (proposing but not
(1992) followed Cronquist in treating missoulensis as a variant of kelseyi
P. P.
publishing subsp. but cited "extremely different habitats" plus "complete geographical
kelseyi missoulensis),
P.
isolation coupled with differences in 10 morphological characters" in support of recognition of missoulensis
and
as a distinct entity within the complex. Phlox missoulensis kelseyi are recognized here as sepa-
kelseyi
P.
P.
endemic Northern Rocky Mountains
rate species (see key below). Phlox missoulensis to the in west-central
is
where Anaconda,
Montana, chiefly along or just west of the Continental Divide occurs in the Garnet,
it
and and and on mountains with
Lewis Clark, Sapphire ranges small in the vicinity of Missoula, outlying
occurrences east of the Continental Divide in the Little Belt Mountains.
and
Leaves dark green, linear to linear-lanceolate, thinnish but with subacerose ciliate surficially pilose with
tip,
gland-tipped exposed mountain slopes and habitat dry to stony and shallow
fine ridges; xeric; soils
hairs;
Phlox missoulensis
and and (becoming
wrinkly
Leaves pale green, te cuspidate, thickish succulent, fleshy
line;
on drying), the margin slightly thickened, basally c glabrous to moderately pubescent;
mountains and intermountain basins and habitat seasonally moist to wet; fine-
valleys soils alluvial,
flats;
Phlox
kelseyi
to highly-alkaline
aline
ACKNOWLEDGEMENTS
The author wishes to acknowledge the curators and staff of the following herbaria for the loan of specimens
.
653
Nomenclatural innovations Phlox
Locklear,
in
MONTU,
CAN, OKLA, RSA-POM,
ALA, BM,
and from PH,
and/or providing images label data specimens: K,
WS. NEB, was Thanks
Robert Kaul, curator of especially helpful in securing these loans. also to the curators of
GH, RM, US work from
BRIT, TEX/LL, for access to their collections. This has benefited greatly conversations
Andrew
with Carolyn Ferguson. thanks anonymous reviewers as well as Hipp, Barney Lipscomb,
Finally, to
Guy Nesom and comments on
Tony Reznicek, and valuable review helpful the manuscript.
for
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