Table Of ContentNEW
A
COMBINATION THE BROMUS
CATHARTICUS
IN
COMPLEX BROMEAE CERATOCHWA)
(POACEAE: SECT
Ana
Maria Planchuelo
HerbarioACOR
Facultad de Ciencias Agropecuarias
Unlversidad Naclonal Cordobo
CONICEl de Correo 509
Casilla
5000 Cordoba^ ARGENTINA
ABSTRACT
A new
combination, Biivnus ailharlicus var. data Dcsv.) Planchuelo proposed. This taxon has
(f:. is
hitherto been treated either as B. stamincu^ E. Dcsv. or as B. cehadilla Stcud, Recent morphological
evidence supports its recognition as a variety of B. catharticus- Detailed illustrations are included
synonym
along with a morphological description, geographical distribution, and representa-
list,
specimens.
tive
Key Words: Bromiis, Ccratochloa, Poaceae, Bromeae, South and North America
RESUMEN
Sc propone la nueva combinacion Bvomus calhariicus var. data (E. Desv.) Planchuelo. Este taxon fue
previamente tratado indistintamente como B ^tamincus Desv. o cehadilla Steud. Evidcncias
E. B.
como
morlologicas recientes validan su ubicacion una variedad de
B. catharticiis. El trabajo se
complemcntaconunadetallada
ilustracion^descripcion morfologica, listadesin6nimos,distribuci6n
geografica cspccimcnes representativos.
\'
During the course of studying materials of Bromus deposited in different her-
AHUC
specimens from (Holmgren
baria, several 1990) were sent by the
et
al,
Some
curator for identification. of the specimens agree with the morphological
features of a native species of Bromus from South America which was identi-
Camara
fied by Hernandez (1978), Matthei (1986), Nicora and Rugolo de Agrasar
(1987), and Zuloaga et al. (1994), as B. stamineus E. Desv, and by Gutierrez and
Pensiero (1998), Planchuelo and Peterson (2000), and Pavlick (2003 as
et
al. B.
)
The
cehadilla Steud. native area ot distribution of this taxon lies primarily in
Andes
Southern and
the of the Patagonia region of Argentina Chile (Gutierrez
&
Pensiero 1998). In Chile grows along the Andes and coastal plains (Matthei
it
and on Robinson Crusoe thejuan Fernandez
1986), Island in archipelago (Baeza
2002). In Argentina grows in western Patagonia (Camara Hernandez
et
al.
it
1978) and on the island of Tierra del Fuego (Moore 1983).
many
This grass w^as introduced years ago in Central California for experi-
now
mental purposes and North America
(Hall 1955) naturalized
in as re-
it is
Munz
ported and Keck Wilken and
in Painter Kartesz
(1959); (1993), (1994),
Pavlick (1995) and Pavlick (2003). Herbarium specimen observations pro-
et
al.
SIDA 22(11:555-560.2006
BRIT.ORG/SIDA
556 22(1)
common
vide testimony that a garden and orchard weed, growing in dis-
is
it
turbed soils in Central and Northern coastal areas of Cahlornia in the vicinity
of San Francisco. Also, tliere are records of in northern Oregon and southern
it
New
Washington (Pavlick 1995). has also become naturalized in Zealand
It
&
(horde Edgar under the names stami cus and valdiviauus, liaving
1995), B. u B.
been introduced as part of a program oi seed sample trials to screen dilicrcnt
grasses their potential lorage value.
lor
Phenetic analyses, based on 16 morphological characters, types and rep-
ot
resentative specimens of different taxa of sect. Ceratochloa (including ccbci-
B.
(Planchuelo indicated that cchadiUa and catharticus should
diUii) 1991), B. B.
be considered conspecific suggested by Peterson and Planchuelo
varieties, as
(1998). This variability within this group of grasses lias led to the description
of numerous taxa at specific and varietal levels based on size, number of nerves,
lemma
pubescence of the glumes and lem mas, and the length of the awMis. Peter-
son and Planchuelo nomenclatural ambiguity some
(1998) clarified of of
tlie
the names by accepting the new^ combination toiharticus van niprsfn's, leav-
B.
Massa
ing the status ol other taxa for future investigation. et al. (1997, 2001,
2004)— on molecular and morphological 30
the basis ot their studies of
mem-
germ plasm accessions from Patagonia— recommended treating octoploid
complex and
bers of the catharticus as a distinct species, coJoratus, the
B. B.
hexaploid mcmhcrsasB. catharticus. Within catharticus, they recogn\zcc\ two
B.
and compared morphol-
subspecies, subsp. catharticus subsp. stamincus. the
1
ogy and verified the application of the relevant names by examining type ma-
members
Bromus
complex. Recognition
terials for o[ the catharticus of B.
sttN)iincus and B. cehadilhi at tlie varietal level seems warranted. At this ranlc,
the epithet data, based on Bromus unioloidcs var data Desv, taxonomic
a
E.
synonym, A
has priority. detailed description, s)Mionyms, representative speci-
mens, geographical distribution and illustrations Bro}}]us catharticus var
oi
data Desv) Planchuelo are provided below.
(E.
Bromus catharticus var data (E. Desv.) Planchuelo, comb, nov (Fig. l). Basionvm;
Bromus
niiioh>idesydy.c]aUi Desv. in Gay, FLChil. 6:438. 1854. /Jrc/nn.s iiJiio/(JiJL-.s [.(.7(ini.s(n.
F:.
C
CHILD
Desv.) Allen .^r Thell. ex Kloob, Ned. Kruidk. Arehief 1917:164. 1^)18. TvrP: Santiago,
CONCl
Gen' s./i-CrrcTOTVPE designated here: ISOLrelOTVrr: [4ioto ACCIR'. SGO). SVNTYPrs
K!, P;
Andes
in text: CI LI:: Santiago, 67 Ga^'s.n.: de Santa Rosa, E.F.Pocppl\:^s.)i.,Conccpc\6n,J.S.C.D.
1 1
rUrviUc
s.fi.
i
BwnuiscchadiUa Stcud., Syn. PI Glumac. 1:32L Apr 1854. TvPR PROTOl.OGun: Cldll !: Raneagna,
BAA
1828, BcrUnv 117 ILncTOTVPr designated here7 P fragm. ex DS-865524!; photo SCO,
i;
P.
CONC; CHILD
Syntyphs
iSOrrCTOTYPi:: MO!), in text: his.Juan Fernand,, C.G. Bcricro //7, IIS,
861,1411 {Bcricw ii7aLso holotypc of Broiuiis stamincus Desv).
F.
Biomus :^idmn\cus F. Desv. in Gay FL C7hil. 6:440. 1854. CcrtUochloa sUu^iinca (E. Des\'.) Staee.
Watsonia 18:413- 190L Bwinus talhdrl'uus subsp. sUimincus Desv) Massa.. Canad.
(E. l3ot.
J.
C
82:136-144, 2004. Typi: i^-iOTOLOGun: CI IILF: Raneagua, 1829, G. Bciicw 117 (MO]_OTYPE:
P,
CONC,
fragm. ex US 865470!; ibOTYPP: MO': photo. SCO).
P,
PLANCHUELa BROMUS CATHARTICUS COMPLEX
557
/
ji
^
L
/;
(li
mm
I
1
\Nhi
^1
*
N
m.H
]\
>p
mm
1
mm
i
»
t /,
i
\f
«
'ii
t\
J
eromu5
Fig cflf/?flrf/c(/5 var. etofl. A. Habit. B, C, D. Spikelets. EJIoret. A, B, E, /W^
.
mUQ,
Howell 29056
D,
BRIT.ORG/SIDA
22(1)
558
BriMn us valJiviti Phil., Lmnaea 29:102. 1858. Cerah) J) ioti va/Ji^'icnuKPhiUHolub., Folia Gcobot.
/I it.s
Phytotax. 8:171^973. Typi^ CHILI!. Pmv.Valdivia;R,AVPfi /ipi^iyiLCilOi.eTrvi^^^
I
cm
Culms 40-100
base
Usually perennial occasionally annual, caespitosc. tal
1,
with some sheaths from previous seasons. Sheaths glabrous or
of the plant
membranous,
pubescent open on the upper quarter part. Ligule gla-
slightly
mm
cm mm,
10-30 4-7
brous, 2-4 long, apex dentate. Auricles absent. Blade x
glabrous or slightly pubescent on the upper side. Panicles erect, semi-con-
flat,
cm
cm more 2-10
tracted, 10-20 long, with 15-30 or spikelets, pedicels long.
mm
com-
Spikelcts 15-25 long, with 4-7-florcts, florets imbricate, laterally
mm
6-8
Glumes glume narrowly ovate-lanceolate,
glabrous, the
pressed. first
5(-7)-nerved, occasionally only 3 nerves conspicuous, 2 additional nerves
long,
mm
glume 7-10 7(-9)-
evident the base, the second ovate-lanceolate, long,
at
Lemmas
nerved. ovate-lanceolate, glabrous, usually smooth, occasionally sca-
mm
awn
brous on 10-12 long, 7-9-nerved, apex bi-denticulate; the
parts,
all
mm mm
with
sub-apical, straight 5-10 long. Paleas 10-12 long, strongly keeled,
mm
nerves and adherent the caryopsis. Anthers 3-4.5 long. Caryopsis
ciliate to
mm
narrow
7-8 long, with a deep, furrow.
South America, B.ai(liarticusvare/nfa
ri/nau)ntniJ/ud^it^U.-Native
Di5( to
grows along the Cordillera de Los Andes from Peru to western Patagonia of
Robmson
Argentina and and on the islands of Ticrra del Puego and
Chile,
North America where occurs Central and North-
Crusoe. adventivc in in
It is it
grows
north-
ern coastal areas of California. Pavlick (1995) reported that also in
it
Oregon and southern Washington but no records substantiating his report
ern
New
known comm.
(Barkw^orth, pers 2006). Reported as naturalized in
are
m
&
Zealand (Forde Ldgar 1995). Grows disturbed soil, waste places.
Common names—
\n Argentina "cebadilla," in Chile: ianco,'' "llanco," "ce-
USA
badilla,""pasto del pcrro" (Matthei 1986) and in "Chilean brome" (Pavlick
1995).
specimens ARGENTINA. Chubuc npto.CushaiTien:Cholila,15Jan IQOlJJhns
Representative studied.
J92 (CORD). Dpto.Futalcufu: ragoFutahiuf quen^9jLUilQ48, S()ria no 2868 (BAA); 3 Feb N55^
Region Corcovado, 20 Dec
197S1 US); 5 Feb 1955, Bnrkart 19818 (SI, US). Dpio. l.anguinco: del rio
(Si,
W
1901, niiu 2S (CORD), 71- 43'^ S, 4-6 Mar 1901, lUm 7.1 {CORD). Ncuqucn: Dpio. Lacar: Farque Na-
cional Naiiucl Huapi, LagoTraful,? Nov 1949, Bockkc cW.3646(BAA);San Martin de los Andes, Dec
ci
Pdvodl 13212 (BAA). Rio Negro: Dpto. Barilochc: Fago Nahucl Huapi. 900 msni, 8 Feb 1934,
1938,
PdWiU 11748 (BAA). Dpto. Barilochc: Barilochc, Farodl 155691/2 (BAA); Ccrro Catcdral, Dec 1961.
Ell€nbc}-^1059 (BAA). ClIlLt. Region dc Aniofagasta: AntoFigasta, Qucbrada Cerisso, road to air-
II
port S of Antofagasta. 27 Feb 1939. Beetle 26i8S (MO, US). Ill Region dc Atacama: Atacatna, 27 Feb
1939. Beetle 26J8S(MC^). IV Region de Coqnimbo: Dpto. lapel, Cboapa.desvio Pola iiasta tunel, 1200-
11
An
1350 msm,12 0ct 1945, J]iCse2]2S(Llij.V Region de Valparaiso: Valparaiso. WegcnJ895, Bu,htien
SM (US); Zapallar, Feb 1920, Holwayet al30S(VS).\n Region del Maule: Maule. Ar?-o_)'o960S9(CONC,
1
MO). ConstitLicion, Oct 1891 Philippi (US). VIIl Region del Bio Bio: Concepcion: 29 Oct 1919Jiolway
s,n,
al L5t}(US), Oct 1925. Claude-Joseph 179 lUS). IX Region de la Araucania: Cajon, Dec 1942, Claude-
ei
Joseph 58/2 (US); X Region de los Lagos: Valdivia, 1888. Ph ilippi s.n. (US). PliRU.Jimin: Tarma, 11^^ 56' S
75'- 56' W, 3780 msm, Jan 1983, Smith 2980 (MO). La Libertad: Trujillo, Cerro Cabezon, 500 insm, 4
7
PLANCHUELO, BROMUS CATHARTICUS COMPLEX
559
&
Nov Sagastcgui Lopez 11005 (MO). UNITED STATES OF AMERICA: CALIFORNIA: Alameda
198.3^
Co.: Gill Tract Albany, 5 Junl944, Sf c^^InsJK 436-i (AHUCJ; Berkeley, 29 Apr 1916, S/ii([h
^^
2029 (AHUC); 14 May 1942, Beetle 3267 (AHUC); 27 May 1942, Harlan 2745 (AHUC); U.C. Campus,
H
Oxford
St. opposite University Ave., 3Jun 1924, Buckley (A) lUC); in front of Helgard near Oxford. 4
AHUC
May
2028(AHUC)-
1923, Kennt^Jy^s.n, 2027, Fresno Co: Kings River Canyon near Copper Creek,
5000 ft elevation, 2 Aug 1958, Howell 34232 (AHUC). Humboldt Co.: Fortuna, Fair grounds experi-
AHUC
mental area, 19 Aug 1947, Murphy 26298 (AUIJC). Mendocino Van Damme
s.n, Co.: State Park,
Damme
Mendocino and ca. 1 mi up Little River from Van Beach, along Fern Canyon trail, Jul 1988,
14
Bowcnlt 1099 (A\ IC)- Marin Co.: Tiburon Blvd., 150 vards F of Blackfield Dr, 26 Autz 1961 Penalosa
1
1
2220 (AHUC). Napa May
Co.: Oakville, Flev ca. 160 20 1951, Raven 2836 (AHUC); Calistoga.
ft, alti-
tude ca. 300 ft, 5 Mayl957, Raven 10788 (AHUC). Sacramcnio Co.: Brown's Ranch, Flk Grove, 25 Apr
W
DAV (AHUC
Cakin
1947, .s.n., 10032 San Luis Obispo Co.: Adelaide Road, 4.7 mi of Paso Robles,
),
May
1700 ft, 21 1960, Twisselmann 5970(A1 lUC). San Francisco Co.: San Francisco, Jackson Street, 2
Jun 1953, Howell 29056 (AHUC). Sonoma Co.: Bluffs above the sea, Sonoma Coast State Park, along
Hwy
State 0.9 ml N of Salmon Creek, 11 Jul 1957, Cramplon 4326 (AHUC). Yolo Co.: UC Davis
1,
agronomy May AHUC
experimental area, plant introduction nursery. 1950, Cramplon 042728
s.n.
(AHUC).
Planchuelo and
Peterson (2000) recognized eight species in the South Ameri-
members
can of Ceratoch Massa al (2004) proposed recognizmg only
sect. loa. et
two species, catharticus and coloratus, with two subspecies within
B. B.
B.
catharticus. In this paper, one of the species recognized by Planchuelo and
Peter-
son (2000) as B. cchadilla is reduced to a variety The status of the otlier species
recognized in the mentioned paper merits further study
ACKN0WLEDGN4ENTS
The Mary
author thanks Barkworth for the loan of specimens and her construc-
comments
on
tive the manuscript, Victor Finot for sending photographs and
information of type materials and to the curators of the listed herbarium for
sending specimens on Kanchi Gandhi comments
provided on
loan. useful the
nomenclature.
RFFFRENCHS
CM,TF
and
Baeza Siuessy, C. Marticorena. 2002. Notes on the Poaceae of the Robinson
Crusoe (Juan Fernandez) 54-1
Islands, Chile. Brittonia 54:1 63.
CamaraHernandezJ.1
978. Srom^ys. M.N. Correa,ed. Flora Patagonica.Coleccion Cientifica
In:
Buenos
INTA
8 (3):77-93. Aires.
New
and
FoRDE, M.B. Edgar. 995. Checklist of pooid grasses naturalised Zealand.
E. in
3.
1
New
Bromeae and
Tribes Brachypodieae. Zealand 33:35-42.
Bot.
J.
Gutierrez, H.F.and J.F Pensiero. 998.Sinopsis de las especies argentinas del genero Bromus
1
(Poaceae). Darwiniana 35:75-1
14.
Hall, B.M. 1955. Genetic analysis of interspecific hybrids the genus Bromus, section
in
Ceratochloa. Genetics 40:1 75-1 92.
HoLMGREN,RK.,N.H.HoLMGREN,and L.Barnett. 1990. Index herbariorun^.Part l:The herbaria of
New
the world. 8th ed. York Botanical Garden, Bronx, NY.
1
560 BRIT.OR6/5IDA 22{1)
KarteszJT. 994. A synonymized checklist of the vascular flora of the United States, Canada
1
Volume Timber
and Greenland Checklist. Press, Portland,OR.
1
Bromus
and Morphological
Massa, A.m., K.B Jensen, S.R Lakjun, D.J. Hoi 2004. variation in
f.
germplasm Canad 136-
[.Ceralochloo of Patagonia. J. Bot. 82: 144.
Massa, A.N,. S.R. Larson, K.B. Jensen, and DJ. Hole. 2001 AFLP variation in Bromus Section
.
Germplasm Crop 41:1609-1616.
Ceratochloa of Patagonia.
Sci.
A.H.Zapp[,R.Gndullo, AcuNA, and 1997. Collecting Bromus Lin the
Massa, A.N., H. I.Seguel.
Patagonian Andes. Plant Genet. Resour. Newsl.
1
genero Bromus L(Poaceae) en Gayana 43:47-1
MATTHEt,0. 1986. Chile. 10.
El
Moore, D.M.I 983. Flora ofTierra del Fuego. A, Nelson-Missouri Botanical Garden, England-
USA.
Combine
MLiNz,RA.and D.D.Kick.1959.A California Supplement, 1968; edition 1973.
flora,
University of California Press, Berkeley.
NicoRA, E.G. and Z.E. Rugolo df Aurasar. 987. Los generos de Gramineas de America Aus-
1
Hemisferio
Sur.
tral.
]995. Bromus L of North America. Royal British Columbia Museuna, British Co-
Pavlick,
L.
lumbia, Canada.
RM. and 2003. Bromus. Soreng, PM.
Pavuck, A.M. Planchuelo, PriERSON, Soreng.
L.E., R.J. In: R.J.
Peterson, G. Davidsc, Judziewicz, PO. ZuloagaJ.S, Filgueiras and 0. Morrone, 0. eds.
E.J,
New
US
Catalogue of World grasses (Poaceae): Subfamily Pooideae.Contr. Natl. Herb.
IV,
Smithsonian Washington, DC. 54-1
48:1 -730, Institution. Pp. 89.
1
and A.M. Planchuelo. 1998. Bromus catharticus South America (Poaceae:
Peterson, in
P,
Bromeae).Novon
8:53-60.
Planchuelo, A.M. 1991. Estudios sobre complejo Bromus catharticus (Poaceae),
el
I.
Evaluacion estadistica de los caracteres taxonomicos. Kurtziana 21 :243-257.
A.M. and RM.PLTERsoN.2000.The species of SromLys (Poaceae: Bromeae) South
Planchuelo, in
and and CSIRO
America. ln:W.L.J. Surrey J.Everett, eds. Grasses systematics evolution.
89-1
Publishing, Australia. Pp. 01.
WiLKEND.H.and 1993. Bromus.l Hickman, ed.Jepson manual: higher plants
E.L.Painier. n: C,
J.
of California. Univ. California Press, Berkeley.
and
Zuioaga, RO., E.g. Nicora, Z.E. Rugoio df Agrasar, O. Morrone, Pensiero, A.M. Cialullla.
J.
1994.Catalogodela Poaceae en Republica Argentina. Monogr.Syst.Bol. Mis-
familia
la
souri Bot. Card, 47:1-1 78.