A NEW HYBRID GENUS AND 12 NEW COMBINATIONS IN NORTH AMERICAN GRASSES

2006·3.9 MB·English

by  Mary E Barkworth

NEW
NEW
AND
A HYBRID GENUS COMBINATIONS
12
NORTH AMERICAN
GRASSES
IN
Mary
E.Barkworth
Intermountoin Herbarium
Deportment
of Biology
Utah
State University
USA
Logan. Utah 84322-5305,
mary@biology. edu
usu.
ABSTRACT
One new hybrid genus, XPascoelymus = Pascopyrum x Elymus, and 12 new combinations are pre-
sented North American grasses. Six of the new combinations arc for hybrids in the Trlticcae.
for
Stipa arnowiae transferred to Achnatherum and Hystrix californica (= Elymus calijornicus) to
is
Ammophila
Elymus
Leymus- Agropyron riparium recognized as a subspecies of lanceolatus\
is
champlainensis reduced subspecies, based on the findmgs of others, and the previously recog-
to a
is
nized subspecies of Pseudoroegneria spicata are reduced to forms. Reasons for the changes are
provided.
RESUMEN
X combmaciones
Se presenta un nuevo genero hibrido, XPascoelymus - Pascopyrum Elynms, y 12
nuevasdegramineasnorteamericanas. Seisde nuevascombinaciones sonde hibridos en Iriticeae.
las
Achnatherum (=Elymus Lcymus.
arnowiae nansiiere Hystrix californica calijornicus)
Stipa se ^
Y -d
Agropyron riparium reconoce como una subespecie de Elymus lanceolatus\ A^nmophila
se
champlainensis reduce una subespecie, basandonos en fos hallazgos de otros, la subespecie
se a y
reconocida prcviamente de Pseudoroegneria spicaia se reduce a formas. Se aportan razonamicntos de
cambios.
lo5
INTRODUCTION
two
Volume 24 in the Flora ojNorth America series will be the second of the
names.
volumes on completion requires publishing the following 12
grasses.
Its
Eleven of the 12 new combmations are for previously recognized taxa; one is a
new
hybrid
genus.
Six of the new combinations, and the new genus, are for hybrids in the
knowing which
hybrids without species
Identifying the parents of
Triticeae.
m
were present the vicinity always difficult. In most instances, have accepted
is
1
when naming one
parentage suggested by previous workers these taxa; in
the
many
known
The
instance, have compelled to disagree. distribution of of
felt
1
Two
these hybrids limited. The remaining six names affect non-hybrid taxa.
is
reflect a generic change and four a change in rank.
As
have attempted examine type material of the taxa treated. indi-
to all
I
mentioned
cated below, however, this has not always been possible. All the taxa
The
America volume
will be described and illustrated in the Flora oJNorth 24.
SIDA22(1):495-501.2006
:
496
BRIT.ORG/SIDA
22(1)
illustrations will also be available http://herbarium.usu.edu/webmanual/.
at
To the extent permitted by own
the institutions that them, photographic im-
ages of the type specimens examined are also available via this site and will be
TROPICOS. The
available via images include ups
close of individual parts
in
addition images
to of the wdiole specimen.
NOMnNCLATURAL
TREATN4ENT
POEAE
Walk
Barrington ex Barkworth, comb.
nov.
Basionym: Ammophila champlamcnsi^
F.
Nrw
Seym., Sida 2:349. 1966. Typi-: U.S.A. YORK: Lake Champlam, Au Sable Poitu, insancl
3Ju]
1902, Nellie t'lynn (llOLOTYPLi: VT).
F. s.n.
Walker, and
Paris, Barrin
sensu and
riclo Fern, A. completely glume and
St cli^nip/cn'ne^risis differ in length
flowering and
time, tend to differ in inflorescence length. They noted, however,
morphological and
that the molecular uniformity com-
of A, champlaiucnsis,
bined with the prevalence
of vegetative reproduction both
in taxa, consis-
is
with
tent the hypothesis that all populations of A. champlaincnsis^re derived
from
a single genetic individual. For recommended
this reason, they acknowl-
edging between
the distinction the two taxa at the subspecific, rather than spe-
new
cific level. I present the combination here so that may be used in volume
it
ijH
STIPEAE
&
Aclinatherum arnowiae Welsh Atwood)
N.D. Barkworth, comb, nov
(S.L.
&
Basionym: Slipa iuiwwide S.L. W^elsh N.D. Aiwoocl Utah 2003. Utah.
l-l (eel. 3J:799. U.S.A.
Kane
T43S, R4W^, mi Canyon
Co.: S 13. ca 19 o[Johni>un pinyon-juniper-sagcbrnsh-com-
1: Jet:
munity at 1740 m, on white, gypsiferous member of the Moenkopi Formation, 30 May
2001,
&
O'DeU 28062
S.L. VVeLs!) lOLOTYi'H: BRY; ISOTYPE UTC-245001!).
1. (i
&
This taxon very similar Achnathcrum
hymenoides (Roem.
is to Schult.)
havmg
Bark worth,
differing
in loosely contracted panicles with non-divaricate
branches.
TRITICEAE
XElyleymus
hultenii (Melderis ex Hultcn) Barkw^orth, comb, nov
Basion M
\'
xAgrodymus
hultenii Melderis ex tiuhen, Ark. Agropyron alaskanum
Bot., n.s. 7(1):21. 1968.
Twr
van arcticum Hulten, Aeta Univ. Lund.,
n.s. 38: 257. 1942, U.S.A. Alaska- Deerine IP
Anderson 4790
(molotype:
ALA!).
Hulten
(1968) listed the parents of this hybrid as Agropyron borcalc subsp.
&Merr.) and
67/67sfcanus(Scribn. Melderis
h'/ymiLsarenan subsp.moZ^
us
Hulten. The new combmation
is needed because these two species will be treated
&
as E. aJashanus iScrihn. Merr) A. Love subsp. alaskaniis and Leymus nwUis
(Trin.) Pilger, respectively in the Flora North America
of vol. 24.
BARKWORTH, NEW NAMES AND NEW COMBINATIONS NORTH AMERICAN GRASSES 497
IN
name
Although Hulten (1968) listed Melderis as the author of the
xAgroelymus huUenii, he did not explain Melderis' contribution in the article.
For this reason, the authorship ''Melderis ex Hulten" rather than "Melderis in
is
would
Hulten". using short form citations, be Hulten.
If it
XElyleymus ontariensis (Lepage) Barkworth, comb. nov. Basionym; xAgroelymus
CANADA.
James
ontaricnsis Lepage, Nat. Can. 79:254-2^7. 1952. Type: Ontario: Bay, Riv.
&
E
Aug
Attawapiskat, 21 1946, A. Diitiliy Lepage 16,^23 iolotype: originally deposited in
(i
NA
LCD, transferred to NA, then to US. Despite a search at both and US, has not been lo-
it
cated).
Bowden
xElyleymus according comprises hybrids be-
ontariensis, to (1967),
tween E/^/mus Gould ex Shinners[^Agrop}^rontrach_yc'auIum
t/'flch^/aiuIiLsCLink)
L
(Link) Malte ex Lewis] and umovaf us (Beal) Pilger [- Elymus innovatus
H.F.
synonym
Bowden xElyleymus
regarded as a of hiriijlorus (Hitchc.)
Beal].
it
&
Dewey
Barkw^orth D.R. but the holotype of that hybrid grows outside the
Leymus TA.
having
range of innovatus. interpreted as sintplex (Scribn. &r
L. It is
Dewey Leymus
Williams)
D.R. as the parent.
XElyleymus mossii (Lepage) Barkworth, comb. nov. Basionym: xAgroelymus
mossii
CANADA.
Near Lake open
Lepage, Nat. Canad. 92:214-215. 1965. Typh: Alberta: Louise, in
Aug ALTA
woods, 22 1946, E. U. Moss 1151 (holotype: 1044D.
The proposed combination reflects a difference of opinion concerning the par-
Lepage
ents of this hybrid as well as a difference in generic interpretation. (1965)
stated that was obvious that Elymus canadensis L. was one parent of this hy-
it
would which Agropyron
brid, but that be necessary to discover species of [in
it
the traditional sense] grew in the neighborhood to determine the other parent.
He
gave ''Agro'gyron trachycaiJun?" as a possibility Boivin (1967) agreed that
(?)
E
was
canadensisbut suggested that the Agropyron parent kgropyron violaceum
(Hornem.) Lange [= Elymus violaceus (Llornem.) Feilberg].
Elymus canadensis, however, generally absent from the region around
is
was Other
Lake Louise (Moss 1983), the area wdiere the holotype collected. fea-
tures of the holotype that are difficult to reconcile with Lepage's suggested par-
rhizomes and abundance long
entage are the presence of the of relatively soft
hairs on both the lemmas and the glumes. Elymus canadensis not rhizoma-
is
has coarse hairs on the lemmas, and has scabrous rather than hairy glumes.
tons,
Elymus trachycaulus and high elevation counterpart, alaskanus subsp.
E.
its
latiglum (Scribn. &r Sm.) A. Love, usually lack rhizomes, and have glabrous
is J.G.
or shortly hairy lemmas and glumes. More probable parents are Elymusglaucus
common
Buckley and Leymus n novatus, both of which are in the region. Elymus
i
more
glaucus has spikes that closely resemble the holotype in their posture,
awn
thickness, and length than those of E. canadensis; L. n novatus has sol hairs
t
i
on glumes and lemmas. Elymusglaucus sometimes shortly rhizomatous;
its is
Leymus innovatus always rhizomatous. Based on this interpretation of the
is
hybrid has be included xElyleymus.
parentage, the to in
498
BR1T.ORG/SIDA
22(1)
Elymus Xcayouetteorum (Boivin) Barkworth, comb. nov. xAgrohordeum
Basionym:
cayouetteorum Boivin, Nat. Canad. 94:520. 1907. TYPE; QUE!
name
Boivin published hybrids between Elymus
(1967) this for rachycauhis [=
t
Agropyron trachycaulum] and
canadensis.
E.
&
Elymus
lanceolatus subsp. riparius (Scribn. Smith) Bark worth, comb.
&r
J.G.
Stat. nov. Basionym: Agropyron nparium Scribii. & J.G. Smith, Bull. Div. Agrosrol, U.S.D.A.
Montana:
4:35. 1897. TYPE: U.S.A. Garrison, 10 Jul 1895, P.A. Rydherg 2127 (Lectoty?^: US
556672!, designated by Hitchcock 1935:776).
many
After examining specimens, agree with Dorn taxon
(1988) that this
I
merits recognition but prefer to treat as a subspecies rather than
a variety.
it
It
&
common
more
than Elymus Gould
is lanceolatus (Scribn. Sm.) subsp.
J.G.
lanceolatus.
WW
&
XLeydeum
littorale Hodgs. Mitch.) Bark worth, comb, nov
Basionym:
(H.J.
&
WW.
xHlymordeum
httorak Hodgs.
H.J. Mitch.^ Canad. J. But. 43:1355. 1965. TYPE: U.S.A.
ArASKA: Matanuska Valley, in tidal flat area along Cottonwood Creek near juncture with Knik
Arm SW WAV &
mi
about 18 of Palmer, 2 Sep 1964, MitchcU Hodgson 15S4
HJ. (l-iOLOTYPF:
ALA
29247!).
xEeydeum
httoralc consists of hybrids between Leymus mollis and Hordcum
hrachyantherum Nevski. has been Matanuska
collected in the Valley, Alaska,
It
and on the coast of Vancouver Island, British Columbia and may be more
it
widespread. Hodgson and Mitchell (1965) stated that grows along the mar-
it
gins of the high-tide zone, where waters flow The hybrid
tidal into the creek.
plants can easily be distinguished from mo by their shorter narrower spikes
L.
s
1 1 i
and more yellow and dense The new combination
foliage. reflects recognition
of the segregate genus Leymus.
Leymus
californicus ex Thurber) Barkworth, comb.
(Bol. nov. Basionym:
Gymnostichum
californicum Bol, ex Thurber in Watson, Bot. California 2:327. 1880,
S.
Gymnostichum
ailijornicum Bol. ex Thurb.. Bot. California 2:327. 1880, Hyslnx calijornka
(Bol. ex Thurb.) Kuncze, Revis. Gen. PL 2:778. 1891. TYPE: Elymus californicus (Bol. ex Thurb.)
Madrono
Gould,
9(4):127. 1947. U.S.A. Calieornia: Redwoods, near San Francisco, H.N.
&
Bolandcrs-n. (lectotype: GIl-19493!, designated by Baden, Frederiksen Nordic
Seberg,
J.
Bot. 17:457. 1997).
Transfer of this taxon Leymus supported by chromosome number 2n
to is its of
&
RAPD
and Wang
= genome-specific
56, assay results (Jensen con-
1997).
It is
with Mason-Gamefs
sistent (2001) examination of granule-bound starch syn-
thase genes in allotetraploid Triticcae. In her tree, groups with PsathyrostcJchys,
it
m
Nevski, a genus very close to us (Bod varsdottir and Anamthawat-Jonsson
J-e_y
2003). Morphological characteristics that tend to place in Leymus rather than
it
Elymus
include its well-developed rhizomes and the more or less equally promi-
many L
nent veins in its leaf blades. There are also specimens of innovatus,
particularly those from Alaska and Yukon Territory, that lack or have very
re-
duced
glumes.
BARKWORTH, NEW NAMES AND NEW COMBINATIONS NORTH AMERICAN GRASSES
IN 499
L
The
habitat of calijornicus, coniferous on non-alkaUne
forests
soils,
is
among
unusual North American
Leymus.
species of however, similar
It to
is,
some
Chinese Moench
that of species that are currently included in Hystrix
because of their lack of glumes.
Hitchcock and Baden Leymus
(1935, 1951) et (1997) included californicus
al.
in Hystrix because lacks glumes. The type of Hystrix however, Elymus
it
is,
hystrix, a species that, apart from lacking glumes and having strongly diver-
gent spikelets, morphologically, genomically, and molecularly similar other
is to
North American Elym
eastern species of
us.
XPascoleymus Barkworth, gen. hybr. nov = Pascopyrum A. Love x Leymus
Hochst.
Type; Pascoleymus howdenii (Boivin) Barkworch.
XPascoleymus bowdenii (Boivin) Barkworth, comb. nov.
Basionym: xAgroelymus
CANADA.
howdenii Boivin. Nat. Canad. 94:520. 1967. Type: Al.BERTA: Beaverlodge, 19Jul 1921,
Make
M.O. 108211 (holotype: QFA!).
xAgroelymus howdenii appUes hybrids between Agropyron Rydb.
to smithii
[=Pascopyrum and Elymus Leymus
smithii (Rydb.) A. Love] innovatus [=
new
innovatus] (Boivin Recognition genus Pascopyrum
1967). of the requires a
name new
intergeneric hybrid as well as a combination at the species level.
& &
Pseudoroegneria spicata inermis (Scribn. Sm.) Barkworth, comb. nov.
f J.G.
.
Stat. TiOV. Basionym: Agropywndivergensv'dY. inermeScribncr&J.G.Sm.,Bull.Div.Agrostol.
U.S.D.A. 4^:27. 1897. TYPE: U.S.A. IDAHO: 1895, Henderson 3058 (lectotype US-5566761
L.F.
Hitchcock
1935:773).
Daubenmire awn
(1939, 1960) reported that rhizome development and length
&
in Agropyron spicatum (Pursh) Scribn. Sm. [= Pseudoregoegneria
spicata
J.G.
(Pursh) A. Love] varied continuously within plants grown from He con-
seed.
cluded that the ability to produce rhizomes and unawned plants heritable,
is
that the two characters are not linked, and that which form becomes dominant
determined by environmental
at a local site conditions.
is
Plant breeders working with Pseudoroegneria spicata consider that awn
presence determined by major modified by some minor
a single gene, genes,
is
unawned
w^ith the condition being dominant. Although no one has gathered
work
the data that explicitly tests this hypothesis, extensive with both av/ned
and unawned means
accessions of the species suggests that true. that a
it is It
pair of heterozygotic unawned parents will give rise to around 50% awned off-
spring. This hypothesis consistent with Daubenmire's observations.
is
The above observations make awned
clear that the (P spicata spicata)
it
f.
and unawned
(P spicataL inermis) phases of Pseudoroegneria spicata are of
little
taxonomic The
significance despite their evident morphological difference. rea-
son making names form accommodate
for available at the level of to those
is
who
wish two
distinguish the
to
entities.
&
Pseudoroegneria spicata pubescens (Elmer) Barkworth, comb.
nov. nov.
f stat.
.
m
BR1T.ORG/SIDA
22(1)
500
Agropywn
BasiON YM: A\jj-opyron spicalum var. puJicsccns Elmci; Bot. Gaz. 36:52. 1903. spicaiu
subsp./nJ;crN!L-iiU{n]P!pcrXonirL\SAfatLllerbJl:147.1906, na/n.su|5crf/TYPE:U5^
m
INGTON: Kittitas Co.: Ml. SLuariJul 1898, A.D.E. FAmcr 115S(\iOLOJ\vii: Deposited Stanford
Herbarium which has been transferred to CAS; despite a search, could not be located at
it
CAS:
ISOTYPE: US-1817092').
known
Plants of Pseudoroegncria spicata with densely pubescent leaves are
from the east slope of the Cascade Mountauis in Washington. Because plants
Wash-
with nearly as densely pubescent leaves are found elsewhere in southern
them
ington and northeastern Oregon, seems best to recognize as a form,
it P.
spicata pubesccns, commensurate in this respect with the level of recognition
f.
awned and unawned
given the phases.
Publication of the above connbinations automatically generates
from
Pseudoroegneria spicata (Pursh) A. Love spicata with priority dating 1814,
f
.
basionym
the year of publication of the for the specitic epithet.
ACKNOWLEDGMENTS
thank the curators of the herbaria from wdiich have borrowed the specimens
I I
m
who me
cited in this paper, and those at other herbaria have assisted kicating
the types that have been transferred from their original institution. also thank
1
Kanchi Gandhi clarifying some of the nomenclatural issues involved James
for
Zarucchi, and Kathleen Capels for assistatice in obtaining copies of the relevant
literature, and Paul Peterson for his prompt and helpful review
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COMBINATIONS NORTH AMERICAN
GRASSES
IN
501
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