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INTRODUCTION
Mosses are remarkably successful plants that thrive
alongside more conspicuous vascular plants. They
originated during the Upper Ordovician-Lower Silurian
border (~440 million years) together with the first land
plants (Shaw and Goffinet 2000). They are currently
represented by approximately 10,000 species worldwide.
*Corresponding author: udangya@hotmail.com
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Reproductive
Effort
Life strategy
Spores numerous, very light (<20 m)
fugitives
Annual shuttle
high
few
Colonists
high
many
Perennial stayers
low
few
Life strategy
Reproductive Effort
fugitives
Annual shuttle
Colonists
Medium
- Ephemeral Colonists
Long-lived shuttle
high
high
- Pioneers
Perennial stayers
many
- Competitive Perennial
Dominants
low
RESULTS
The moss collections from Mt. Pulag totaled 58 species
that belong to 28 families bringing overall inventory to
144 species, an increase of 27 species from the previous
count of 117 (from those documented by Bartram in 1939
and Tan and Iwatsuki in 1991). Table 3 summarizes the
collected specimens.
Life Strategy and Vegetation Type
Five out of the six life strategies of During (1979) have
been observed to occur in the moss populations of Mt.
Pulag. The only strategy that was not seen was the
short lived shuttle. As can be observed, the frequency
of colonists in all of the collection sites is quite high,
however lesser in mossy forest 1 (~2725 m). Fugitives
and annual shuttle species are observed to occur only in
the pine forest. The frequency of perennials and long
lived shuttle species vary between the different collection
sites. For perennials, the mossy forests are the habitats
that they prefer basing on the relatively high frequency of
occurrence of this strategy in the two collection sites. The
long lived shuttle dominate the grassland. Abundance of
the different life strategies are shown in Figure 1 and 2.
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80%
60%
40%
20%
0%
rt l
ive
d
lon
gl
ive
d
pe
ren
nia
l
un
kn
ow
n
sh
o
itiv
e
fug
nu
an
co
lo
al
ni s
t
% occurrence
100%
Life Strategy
long lived
unknown
perennial
100%
90%
80%
70%
60%
50%
40%
30%
20%
10%
0%
% occurrence
Figure 1. Abundance of each moss life strategy in each of the vegetation zones in %s.
short lived
fugitive
annual
colonist
Vegetation Type/Elevation
Figure 2. Abundance of life strategies from the different vegetation zones/elevation in %s.
Table 3. Moss species collected from Mt. Pulag, Benguet Province Philippines together with their family,
vegetation type of location, expressed life strategy, and substrate (During 1979)
Species
Family
Vegetation Type**
Life strategy
Anoectangium aestivum
Pottiaceae
pine
Colonist
Anomobryum gemmigerum
Bryaceae
pine
Colonist
Anomobryum gemmigerum
Bryaceae
pine
Colonist
Barbella flagellifera
Meteoriaceae
m1
Perennial
Bryum clavatum
Bryaceae
pine
Unknown
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Bryum ramosum
Bryaceae
pine
Colonist
Calyptrochaeta remotifolia
Hookeriaceae
m1
Calyptrochaeta remotifolia
Hookeriaceae
grassland
Campylopus austrosubulatus
Dicranaceae
grassland
Colonist
Campylopus austrosubulatus
Dicranaceae
m2
Colonist
Campylopus hemitrichius
Dicranaceae
grassland
Colonist
Campylopus hemitrichius
Dicranaceae
m1
Colonist
Campylopus hemitrichius
Dicranaceae
grassland
Colonist
Campylopus hemitrichius
Dicranaceae
pine
Colonist
Campylopus laxitextus
Dicranaceae
pine
Colonist
Campylopus laxitextus
Dicranaceae
grassland
Colonist
Campylopus sp.
Dicranaceae
pine
Colonist
Campylopus umbellatus
Dicranaceae
m2
Colonist
Campylopus umbellatus
Dicranaceae
m2
Colonist
Campylopus umbellatus
Dicranaceae
pine
Colonist
Clastobryum caudatum
Sematophyllaceae
pine
Ctenidium andoi
Hypnaceae
m2
Perennial
Ctenidium andoi
Hypnaceae
m1
Perennial
Ctenidium andoi
Hypnaceae
m1
Perennial
Ctenidium lychnites
Hypnaceae
m1
Perennial
Dicranodontium fleischerianum
Dicranaceae
m2
Colonist
Dicranodontium fleischerianum
Dicranaceae
m1
Colonist
Dicranoloma brevisetum
Dicranaceae
m1
Colonist
Dicranoloma brevisetum
Dicranaceae
m2
Colonist
Dicranoloma brevisetum
Dicranaceae
m1
Colonist
Dicranoloma reflexum
Dicranaceae
m2
Colonist
Dicranoloma sp (billarderi)
Dicranaceae
m1
Colonist
Dicranum psathyrum
Dicranaceae
pine
Colonist
Ditrichum dificile
Ditrichaceae
pine
Colonist
Ectropothecium falciforme
Hypnaceae
m2
Entodon sp.
Entodontaceae
pine
Perennial
Entosthodon buseanus
Funariaceae
pine
Annual
Eurhynchium asperisetum
Brachytheciaceae
m2
Perennial
Eurhynchium asperisetum
Brachytheciaceae
m2
Perennial
Fissidens nobilis
Fissidentaceae
m1
Colonist
Fissidens nobilis
Fissidentaceae
m2
Colonist
Fissidens plagiochiloides
Fissidentaceae
m2
unknown
Funaria hygrometrica
Funariaceae
pine
Fugitive
Gammiella ceylonensis
Sematophyllaceae
grassland
unknown
Gammiella ceylonensis
Sematophyllaceae
pine
unknown
Gollania benguetense
Hypnaceae
pine
unknown
Gollania benguetense
Hypnaceae
pine
unknown
Gollania benguetense
Hypnaceae
pine
unknown
Homaliodendron flabellatum
Neckeraceae
m2
Perennial
Hookeria acutifolia
Hookeriaceae
m1
Isothecium trichocladon
Lembophyllaceae
m2
Perennial
Bartramiaceae
m1
Leiomela javanica
see next page for continuation . . . .
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Leucobryaceae
pine
Perennial
Macromitrium reinwardtii
Orthotrichaceae
pine
Macromitrium reinwardtii
Orthotrichaceae
grassland
Macromitrium reinwardtii
Orthotrichaceae
m2
Macromitrium sulcatum
Orthotrichaceae
pine
Meteoriopsis reclinata
Meteoriaceae
pine
Miothecium microcarpum
Sematophyllaceae
pine
Mnium laevinerve
Mniaceae
m1
unknown
Mnium laevinerve
Mniaceae
m1
unknown
Papillaria fuscescens
Meteoriaceae
m2
Plagiomnium integrum
Mniaceae
m1
unknown
Plagiothecium neckeroideum
Plagiotheciaceae
m1
Perennial
Plagiothecium nemorale
Plagiotheciaceae
m1
Perennial
Platyhypnidium muelleri
Meteoriaceae
m1
Perennial
Pogonatum perichatiale
Polytrichaceae
pine
Colonist
Pogonatum proliferum
Polytrichaceae
m2
Colonist
Pogonatum proliferum
Polytrichaceae
m1
Colonist
Pogonatum urnigerum
Polytrichaceae
m2
Colonist
Pogonatum urnigerum
Polytrichaceae
m1
Colonist
Pohlia elongata
Bryaceae
pine
Colonist
Racomitrium subsecundum
Grimmiaceae
pine
Unknown
Racopilum johannis-winkleri
Racopilaceae
m1
Unknown
Racopilum johannis-winkleri
Racopilaceae
m1
Unknown
Rhodobryum giganteum
Bryaceae
m1
Colonist
Sphagnum cuspidatulum
Sphagnaceae
m1
Syrrhopodon tjibodensis
Calymperaceae
pine
Colonist
Taxiphyllum taxirameum
var. recurvifolium
Hypnaceae
m1
Perennial
Thamnobryum subserratum
Thamnobryaceae
m1
Perennial
Thuidium cymbifolium
Thuidiaceae
m1
Perennial
Trachyloma indicum
Pterobryaceae
m1
Perennial
Trachyloma indicum
Pterobryaceae
m1
Perennial
Trachypodopsis serrulata
Trachypodaceae
m1
Perennial
DISCUSSION
Overall affinities of specific life strategies to the different
altitudinal vegetation zones have been found. The
following discuss the observed moss based vegetation
pattern correlated with the dominant vegetation pattern
mentioned by Merill and Meritt (1910).
Grassland Moss Flora
The absence of a tree canopy that could protect against
intense light and high wind velocity makes the grassland
environment particularly desiccating for such organisms,
and may be the prime reason why the grassland is the
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CONCLUSION
Mt. Pulag bryoflora is one of the more diverse communities
of bryophytes especially of mosses. It is here that many
of the mosses and other organisms find their last refuge
in view of the continuous and large-scale land conversion
happening throughout the country. It is also where life
strategy distribution has never been investigated. In this
view, this research is pioneering. The study on these facets of
moss biology has resulted in the establishment of moss based
zonation pattern in Mt. Pulag conforming to the boundaries
of Merill and Merrits (1910) 3 distinct vegetation zones.
The following moss-life-strategy based zonation would be:
(1) Mixed strategy Zone where those with short life spans
are present (Fugitives and Annual Shuttle species) at the
pine forest, (2) Perennial Stayers and Long Lived Shuttle
Zone within the mossy forest, and (3) Colonists-Annual
Shuttle Species Zone at the grassland. The dominance of
these life strategies unique to each of the vegetation zones
illustrates the distinctly different environmental conditions
offered by each of these ecosystems to which the mosses
have to adapt to. It would be very interesting to determine
what these factors are that specifically favor the abundance
of specific strategies. In the words of During (1992), results
of these types of researches are a valuable guide in the further
investigation regarding environmental key factors that define
specific environments or regions.
ACKNOWLEDGEMENTS
We would like to thank Dr. Benito Tan of the National
University of Singapore for his invaluable help in the
identification of the moss collections, and Dr. Inocencio Buot
and Prof. Maria Fe Sangalang of UP Los Baos for the help in
the preparation of this research. Our gratitude also to the Tan
Kin Chee Foundation for a small grants award, SEARCASEAMEO for the thesis grant, and UP Baguio for the Local
Faculty Fellowship grant to the corresponding author. The
assistance of Mr. Orlando Apostol, Ronnel Almazan, and Rexel
Almazan during field work is also gratefully acknowledged.
REFERENCES
BARTRAM ED.1939. Mosses of the Philippines. The
Philippine Journal of Science 68(1-4).
BUOT IE and OKITSU S. 1998. Vertical distribution and
structure of the tree vegetation in the montane forest
of Mt. Pulog, Cordillera mountain range, the highest
mountain in Luzon Is., Philippines. Vegetation Science
15:19-32.
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EDDY A. 1996. A handbook of malesian mosses. Vol. 13. British museum of natural history. London: Natural
History Museum Publications
FLOWERS S.1973. Mosses: Utah and the west. Utah:
Brigham Young University Press. 567p.
FRAHM JP. 1990. Bryophyte phytomass in tropical
ecosystems. Botanical journal of the linnean society
104:23-33.
FRAHM, J.P. 2002. Ecology of bryophytes along
altitudinal and latitudinal gradients in Chile. Tropical
Bryol 21:67-79.
GONZALES-MANCEBO JM and HERNANDEZGARCIA CD. 1996. Bryophyte life strategies along an
altitudinal gradient in El Canal y Los Tiles (La Palma,
Canary Islands). J Bryol 19:243-255.
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