Table Of ContentAmerican Fern 97(3):155-165
Journal (2007)
Holub
Diphasiastrum multispicatum Wilce)
(J.H.
Thailand
(Lycopodiaceae)
in
H. WlLFRIED BENNERT
Bochum, D-44801 Bochum, Germany
Spezielle Botanik, Ruhr-Universitat Universitatsstrasse 150,
SUKSATHAN
PlYAKASET
Queen Botanic Garden, P.O. Box Mae Rim, Chiang Mai 50180, Thailand
Sirikit
7,
Horn
Karsten
Germany
Frankenstrasse D-91077 Dormitz,
2,
—
Review herbarium specimen (QBG) and subsequent studies have revealed that
Abstract. of field
a
Diphasiastrum multispicatum occurs near the summit of the two highest mountains in Thailand.
submontane
This species restricted to SE Asia and grows exclusively at higher elevations in the to
is
Taiwan, and Vietnam,
montane zone. Previously, has been reported from China, the Philippines,
it
A
but not from Thailand. comparison of the plants from Thailand with those from the type locality of
Thomas
D. multispicatum in the Philippines (Mt. Santo near Baguio City, Province of Benguet,
Luzon), where species present, shows the plants from Thailand to be D. multispicatum.
this is still
Diphasiastrum multispicatum weak competitor and grows on (disturbed)
Like other species, D. a
is
We
immature on slopes with more or open and low growing vegetation. also present
soils less
morphological evidence Diphasiastrum multispicatum distinct from Diphasiastrum
that is
and
complanatum which a north temperate, circumboreal species (in northern central
s.s., is
Europe, Greenland, northern North America, Japan and northern Asia, excluding the tropics).
The genus Diphasiastrum comprises a relatively small group of lycopods
Lycopodium
morphologically from the genus in several
that differ s.s, traits,
such leaves (mostly) 4-5-ranked (and not spirally arranged), leaves (mostly)
as
and
even trimorphic (and not isomorphic), upright shoots (mostly)
or
di-
was
quadrate flattened (and not rounded). This group formerly treated as
to
section Complanata of genus Lycopodium (Wilce, 1965; 011gaard, 1987, 1989,
and
Holub proposed genus Diphasiastrum,
1990), but (1975a) as a separate
it
many modern
treatment has been accepted in floras Dostal, 1984;
this (e.g.,
The
Wagner and Kukkonen, genus includes
Beitel, 1993; Jermy, 1993; 2000).
Only
and
about 25 species of mainly north temperate subarctic distribution.
limited number of species occur in the tropics or subtropics, where they are
a
mountainous Species with southeast Asian distribution
restricted to areas. a
include Diphasiastrum angustiramosum (Alderw.) Holub, D. multispicatum
and
(Wilce) Holub, D. piatyrhizoma (Wilce) Holub, D. veitchii (Christ) Holub,
& Holub
D. wightianum (Wall, ex Hook. Grev.) (Wilce, 1961, 1965).
and were
Additionally, taxa of unresolved relationship occur in this area
"New
called the "Chinese plant" and the Guinea plant" in the monograph by
Wilce The "Chinese plant" might be conspecific with the plants
(1965).
yueshanense M. Kuo and wilceae Ivanenko (Kuo, 1985;
described as D. C. D.
Ivanenko, To our knowledge, no recent comparative studies are
2003).
available for these critical taxa.
AMERICAN VOLUME NUMHKR
KKRN JOURNAL:
L56 97
3 (2007)
the century, Diphasiastrum was unknown Thailand (Tagawa
In last still in
and was name
Iwatsuki, 1979). Later, a single species reported under the
Lycopodium complanatum
(Boonkerd and Pollawatn, However,
L. 2000).
=
Diphasiastrum Lycopodium) complcmatum
north temperate, circum-
a
is
[
which
and
boreal species occurs in northern central Europe, Greenland,
northern North America and extending Japan and northern Asia. Wilce
to
complanatum
(1905, 103) states explicitly: ''There no true either in the
p. is L.
We
tropics (even on mountain southern hemisphere".
slopes), or in the present
evidence Diphasiastrum multispicatum Holub
that (Wilce) occurs Northern
in
where
Thailand, occupies the peaks of the two highest mountains.
it
Matkkials and Methods
Q
QBG)
September 2005 and February
P.S.) in in 2006. Field collection of plant
specimens on Doi Inthanon were made same months W.B. and
in the (by P.S.)
and on Doi Pha Horn Pok October
(by P.S.) in 2005. Additionally,
Diphasiastrum specimens
kept in the herbaria of Berlin and Aarhus
(B)
(AALJ) were annotated by K.H. The type Diphasiastrum
locality of multi-
spicatum on Thomas
Mt. Santo (Luzon, Philippines) was
revisited
in
November A
2005 (by P.S.) and February 2006 (by W.B.). Global Positioning
System (GPS) instrument (Garni GPS was used determine and
in 72) to altitude
coordinates of the Philippine and Doi Inthanon populations; Doi Pha Horn
for
SEM
were
Pok, these data obtained from topographical map. images
a of spores
made
were (DSM
using a scanning electron microscope 950, Zeiss).
Results and
Discussion
—
A
Morphology. herbarium specimen was "Lycopodium
that labeled
as
complanatum"
on peak mountain
(collected the of the highest in Thailand,
QBG
Nanakorn
Doi Inthanon, was
no. 6601) discovered while studying
s.n.
Queen
pteridophytes in the herbarium of Botanic Garden (Mae Rim,
Sirikit
Chiang The
Mai, Thailand; QBG). morphology
plant clearly different in
is its
from Diphasiastrum complanatum Lycopodium complanatum
(=
the type
L.;
specimen unknown
of species Wilce and Holub
this (see 1995, 143
is
p.
IT.
and was
1975b)) identified as Diphasiastrum multispicatum. The and the
site
population on Doi Inthanon were subsequently studied on several
field
trips.
Later, another population was discovered on second mountain
the highest
in
Thailand, Doi Pha Horn Pok
(by P.S.). Plants from the type locality on Mt.
Thomas
Santo (Luzon, Philippines) were studied comparison.
for
Table summarizes
diagnostic characters that distinguish D. complanatun
1
/
from D. multispicatum. The latter the more robust species, which becomes
is
evident especially from the dimensions of the rhizomes and peduncles.
Its
most
distinguishing characters are the long branchlets with very glaucous
a
lower and
side strongly incurving moderately well-developed
lateral leaves,
BENNERT DIPHASIASTRUM MULTISPICATUM THAILAND
ET IN 157
AL.:
Table Morphological comparison of Diphasiastrum complanatum and Diphasiastrum
1.
&
multispicatum (mainly after Wilce, 1961 1965).
complanatum Diphasiastrum multispicatum
Diphasiastrum
Characters
Characters being different
mm
somewhat
rhizome terete, 1.7 (1.1-2.7) in terete to flattened,
mm
broad and
diameter 2.2 (1.3-3.2)
mm
1.7 (1.3-2.3) thick
color of lower side pale, lighter in color than strongly glaucous
upper
surface, but not
glaucous
mm mm
(1.2-2.5)
ventral leaves (free blades) 1.3 (0.7-2.1) 1.8
mm
mm
leaves (including 5.0 (2.6-7.3) 4.6 (3.2-6.2)
lateral
bases)
mm
mm
dorsal leaves (including 4.8 (2.8-7.0) 3.8 (2.8-5.2)
bases)
mm
mm
diameter of peduncles 0.6 (0.4-0.9) 1.0 (0.8-1.1)
numerous,
number of strobili strobili few, 3-4, strobili generally 8
more, though occasionally
occasionally 5 or 6 per or
peduncle as few as 4 per peduncle
common
Characters branch of upright shoots distinctly flattened, leaves of
in lateral lets
ultimate branchlets 4-ranked, scale-like, trimorphic, ventral leaves
pedunculate
of branchlet less conspicuous than dorsal ones, strobili
sharply defined
lower leaves with evident decurrent leaf bases (Fig. a
1),
number
between branch and peduncle, and the of strobili per
transition fertile
complanatum
peduncle being significantly greater than in D. (Wilce, 1961,
number
Not surprisingly, the two species share a of traits that are typical
1965).
genus Diphasiastrum, branchlets of upright shoots flattened,
the like lateral
for
and
trimorphic (Table
leaves of ultimate branchlets 4-ranked, scale-like 1).
and
shows photographs of the habitat, part of the population
Fig. 2
mor
almost silvery lower surface are good field characters to identify the
bright,
Diphasiastrum multispicatum. At the type locality on Mt. Santo
plants
as
Thomas (near Baguio City, Province of Benguet, Luzon, Philippines), the
was and populations
Diphasiastrum multispicatum confirmed, 4
occurrence
of
The growth and
were found (Table habit the
different altitudes
at 2).
om
microm
Lycopodiaceae the spores are and subglobose to subtriangular in outline
trilete
show
Lycopodium
Those very
(Tryon and Lugardon, 1991). of s.L a
mposed
elements
of reticulate
mb such
like pattern. Wilce (1972) recognized 4 types of
)d
common
with most clavatum type being present in
spores the
reticulate
Lycopodium [Lycopodium and Complanata [Diphasias-
section
section
s.s.)
may
on be
trum). While the reticulum always continuous the distal face,
is it
(W Tryon and
proximal 1972;
face
AMERICAN FERN VOLUME NUMBER
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(2007)
3
mm
5
o
Fie. I. Line drawing of Diphasiastrum complanatum (above) and D. multispicatum (below)
showing
branchlets dorsal and view
in ventral
(left) (right).
Lugardon,
Diphasiastrum
1991). In multispicatum,
the outer one-third of the
proximal meshwork
face reticulate, but the gradually reduced towards
is the
is
aperture and lacking between
is the triradiate arms. This in contrast to D.
is
complanatum
and European
other Diphasiastrum where
species, the
re-
THAILAND 159
MULTISPICATUM
DIPHASIASTRUM IN
BENNERT ET
AL.:
Thomas
(Luzon,
on Mt. Santo
Philippines (a-d, h-i)
Diphasiastrum multispicatum in the
Fig. 2.
growth
open along roadside
Inthanon habitats (a, e).
and Thailand on Uoi (o-g);
type locality), in
rhizome
branches part of colony with strobili (d),
form of upright shoots (b, f], lower side of (c, g),
growth over blank soil on steep roadside slope (h-i).
160 AMERICAN
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Tabu;
Geographical
2. position of the two Diphasiastrum
multispicatum
localities in Thailand
and
of the type
locality in the Philippines.
Country
Altitude
(in asl) Coordinates
Thailand
Doi Inthanon
2,243 N
18 33.754' E 98 29.046
Doi Pha Horn Pok
N
1,950
20 06' E 99 07'
pp
Phili
IK'S
I I
Mt. Santo Thomas
population
1 2,260 N
16 20.115' E 120 33.651'
population
N
2
2,215
16 20.203' E 120 33.592'
population
3 2,196 N
16 20.204' E 120 33.585'
population
4
2,003 N
16 20.546' E 120 33.444'
ticulum
well-developed
on
is also the proximal and
face extends
the base
to of
the raised arms
of the aperture
(Fig. see also
Ferrarini Another
3; et 1986).
al.,
distinguishing
feature of D. multispicatum
the frequent occurrence
is of
perforations
the walls
in of the reticulum, on
visible both
faces
(Fig.
2i-j).
Diphasiastrum
remarkable
is in North America and Europe
for ability
its to
form homoploid,
apparently
fertile interspecific hybrids
(Flora of North
America,
1993; Sloor which
et al, 1996), are morphologically
intermediate
between
the putative Kuo
parents.
(1985) describes Lycopodium
yueshanense
Diphasiastrum
(- yueshanense) new
endemic
as a species from Taiwan and
reports
that intermediate between
is
it
L. veitchii D. and
veitchii)
L.
(
multispicatum
D. multispicatum)
both morphologically and
( ecologically.
Thus, yueshanense may
D.
well
represent the example homoploid
of
first a
hybrid
in the
tropics.
and
Ecology
Distribution.—
Diphasiastrum
multispicatum weak
com-
a
is
and
petitor colonizes
(disturbed) immature
on
soils slope
cuttings have
that
been man
created by with more
or open and low
less growing
vegetation.
All
checked
sites in Thailand and
in the Philippines
are located on
steep road
cuts
most
(Fig. In two
2). cases, other lycopods, Lycopodium
clavatum
and
L.
Lycopodiella
cernua
were
(L.) Pic. Serin., observed
the
in
vicinity.
other
In continents Diphasiastrum
species often grow on secondary
sites as well,
both
and
in the tropics
in the temperate
zones. Diphasiastrum
thyoides (Willdj
Holub
from South and
Central America,
example,
for reported from way- and
is
roadsides, clearings, scrub and
fallow land
(011gaard, 1988, Also
1995).
D.fawcettii
Lloyd
& Underw.) Holub
(F. L. from Jamaica and known
Hispaniola
occur on
is to
clearings, sunny embankments
and open
slopes with
scattered shrubs
(Proctor,
most
1985). In parts of North American
their and
Central European
range, the
Diphasiastrum
species display
a preference younger
for secondary on
habitats
immature
soils with an only fragmentary
plant Examples
cover.
are roadsides,
slopes
of forest roads, other
cuttings, runs and
ski their margins, abandoned
pits,
firebreaks along
railways,
forest aisles, clearings under power and
younger
lines
Ardelmann
afforestations
(e.g., 1995; Horn,
et al., 1997; Bennert,
1999).
THAILAND 161
MULTISPICATUM
DIPHASIASTRUM IN
BENNERT ET
AL.:
and
Doi Inthanon)
(Thailand,
multispicatum
SEM Diphasiastrum
photographs of spores of
Fig 3
column) and proximal face (right
showing
European Diphasiastrum species (a-h) distal (left
complanatum, e-f: D. issleri, g-h: D. zeillen, i-j:
alpinum, c-d: D.
column); a-b: D.
multispicatum.
D.
AMERICAN
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3
The
altitudinal range of Diphasiastrum
multispicatum
observed was
2,003
m ™A
on Thomas
260 Mt. Santo tVm Pliilinninoc t-v.^1,^^
in ;„
r^r-^
11,950-
m
on
2,243 Doi Inthanon and Doi Pha Horn Pok
(Table thus Diphasiastrum
2);
multispicatum
montane
represents
a species both
in countries.
Doi Inthanon
National km
Park, approximately
80
south Chiang
of Mai,
encompasses
the highest mountain
in Thailand, Doi
(Mt.) Inthanon, as well as
several summits. The
lesser park
covers an
area of 48,240 lowland
ha.
areas
Its
800 m)
(< are covered by dry
a dipterocarp which
forest gives an
rise to
evergreen
forest (above 1,000 m), where annual mm.
exceeds
rainfall 2,500
The
slopes around summit
the
area (2,300-2,565 m)
carry moist
a
hill
evergreen many
forest ('cloud forest') with The
epiphytes. temperature maj
-8
drop C
and
to
frosts are not unusual during
the dry season. Doi Pha Horn
Pok
approximately
lies 80
kilometers north Chiang Mai
of
second
the
is
highest mountain
Thailand
in
(2,285 m). Vegetation
types are almost the same
on
as Doi Inthanon.
Thomas
Mt. Santo
above
Bagnio
lies City (approximately 250
kilometers
north
of Manila)
in the heart of the Province
of Benguet. Baguio
City
itself
is
m
situated an
at elevation of about
1,400-1,500 and
has an
average annual
temperature C and mm
of 18.2 an annual
precipitation of 4,179
(recording
period
17 years; see: www.globalbioclimatics.org
climate diagram
- of Baguio
we
assume
City).
the temperature
If drop by C when
to
ca. 0.5 the elevation
increases by 100 m, an average annual
temperature C
of about 14 would
result
for the peak area of Mt. Santo Thomas. Annual
precipitation expected
be
is to
somewhat
higher than in the The known
city. presently
area of distribution
in
Thailand
comprises
only two and few
sites, additional
localities are be
to
expected,
as there number
are limited
a of higher mountains
with an
elevation
exceeding
2,000 m. Other
countries SE
in Asia from which
Diphasiastrum
multispicatum
has been
reported China
are
(southern
part), the Philippines.
Taiwan, and
Vietnam.
the
In recent treatment
of Lycopodiaceae
in the Flora
of
China
(Xianchun and
Libing,
2004), Diphasiastrum
multispicatum
not
is
recognized
as separate
a species, but united with
Diphasiastrum
complaua-
The
tum.
latter has been two
split into
varieties, the complanatum
typical
var.
and
glaucum. From
var.
the short diagnosis given by Ching
(1982)
in his
description glaucum
of and
var. from
the appearance
of the type specimen
(a
photo which was
of
supplied by we
Y. Ivanenko), conclude
that this variety
Diphasiastrum
refers to
multispicatum.
This
already has been proposed
bv
Ivanenko The
(2003).
total altitudinal
distribution
in these countries ranges
m
m
from
1,165 to 2,415 corresponding
submontane
to the montane
to /ones
(W
ig65
Tab]o summarizos
3 tho
rocor( oc Thus,
j localities. the
]
j
discovery
of Diphasiastrum
multispicatum
Thailand
in remarkable
a range
is
extension, and the mountains
in northern Thailand harbor
the westernmost
known
populations
of
this species. Diphasiastrum
multispicatum
likely
is to
occur
in the neighboring country
of Laos where
as well,
several
sufficiently
high mountains
are located with
the mountain, Phou
tallest
Bia
in the
Annamese m
Cordillera,
attaining 2,817 above
sea
level.
by B [Botanischer z
compiled from and obtained from herbarium revision (indicated
Diphasiastrum multispicatum; data literature
Records
Table 3 for z
AAU The given the Chinese literature
Aarhus, Denmark]). distribution in
Museum Germany] [University of
und Botanisches Berlin-Dahlem, or
Garten
synonym Diphasiastrum multispicatum. H
which
complanatum glaucum of
Diphasiastrum a
var. is
refers to
H
Reference
Locality >
Region/province/district
—
State
Xianchun & 2004
Libing,
Guangxi * *
China Prov.
Yunnan
Prov.
Xizang
Prov. (Tibet) 1
1982
Ching,
Yunnan Pingbian Xian
Eastern CO
DeVol & Kuo, 1979
Taipingshan
Taiwan Co
Distr. Ilan
DeVol & Kuo, 1979
Tapachienshan
Hsinchu
Distr.
DeVol & Kuo, 1979
Kuantaochi
Nantou
Distr.
DeVol & Kuo, 1979
Alishan
Mt.
Chiayi
Distr.
CM.
Kuo
L
M.T. Kao (7486), 13.12.1968, det. as
AAU
Horn
complanatum,
rev. K. (2003),
DeVol & Kuo, 1979
CO
Kuanshanakou
Kaohsiung
Distr.
DeVol & Kuo, 1979
Mukuashan
Hualien
Distr.
2
Wilce, 1961
Mt. Taiheizan H
c
1961
Wilce,
Arisan
paper
Mai Doi Inthanon this Z
Chiang
Thailand Prov.
paper
Pha Horn Pok this
Mai Doi
Chiang
Prov.
=
>
Nam-kep, Massif du Pia-Quac Wilce, 1961
Vietnam Bac Phan (Tonkin)
North Prov.
Chapu, Lo Qui Ho between Cao-Bang Wilce, 1961 >
Z
and Nguyen-Binh
complanatum,
Col de Lo Qui Ho, pres Chapa: Cha-pa E. Poilane (12641), 13.5.1927, as L.
AAU
Horn ex
Cho-bo rev. K. (2003), P,
et
(VH
Averyanov 183), 17.3.1995, as L.
Kontum no exact locality given et al.
Vietnam
South Prov.
AAU
Horn
complanatum,
rev. K. (2003),
complana-
Hb. E.B. Copeland, 25.10.1905, as L.
Tabiao
Benguet Mt.
Philippines Luzon, Prov.
Horn B
tum
var. thyoides, rev. K. (2005),
1965
Thomas; Baguio Wilce; 1961,
Luzon, Prov Benguet Mt. Santo
1965
Wilce; 1961,
Tonglon
Nueva Viscava Mt.
Luzon, Prov.
(?)
1965
Banahao Wilce; 1961,
Maquiling; Mt.
Luzon, Prov Laguna Mt.
104 AMERICAN
FERN VOLUME
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3
Acknowledgments
We
thank
Mrs. Wessel, Bochum,
tlso for preparing the drawings complanatum
of D. and
D.
multispicatum, and Mr. Marcus
Strockonbach, Bochum, SEM
providing
for the photos
of the
spores
of D. multispicatum. Also, thanks due
are
to Prof. Dr. Brigitte Zimmer, and
Berlin, Prof. Dr.
Benjamin
Ollgaard, Aarhns, sending herbarium
for specimens on
loan (from the herbaria B and
AAU).
Ynry
Dr. Ivanenko, Petersburg
St. (Russia), sent us a photograph of the type specimen
of D.
complanatum
glaucum PE
var. kept
in (Institute of Botany. Chinese Academy
of Sciences,
Beijing),
and
Prol. Dr. Zhixiang Zhang,
Beijing (China) contributed information on
the occurrence
of
D.
multispicatum
China;
in both are gratefully acknowledged.
Wrightwood USA)
Jeff Stauffer, (CA,
and
Dr. Nicola Bennert, USA)
Riverside; (CA,
contributed improving our
to English.
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