MUHLENBERGIA TARAHUMARA (POACEAE: CHLORIDOIDEAE: CYNODONTEAE: MUHLENBERGIINAE), A NEW SPECIES FROM CHIHUAHUA, MEXICO

2009·13.3 MB·

by  Paul M Peterson

MUHLENBERGIA TARAHUMARA
(POACEAE: CHLORIDOIDEAE: CYNODONTEAE:
NEW
MUHLENBERGIINAE), A FROM
CHIHUAHUA,
SPECIES MEXICO
M.
Paul Peterson
Columbus
Travis
J.
Department
of Botany
Rancho Ana
Santa and
Botanic Garden
Museum
National of
Natural
History
Claremont
Graduate
University
Smithsonian
Institution
1500 North Avenue
College
DC
Washington,
20013-7012,
U.S.A. Claremont, 9171
California
7,
1
On
2003
a collecting the Madre
trip to Sierra Occidental Chihuahua,
in Mexico, two unusual specimens
&
were found and
initially given fieldnames oi Aegopogon
(Peterson Catalan and
17542) Bouteloua
P. (Peterson
&
when PMP
Cataldn was
17621). Later, preparing
treatments and
of Bouteloua
the Muhlenbergiinae
tribe
northeastern Mexico
for
(Peterson 2007b;
Herrera
et al. Arrieta et 2008), these specimens were
al. reevalu-
them
ated after seeing in the United National Herbarium
States
(US)
in separate
unidentified
folders of
Aegopogon and Upon
Bouteloua,
respectively.
closer inspection, these same new
collections represented
the
species that could not be
easily placed in Aegopogon nor any
or Bouteloua, remaining
of the genera
currently
placed
in the Muhlenbergiinae
(Peterson 2001b, new
et al. 2007a). Individuals of this species are unique in
having
terminal, raceme-like panicles with only two end A
spikelets the each primary
at of branch. second
PMP
opinion seemed
warranted; then
sent a duplicate JTC examination and
off to for incorporation
into
DNA
sequence
data could
sets that possibly
elucidate
of enigmatic
affinities this species. Subsequently,
a
specimen was DNA
third found
As
(Peterson discussed
et al. 8032). below, analyses of sequences from
the
new
species placed
in subtribe Muhlenbergiinae.
it
Members
of subtribe Muhlenbergiinae
are grasses (Poaceae) in the subfamily
Chloridoideae,
Cy-
tribe
nodonteae
(Peterson 2001b, 2007a,
et in review). Muhlenbergiinae
al. are highly
variable morphologically,
although the group can be
generally characterized membrane
as follows: ligule a (rarely a line of
hairs);
inflorescence rebranched composed
a panicle, or only primary
of branches;
spikelets sometimes
solitary, in
pairs or cleistogamous
triads,
spikelets occasionally present
in the
leaf sheaths;
floret (rarely more),
1 perfect,
staminate, or glumes awned unawned; lemmas
awned
sterile; or 3-nerved, unawned; chromosome
or base
&
number
x 8-10
=
(Peterson 2000;
Peterson Herrera
Arrieta 1995; Peterson Colum-
et 1997, 2007a,
al. b;
of Texas
Journal of the Botanical Research Institute 3(2)
528
NAD-ME
adenine dinucleotide cofactor
C (nicotinamide
Two photosynthesis,
bus 2007). subtypes of
et
al.
4
and by biochemi-
been found
PCK carboxykmase), have verified
(phosphoenolpyruvate
and
enzyme)
malic
&
Watson
Brown 1992).
Hattersley
al 1974; 1977;
Muhlenbergiinae (Gutierrez
occur in et
assays
cal to
important
which has 154 species including the
Muhlenbergia,
By the largest genus in the subtribe is
far,
and M.
Buckley
M.
amphitropical disjuncts arenicola
Hitchc,
montana
North American range grass M. (Nutt.)
Peterson
Asia (Herrera Arrieta 1998;
southeast
and seven species located in
ex Bush,
(Kunth) Hitchc.
torreyi
&
&
Wu
& Peterson
Peterson 2007; et
Peterson 2006; Herrera Arrieta al.
Diaz Peterson 2003;
Ortiz 1998;
where
Mexico,
number and 125 these occur in
America 127 of
North (86%),
indigenous
2007b). Species to
&
Herrera Ar-
Peterson
Davila-Aranda 2004, 2006;
2000;
endemic (Espejo Serna et al.
56 species are et al.
Peterson 2007a).
2005;
et
rieta al.
and monotypic. All are limited to the
four are
The remaining nine genera have four or fewer species,
& New
Papua
Humb.
Bonpl. ex Willd. in
New Aegopogon cenchroides
World an occurrence of
except
for
New
Papua Guinea,
North America. Apart from presence in
endemic
its
Guinea (Veldkamp 1985). Five are to
north-
restricted to
North and South America. Bealia (one species) is
distributed in
Aegopogon (four species)
is
North America
found
in
(two species)
Blepharoneuron is
Mexico 1989; Peterson 1993).
ern (Peterson et al.
and northern
southwestern U.S.A.
an important range grass in the
Nash,
and
includes (Torr.)
tricholepis
B.
& Mexico
has three species in central
Chaboissaea (four species)
Annable
Mexico 1990, 2003).
(Peterson
&
&
Annable
1992;
and (Peterson
Annable Argentina Bolivia
in
Peterson
and atacamensis (Parodi) P.M.
C.
North
&
has one species limited to
Lycurus (three species)
Sykes 1997).
Herrera Arrieta 1995; et
Peterson al.
&
Sanchez
Reeder (Reeder 1985;
C.
including setosus (Nutt.)
America and two amphitropical disjuncts, L.
&
Cen-
North,
distributed in
Morrone Pereilema (four species)
1998). is
Rugolo de Agrasar 1986; Peterson
(one
(Reeder
endemic the U.S.A. 1976). Schaffnerella
to
(one species)
and South America. is
Redfieldia
tral,
Schedonnardus (one species)
Mexico (Columbus 2002). is
known
from San et
only Luis Potosi, al.
species)
is
genus with an amphitropical distribution.
another
yet
mG
andnrDNA
rpsl6-tmK, trnL-F,
cpDNA
rpl32-trnL rps3, rpsl6-intron,
ndhF,
Analyses ndhA-intron, \
of
(Columbus Peterson al in prep.) indicate
in press; et
Muhlenbergiinae
et
ITS sequences of species in the al.
genera
smaller
monophyletic because these nine,
not
delimited above,
genus Muhlenbergia, as is
that the large
nine genera
include these
Muhlenbergia to
expanding circumscription of
the
within
nested therefore,
are
it;
&
Peterson
Herrera Arrieta 2005;
2001a, 2004; Peterson
Peterson
seems warranted (Duvall 1994; et
et al.
al.
we
new phylogenetic
present a
science,
describing a species to
addition
In paper, in to
2007a). this
et
al.
unusual
placement taxon.
of this
and support the generic
hypothesis for
&
MEXICO. Muni-
Muhlenbergia tarahumara Peterson Columbus, sp. nov. (Figs. 1A-J; 2). Type: Chihuahua.
P.M.
W
1960-2040
km 107°31'5.0"W),
and E Osichi (27°28'15.0"N,
Madre 2 of Rio Corareachi of
Occidental,
Guachochi,
Sierra
cipio
& MO!, RSA!, US!).
US-3470469!; CIIDIR!, K!,
Aug PM. P Catalan 17621 (holotype: isotypes:
m, 30 2003! Peterson
nodes
cm glabrous below the nodes, usually 3
Culms 18-35 near base,
terete
erect,
Caespitose perennials. tall,
cm
than internodes above,
0.8-8 shorter the
and Leaf sheaths long,
glabrous shiny.
per culm; internodes
mm
membranous, pubescent or
abaxially
1-1.8 long,
below;
and mostly glabrous ligules
pubescent above
mm
cm
wide,
0.2-1.3 to
(2-)3.5-13 long, flat
blades
minutely
apex often erose, ciliolate;
glabrous, acute,
0.1-0.4
on both the hairs
surfaces,
somewhat sinuous, antrorsely hirsute
acuminate,
tightly involute, apically
mm primary
cm cm with 5-13 racemosely arranged
terminal
3-6 wide, narrow,
0.7-1.6
Panicles long,
long.
cm deciduous, disarticulation
with two terminal spikelets,
branches 0.5-1.4 long,
per node;
branches,
1
culm
from
near the
curling base)
(bending sharply or
spreading
ascending then
branches
near base,
first
mm
ending
in
inflorescence axis flattened,
0.2-0.4 long;
the hairs
secund, antrorsely hirsute,
usually
axis,
mm
than
one longer the other,
0.2-0.6 long, slightly
fused or
margins pedicels
terminal branch, hirsute;
mm
mm
glumes 3-5.8 long,
one 1-flowered;
appressed another,
4.5-7.8 long, to
appressed. Spikelets
tightly
and new
Peterson Columbus, Muhlenbergia tarahumara, a species from Mexico
&
[PM
tarahumara P
gia Peterson Catalan 17621 A. Sheath, and Two
(US)]. Habit. B. ligule, blade. C. paired spikelets. D. Bran<
two
spikelets (only one seen from this perspective). E. Glumes. Floret on branch axis (glumes, other spike
F.
Stamens and
Lodicules.
pistil. J.
.
Journal of the Botanical Research Institute of Texas 3(2)
=
&JM
[PM 22053 OS
se section of the leaf blade of Muhtenbergia tarahumara Peterson Saarela (US)]. Adaxial surface is at top.
membranous
lemma,
than chartaceous, 1-veined,
narrowly usually equal in length, shorter the to
lanceolate,
mm mm
lemmas awned,
awns
pubescent, apex acuminate, awned, the 1.4-3.2 long; 3.6-7.3 long, lanceolate,
membranous apex acuminate, the
appressed pubescent, acute
chartaceous, to bifid,
distinctly 3-veined, to
mm
mm
awns
awn extending 0.6-1.3
central 2-3.7 long, straight or slightly recurved, the lateral veins into
mm
lemma, membranous, below and appressed pubescent
paleas 3-4.6 long, shorter than the glabrous
long;
mm
awn
mucro stamens
near apex, 2-veined, apex acuminate, each vein extending as a or 0.2-0.9 long; 3,
mm
and
membranous; with two
ovary glabrous
2.2-2.8 yellowish orange; lodicules styles
anthers long,
2,
Comments.—The new can be distinguished from other species of Muhlenbergia by having a
species
all
with only two the end of the branch. closely resembles species
terminal, raceme-like panicle spikelets at It
oiAegopogon, which have the same kind of inflorescence, including the deciduous branches, but with three
who
named
Tarahumara
some The people
per branch, and species otBouteloua. species after the
spikelets is
Madre
northern Occidental.
indigenous the Sierra
are to
—
known
from
and Muhlenbergia tarahumara only three locations in the Sierra
Distribution Habitat. is
Madre Occidental Chihuahua, Mexico, where occurs on rocky slopes, ridgetops, and white-tuff rock
in
it
Ae-
with Vaccinium, Comarostaphylis, Aristida,
outcrops species of Pinus, Quercus, Cupressus, Arctostaphylos,
M. montana, M. Buckley
gopogon (DC.) A. cenchroides, Muhlenbergia lucida Swallen, pauciflora
Trin.,
tenellus
1880-2075 m.
and M. (Kunth) Kunth;
M.
polycaulis Scribn., rigida
and
Peterson Columbus, Muhlenbergia tarahumara,
a
anatomy.—
Leaf Cross-sections of the leaf blade were done with a rotary microtome on fresh-field fixed mate-
&Saarela and mounted (Sharman Columbus
rial (Peterson 22053), stained, prior to observation 1943; 1999;
&
RSA
Peterson Herrera Arrieta 2001). Permanent slides are deposited and US. The following description
at
much
employs
standardized terminology proposed by
of the
Ellis (1976).
The C
Kranz which
leaf blade in transverse section displays anatomy, associated with photosynthe-
is
4
sis (Fig. 2). It is nearly flat (becoming involute on drying) and both surfaces have rounded or flat-topped,
longitudinal costal (above/below the vascular bundles) ribs alternating with V-shaped intercostal furrows;
a
conspicuous rounded midrib The up
projects abaxially. furrows are to 1/2 the thickness of the blade; those
on more The
the abaxial surface are shallow. vascular bundles (VBs) have two sheaths, an inner (mestome,
&
XyMS+; Watson PCR
Hattersley 1976) continuous sheath of sclerenchyma and an outer (parenchymatous,
tissue; Hattersley et 1977) sheath of chlorenchyma. The outer sheath uneven in outline and
al. is cells
its
round
contain
chloroplasts that are distributed centrifugally/peripherally within the (Prendergast
cell et
al.
The VBs
development metaxylem, whether
1987). differ in size, of or not the outer sheath continuous,
is
and amount sclerenchyma on and VBs
the of present the adaxial abaxial sides. There are four primary (with
metaxylem
VBs
large vessels; includes the midvein), four secondary VBs, and four arranged
tertiary as
fol-
The phloem
lows: not sclerosed. In the innermost VBs, the outer sheath
iii-i-iii-i-ii-ii-i-ii-ii-i-iii-iii. is five
is
extended adaxially into the The outer sheath interrupted by sclerenchyma on
rib. the abaxial side of
is all
VBs
except three of the four tertiary VBs, and on the adaxial side of the two primary VBs nearest the
lateral
A
narrow
midvein. layer of smaller, isodiametric, indistinctly radiate chlorenchymatous surrounds the
cells
(PCA
outer sheath tissue; Hattersley et 1977). Sclerenchyma girders (in contact with the outer sheath) are
al.
on and The (many
present the adaxial abaxial sides of all VBs. largest girder cells in five series/rows) forms
VBs
whereas
the midrib, those of the tertiary are smallest (< 10 cells in one or two series). Abaxial girders
associated with primary and secondary VBs, as well as those on the adaxial side of the two primary
lateral
VBs
nearest the midvein, are broadest near the epidermis. The blade margin has two subepidermal
or three
Bulliform midvein and become
fibers. cells are largest adaxially adjacent to the progressively smaller towards
on
the margin; those the abaxial side are smaller than their adaxial counterparts. Colorless one or
cells
two columns wide extend between bulliform on both sides of the blade, dividing the chlorenchyma
cells
with
associated adjacent VBs.
&
DNA
was from
Molecular analyses.— obtained
field-collected leaf material (Peterson Saarela 22053)
mG
and was sequenced
for ndhA-intron, ndhF, rpl32-trnL rps3, rpsl64ntron, rpsl6-trnK, trnL-F, and ITS. To
\
new
determine
the phylogenetic position of the species within Chloridoideae, the sequences were aligned
Columbus
with the et al. (2007) data set, representing 66 chloridoid genera and the Peterson et (in review)
al.
Upon new
96
data set representing chloridoid genera. analysis, the species resolved in the Muhlenbergiinae
clade. To determine position within Muhlenbergiinae, the sequences were aligned with the two separate
its
80-90%
data sets that included ten genera in the subtribe and of the species in Muhlenbergia (Columbus
all
new
et al. in prep.; Peterson et al. in prep). Analyses of the Muhlenbergiinae data sets revealed that the
member
species is a of a subclade that includes Aegopogon, M. subgen. Muhlenbergia, and Pereilema. posi-
Its
was
tion within the subclade not resolved in the trnL-F phylogeny (Columbus in although
prep.), in
et
al.
mG
combined
the ndhA-intron, ndhF, rpl32-tmL \ rps3, rpsl6-intron, rpsl 6- trnK phylogeny (Peterson in
et
al.
new
was
supported and
prep.) the species Aegopogon However, ITS
as sister to cenchroides A. in the
tenellus.
Aegopogon (Columbus
the sister of in prep; Peterson
et et
is al. al.
Journal of the Botanical Research Institute of Texas 3(2)
532
DISCUSSION
The new unique among members of the Muhlenbergiinae in having only two spikelets per branch,
species
is
number branch
per
racemosely arranged along the panicle Reduction of the of spikelets is also
these axis.
member where 2-5
M. subgen. Muhlenbergid) there are spike-
seen in Muhlenbergia diversiglumis Trin. of
(a
&
most Aegopogon
branch Annable However, M. tarahumara resembles closely in
per (Peterson 1991).
lets
(Columbus
morphology, which supports the phylogenetic hypothesis that these taxa are sister et al. in prep.;
The
from M. tarahumara in having three spikelets per branch.
Aegopogon
Peterson in differs
et prep.).
al.
rudimen-
and two (sometimes
dimorphic— and hermaphrodite smaller
one are
three spikelets are larger
is
no
M.
and staminate In tarahumara, spikelet
one often not developed in A. bryophilus Doll) or neuter.
tary;
dimorphism
evident.
is
PCK C The
presence
anatomy M. tarahumara undergoes the subtype of photosynthesis.
Leaf suggests
4
two bundle uneven outline of the outer sheath, round shape of the chloroplasts in the outer
of sheaths,
and the centrifugal/peripheral position of these chloroplasts within each cell are together predictive
sheath,
PCK
PCK anatomy (Columbus Peterson
Aegopogon likewise has 1996).
Prendergast 1987).
of et
(Fig. 2; al.
PCK-like anatomy,
M. Muhlenbergia possess
and Herrera Arrieta (2001) reported that the species in subgen.
NAD-ME
whereas the remaining species have anatomy. However, M. tarahumara differs from species of Ae-
narrow around
gopogon and M. subgen. Muhlenbergia by having mesophyll chlorenchyma that forms a layer
by columns
sheath and not continuous between adjacent vascular bundles, being separated of
the outer
is
&
M. and M. (Peterson Herrera Arrieta
This pattern partially seen in pauciflora polycaulis
colorless is
cells.
columns
where vascular bundles near the middle of the blade have of colorless cells that separate
the
2001),
M.
bundles
adjacent vascular bundles. However, near the leaf blade margins the tertiary vascular of pauciflora
and M. have chlorenchyma that are continuous between each adjacent vascular bundle. In
cells
polycaulis
sclerenchyma
M. tarahumara, has well developed abaxial girders that
Muhlenbergia
addition, like
pauciflora,
&
Herrera
broadest near the epidermis (Peterson Arrieta 2001).
are
We number
and 8087-06)
Committee Research Exploration (grant
thank the National Geographic Society for
Endowments
Fund,
and Smithsonian Restricted the Scholarly
laboratory support; the Institutions,
for field
and Surveys and Inven-
Atherton Foundation, Biodiversity
Program, Research Opportunities, Seidell
Studies
new
Acevedo
Pedro
Program, financial support; Alice R. Tangerini for illustrating the species;
tories all for
and Lopez Spanish resumen; Adolfo Espejo Serna and Yolanda Herrera Arrieta for
Ida C. for correcting the
who
number with
providing helpful comments on the manuscript; and a of colleagues assisted fieldwork.
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