Table Of ContentAND
FESTUCA ALOHA M0L0KA1ENS1S (POACEAE: LOLIINAE),
F.
TWO NEW FROM
HAWAFI
SPECIES
Robert Soreng and Paul M. Peterson
Catalan
Pilar J.
While revising specimens of Festuca deposited at the United States National Herbarium (US) the first author
However,
found two specimens fme-leaved individuals from Kaua'i that were identified as rubra L.
of F.
F
these individuals exhibit several characters that separate them from holarctic red fescues of the rubra
and
complex, including: leaf blades flat with involute margins, sheaths open, relatively long ligules, ovaries
specimens from Molokai'i deposited the Bishop
with densely hairy apices. After reviewing of Festuca at
Museum (BISH) we noticed an additional taxon. Previously, the only species of Festuca and relatives re-
(Hitchcock robust
corded from Hawai'i and the Pacific include the endemic, F. hawaiiensis Hitchc. 1922), a
&
broad-leaved species placed in subg. Drymanthele V.I. Krecz. Bobrov sect. Banksia E.B. Alexeev (Alexeev
F.
and
Aulaxyper Dumort.);
1980), and three Eurasian species: the fine-leaved F. rubra L. (F subg. Festuca sect.
the broad-leaved F. arundinacea Schreb. and F. pratensis Huds. [F subg. Schedonorus (P. Beauv.) Peterm. sect.
Schedonorus Beauv.) Endl]. The taxonomy of Festuca si, the largest genus of monophyletic subtribe
(P.
Loliinae Dumort., is in a state of flux. Much additional research is needed to properly align the taxa, although
DNA
two major
preliminary data from sequencing (Catalan et al. 2007; Inda et al. 2008) suggest that clades
may two have been within the
As elements of the "broad-leaved" clade, the latter species classified
exist.
Dumort
separate genus Schedonorus Beauv., as Schedonorus arundinaceus (Schreb.) and S. pratensis (Huds.)
P.
Snow Drymanthele sometimes recognized
Beauv. (Soreng 2003; 2008). In addition, Festuca subg. is
et
P. al.
F
as the genus Drymochloa Holub. The "fine-leaved" clade of fescues, including rubra, are placed in Festuca
However, equivocal concerning the resolution of the division at the
(Soreng 2003). results are
et still
s.s. al.
two major
base of the clades.
Specimens were examined from the Bishop Museum (BISH), National Tropical Botanic Garden (PTBG),
Herbarium
Missouri Botanical Garden (MO), and the United States National (US).
&
nov
Festuca aloha Catalan, Soreng P.M. Peterson, sp. (Figs. 1A-J, 2). Type: U.S.A. Hawaii: kaua'i, Hanalei district,
isolated hanging side-valley of lowland diverse mesic forest
t,
fwt^
&
Festuca aloha Catalan, Soreng P.M. Peterson (isotype, US-3252239). A. Habit. B. Sheath, ligule, and blade. C. Spikelet. D. Floret
f
with rachilla. E. Lemma, ventral view. F. Palea, dorsal view. G. Palea with ovary, ventral view. H. Palea with immature ovary and mature stamens,
ventral view. Lodicules, mature ovary, and basal remnant of filaments.
I. J. L<
BISH-728771).K. Spikelet.
Culms
Plants perennial, cespitose, with extravaginal innovations; cataphylls conspicuous, brownish.
mm
cm
45-70 slender diam.), nodes 2-3, smooth and glabrous. Leaf sheath margins fused
tall, erect, (ca. 1
mm
when
for 1-2 at base, overlapping more than Mi the length below, sparsely villose abaxially young,
becoming glabrous maturity, purple-brownish the base, greenish above, becoming fibrous in age
at at at
mm
base; auricles absent; collars glabrous; ligules 1-1.5 long, scarious, glabrous, brownish, apex obtuse,
cm mm,
dentate-erose; leaf blades of vegetative shoots 22-33 x (1-)1. 5-1.7 erect to pendant, flat with
and smooth up
involute margins, glabrous abaxially, hirsute along protruding ribs adaxially, the hairs to
mm mm
0.1 long, cross section (Fig. 2) about 1 bearing 13-16(-20) larger vascular traces or veins, each
with prominent ribs adaxially alternating with smaller ribs on the secondary and tertiary veins, scleren-
chyma cm mm,
strands forming trabeculae or girders on most veins; leaf blades of culms about 22 x 2.4
erect to pendant. Panicles 8-11 x 2.5-4 cm, erect, axis and peduncle smooth; loosely contracted with
34-66 4-6 cm
spikelets, spikelets mostly at branch tips; panicle branches 2(3) per node, basal branches
and smooth
ca. 1/2 the length of panicle, angled, erect or ascending, proximally to scabrous, angled dis-
mm
mm,
Spikelets 8.2-13 x 3.5-6.0 4-6-flowered, broadly lanceolate, pale green, sometimes tinged
tally.
mm mm
glumes
with purple; pedicels (l-)1.5-3.5(-5) long; rachilla internodes 0.8-1.2 long, scabrous;
lanceolate, green, margins very narrowly scarious (<0.5 mm), apex acute; lower glumes (3.6-)3.9-4.2(-4.5)
mm mm mm
lemmas
upper glumes
long, 1-veined; 5.2-5. 5(-6.0) long, 3-veined; 5.8-8.0 long, 5-veined,
glabrous below, scabrous near the apex, green or tinged with purple at the apex, mucronate to awned, the
mm
awn
mucro smooth
or 0.5-1.1 long, scabrous; callus rounded, obliquely angled 45°), to dorsally
(to
minutely scabrous at the apex; paleas longer or shorter than lemma, scabrous on and between keels; stamens
mm
anthers 3.1-4.2 long; ovary densely hairy on upper 1/3. Caryopsis not seen.
3,
abilis, Carex meyenti, ca. 3000 17 Jun 1994, K.R Wood 3245 (PTBG-20438); Pohakuao, hanging valley between Kalalau and Hanakoa,
ft,
below Puu Ki and Kaaalahine Ridge, 2000 4 Jan 1992, K.R. Wood 1761, Perlman Lou (BISH-621 133, PTBG-12875, US-3250261);
ft, S. &J.
&
Waimea district, with Panicum lineale and P. beecheyi, 2000 ft, 30 Mar 1993, K.R. Wood 2470, S. Perlman (PTBG-18788), K.R. Wood 2471
&
N
Perlman (PTBG-18779); Awaawapuhi facing slopes above stream, 0.5 mi along growin
S. valley, trail,
—
cm
Comments. Festuca aloha has panicles 8-11 long, narrow, branches erect and ascending; lemmas
mm
mm
cm
30-40
5.8-8 long; and leaf blades 1.5-2.4 wide; in contrast hawaiiensis has panicles long,
F.
mm mm
lemmas and 2-3
widely open, branches patent or patent-erect; 9 long; leaf blades wide.
The illustrator Alice Tangerini noticed that all florets with mature anthers of F. aloha contained
small, undeveloped ovaries (Fig. 1H). Mature ovaries with well-developed styles were found only in florets
that had already shed their anthers (filaments were still present) [Fig. II]. Therefore, F. aloha is apparently
protandrous.
—
name
Etymology. The specific epithet aloha derives from the Kalalau mountains of Kaua'i.
F.
—
Conservation status. Festuca aloha has been found in at least six different localities of Kauaii, covering
km
a distribution area of approximately 700 2 The number of individuals varies among populations but
.
new
usually there are less than 1000 individuals per population. Based on these preliminary data, the spe-
IUCN
cies clearly falls within Vulnerable (VU) category as defined by the (2001). The major threats to in-
F
dividuals of aloha are the allochthonous plants, such Cyperus meyenianus Kunth, Erigeron Kalan-
as: sp.,
choe pinnata (Lam.) Pers., Lantana camara L., Melia sp., Passiflora mollissima (Kunth) L.H. Bailey, Ruhus
=
&
le cross section of vegetative shoots of Festuca aloha Catalan, Soreng P.M. Peterson (US-3252239). Scale bar 0.1 it
Culms
up
Dwnish.
Plants perennial, cespitose,
cm
60-70 scabrous below panicle. Lc base, overlapping
to
tall, it
more than lh the length below, purple-brov fibrous at base in
I
mish, apex obtuse,
age; auricles absent; collars glabrous; ligult
pendant,
dentate-erose; leaf blades of vegetative si r less flat
and smootl
with involute margins, glabrous
25
gins and apex, dull green, cross section sim iring larger vas-
each with prominent
cular traces or veins,
ri
bulliform between adaxial sclerenchyma strands present forming trabeculae or
tertiary veins, cells ribs,
and
on most sclerenchyma strands with thin walls (adaxially) thick cell walls (abaxially);
girders veins, cell
cm mm,
Panicles
leaf blades of culms about 21 x 2 with involute margins, densely scabrous adaxially.
flat
8.5-13 x 4 cm, loosely contracted with approximately 75 spikelets, spikelets located at the ends of the
branches, branches on basal node, 2(3) branches in other nodes, axis and branches strongly scabrous;
3(4)
branches long 7 cm, more than 1/2 the length of the panicle, angled, suberect to ascending.
basal as as
mm
mm,
Spikelets 7-9 x 3-4 5-6-flowered, broadly lanceolate, pale green; pedicels (2-) 4 (-9) long; ra-
internodes scabrous; glumes lanceolate, green, margins very narrow, scarious, apex acute; lower
chilla
mm
mm
glumes upper glumes (6.0-)6.2-6.8(-7.0) long, 3-veined;
(5.0-)5.2-5.5(-5.7) long, 1-veined;
mm
mm
awn
lemmas (5.0-)5. 5-6.0 long, 5-veined, scabrous dorsally and near apex, green, 1.5-2.5 long,
scabrous; callus rounded, obliquely angled (to 45°), scabrous; paleas slightly shorter than lemma, bidentate
mm
at apex, keels scabrous, between keels scabrous; stamens 3, anthers 2.8-3.0 long; ovary densely hairy
on upper Caryopsis not
seen.
1/3.
—
Comments. Festuca molokaiensis differs from F aloha by having more delicate and thinner, more nu-
merous and lax branches; scabrous panicles and spikelets; lemmas scabrous throughout, awns longer
mm; mm;
1.5-2.8 mm; anthers shorter 2-3 lower glumes (5.0-)5.2-5.5(-5.7) and upper glumes (6.0-)6.2-
cm
mm. In comparison to F aloha, culms are scabrous on the nodes; panicles 8.5-13 long, more
6.8C-7.0)
more
and peduncles scabrous, branches densely scabrous, angled; spikelets delicate; rachilla
delicate, axis
mm
mm
more slender; lower glumes (5.0-)5.2-5.5(-5.7) long; upper glumes (6.0-)6.2-6.8(-7.0) long; lem-
mm
mas 5-6 long, evenly scabrous across the back from the base to the apex, 3-veined or infrequently
mm mm
awn and 2-3
5-veined veins obsolete or obscure), awned, the 1.5-2.8 long; anthers
faintly (lateral
long.
Two new
Catalan et species of Festuca from Hawaii 55
al.,
—
F
The from
Etymology. specific epithet of molokaiensis derives the island of Moloka'i.
—
known
Conservation status. Festuca molokaiensis is only from the type locality where it has been re-
Due
corded as occasional (Oppenheimer, on herbarium label). to the scarcity of available data on the po-
F
tential distribution, demography, and population status of molokaiensis, we consider to be a Data Deficient
(IUCN
(DD) taxon
2001).
TAXONOMY AND PHYLOGENY
F
F
Festuca aloha and molokaiensis art morphologically similar to the Hawaiian endemic, hawaiiensis in ligule
size and shape, ovary hairiness, and leaf-blade anatomy (Hitchcock 1922). However, hawaiiensis a more
F. is
cm
robust plant up to 150 tall with longer (30-40 cm) and wider panicles that are open with 3-5 spreading
mm
and drooping branches at the nodes, and larger lemmas up to 9 long. Festuca hawaiiensis grows at
higher elevations, at ca. 2000 m, in rich soil on moist wooded hills on the island of Hawaii.
F F
Festuca aloha and molokaiensis resemble the western North American, calijornica ssp. hitchcockiana
&
and
(E.B. Alexeev) Darbysh. [Darbyshire Pavlick 2007] in leaf blade cross section, panicle features, hairy
F new
ovary apices. However, calijornica ssp. hitchcockiana differs from the species by having a more robust
and cespitose habit, possession of a continuous sclerenchyma layer on the abaxial side of the leaf blade, a
and with
partially hairy collar, longer ligules ciliate apices.
F
The new Festuca taxa also show similarities with Macaronesian volcanic cliff dwellers: agustinii Lind-
&
ing., F.jubata Lowe, and F.francoi Fern.Prieto, C.Aguiar, E. Dias M.I. Gut. (Saint-Yves 1922; Fernandez
and
Prieto et al. 2008) because all share extravaginal innovations flat to inrolled (or conduplicate) leaf-blades
with some complete sclerenchyma trabeculae. However, the latter species all differ from the new species by
mm)
having glabrous ovary apices and short (< 1.5 ligules.
We
analyzed the ITS and trnL-F sequences of two samples of Festuca aloha from different populations
from Kauai US-3252239 GenBank GQ162205 and GQ162208
[F aloha Kalalau, (isotype), (ITS) (trnLF)
1:
accessions; F aloha Pohakuao, US-3250261, GenBank GQ162206 (ITS) and GQ162209 (trnLF) accessions]
2:
F known
and one sample of molokaiensis from the only population from Molokai [F molokaiensis Kupai
1:
We
Gulch, BISH-728771 (holotype), GenBank GQ162207 (ITS) and GQ162210 (trnLF) accessions]. then
random
conducted heuristic parsimony analyses for the combined ITS and trnL-F dataset (10,000 entry
trees, TBR, mulpars off, saving no more than 10 trees of length <10 per replicate; 1000 bootstrap replicates
with the same parameters as in the original search) using these three samples and combined them with our
previous data for Festuca on a worldwide level (Catalan et al. 2007; Inda et al. 2008).
All three samples grouped together within the broad-leaved Festuca clade nested within the
(Fig.3),
F
well supported clade Subulatae + Leucopoa [92% bootstrap(BS)], which were separated from
p.p. calijor-
and Macaronesian Aulaxyper was monophyletic (99%
nica, F. rubra, the Festuca sect. s.l. species. Festuca aloha
F
BS) and sister to molokaiensis (98% BS).
&
F F
The two aloha specimens from different populations on Kauai (F aloha 1 aloha 2) had similar
F
sequences (99% BS; trnL-F nucleotide substitution difference); however, the sample of molokaiensis from
1
Molokai showed several nucleotide differences in both the ITS substitutions) and trnL-F (two substitu-
(five
tions) regions with respect to aloha. These molecular differences also support the taxonomic separation
F.
F
of the two species. Despite the limited infraspecific sampling, the ITS variation found between aloha and
F
molokaiensis, from the same Hawaiian archipelago, remarkable as these species show more nucleotide
is
F Ramond
differentiation than that found between other closely related species the Pyrenean eskia ex
[e.g.
F
DC., and gautieri (Hack.) K. Richt. with only two ITS nucleotide substitutions], and between species from
F
different Macaronesian archipelagos Madeiran F.jubata and Azorean petraea Guthn. ex Seub. with
(e.g.,
F
three ITS nucleotide substitutions). Unfortunately, hawaiiensis could not be included in the molecular
we
analysis because extremely rare and have no material other than the type.
it is
F F
The closest relative of F. aloha and molokaiensis in our analyses (Fig. 3) was the eastern Asian
F
parvigluma Steud. (88% BS), followed by the northwestern North American subulata Trin. (77% BS), both
F F
belonging to subg. Subulatae, and then by the Siberian-North American altaica Trin. (92% BS), placed
Journal of the Botanical Research Institute of Texas 3(1)
-Og:
^hlr
_Castellia_
"_^--^:-^-_L
,
-^
iff
x excluding uninformative characters 0.400;
within subg. Leucopoa sect. Breviaristatae. In a separate ITS
F. s
F
ensis clade fell within the broad-leaved clade, and was closely related to the Asian parvigluma and F.
modesta Steud. [sometimes treated as Drymochloa modesta (Nees ex Steud.) Holub]. In contrast to the ITS
F
data, a trnL-F strict consensus tree placed aloha/F. molokaiensis in a polytomy at the base of the fine-leaved
among
clade, fine-leaved taxa and taxa intermediate between fine- and broad-leaved forms of Festuca.
DNA
Given the different resolutions between separate nuclear and plastid sequence topologies for our
On
species, possible that aloha and molokaiensis are of allopolyploid origin. morphological grounds
it is F. F.
common
and same
aloha, molokaiensis, hawaiiensis art part of the complex, representing a pattern
F. F. F. all
of morphological speciation in upland habitats of different islands along the Hawaiian archipelago.
2. Auricles ciliate on margin; lemmas awned (awn 2-3 mm), more or less scabrous distally_
Auricles not on margin; lemmas muticous, smooth or only spars
2. ciliate
mm
All leaf blades without auricles; leaf blades 0.3-5 wide, involute to flat but whei
.
mm
Ligules 0.1-0.5 long; ovary apices glabrous; leaf blades conduplicate;
3. le
below (open < 14 their length); leaf blades of vegetative shoots in cross sectii
mm
Ligules 1-2.5 long; ovary apices densely hairy; leaf blades flat with involute margins;
l<
.
mm
generally open 16 their length [they are fused only for 1-2 at the base]; leaf blades ofvegeta
in cross section with sclerenchyma girders.
mm
cm
30-40 3-5
4. Panicles long, open, the branches spreading or drooping; leaf blades wi
iwaii
Festuca aloha
.
Lemmas mm;
awns
evenly scabrous throughout, intermediate veins absent or obscure, 1.5-2.8 anthers
5.
mm mm mm
2-3 long; iigules 1.5-2.5 iong; lower glumes (5.0-)5.2-5.5(-5.7) long; upper giumes
mm
known
(6.0-)6.2-6.8(-7.0) long; only from Moloka'i Festuca molokaiensis
ACKNOWLEDGMENTS
This work has been subsidized by the Spanish Ministry of Science and Technology Grant Project CGL2006-
We
00319/BOS and Sabbatical Research Stay Grant PR2008-0020 to Pilar Catalan. thank Stephen Darbyshire
and Neil Snow for their critical review of an earlier version of the manuscript; Napua Harbottle, Tim Flynn,
PTBG
and Gerrit Davidse, for facilitating for us the study of Hawaiian Festuca materials deposited at BISH,
and MO, respectively; Hank Oppenheimer, Neil Snow, and Cliff Morden for valuable information on the
F F
ecology and distribution of aloha and molokaiensis in Kauai'i and Molokai'i; Sasha Savytskyy, Dai
Tsuchiya, and Juan Viruel for helping us with figure preparation; Alain Touwaide for correcting the Latin
and
diagnoses; Alice R. Tangerini for providing the illustration.
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