Philippine Journal of Science

136 (1): 11-18, June 2007

ISSN 0031 - 7683

Life Strategies of Mosses in Mt. Pulag,

Benguet Province, Philippines

Roland M. Hipol*, Danielo B. Tolentino2, Edwino S. Fernando3, and Nia M. Cadiz2

1
Department of Biology, College of Science
University of the Philippines Baguio, Baguio City, Philippines
2
Plant Biology Department, Institute of Biological Sciences
University of the Philippines Los Baos, Los Baos, Laguna, Philippines
3
Makiling Center for Mountain Ecosystems
University of the Philippines Los Baos, Los Baos, Laguna, Philippines
Plant Biology Department, Institute of Biological Sciences

Mt. Pulag, the highest peak in Benguet Province, Luzon Island, Philippines at 2,924 m (Buot
and Okitsu 1998) was investigated for its moss flora focusing on life strategies. Moss collections
totaled 58 species that belong to 28 families. These are distributed among 3 vegetation zones; the
pine forest, mossy forest, and the mountain top grassland. Five out of the 6 main life strategies
of During (1979 and 1992) were observed to occur; namely colonists, fugitives, annual shuttle
species, long lived shuttle species, and perennials. The frequency of colonists, many of them
belonging to the genus Campylopus, was moderately high ranging between 40% and 50% at the
grassland and at the pine forest while it was lower in the mossy forest at 29%. Fugitives and
annual shuttle species were observed to occur only in the pine forest. Perennials were highest
in the mossy forest (45%), most of them belong to the family Hypnaceae. The long lived shuttle
and colonist strategies dominated the grassland. This survey on life strategies resulted in the
establishment of a moss based zonation pattern in Mt. Pulag as follows: colonist-annual shuttle
strategies at ~2,850 m (grassland), perennials at ~2,725 m (upper boundary of the mossy forest),
mixed perennials and colonists at 2,590 m (lower boundary of the mossy forest) and mixed
life strategy zone with the more desiccation tolerant strategies with the highest frequencies at
~2,440 m (pine forest). This zonation pattern coincides with the vegetation pattern suggested
by Merill and Meritt (1910).

Key Words: mosses, Bryophyta, Mt. Pulag, life strategies, altitudinal zonation

INTRODUCTION They colonize diverse habitats from high mountains to

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 Philippine bryology where there is even more of a lack of
deep forests (Shaw and Goffinet 2000); and even in hot
deserts though sporadic (Flowers 1973). They make up
the largest and most familiar group of bryophytes.
The Philippines, being a tropical country, is conducive
to rich growth of bryophytes. However, very few scientists
and researchers have taken interest in this very rich
bryological vegetation (del Rosario 1979). An aspect of
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Philippine Journal of Science
Vol. 136 No. 1, June 2007
information, is the relationship of these organisms with the
unique set of environmental conditions prevailing in the
different ecological zones of the country. Mosses are the
perfect indicator species of the stature of the environment
because the possibility that they are artificially introduced
in an ecosystem is practically nil. As such, their
existence and current state truly reflect what the ambient
environment really is.
Distinct environmental zones with increasing altitude
on Mt. Pulag, Benguet are reflected by the occurrence
of different vegetation zones on its slopes recently
elaborated by Buot and Okitsu in 1998. The first zone
they described was the pine forest (Pinus kesiya). Merill
and Meritt (1910) describe this vegetation as occupying
the main bulk of the mountain slopes up to an elevation
of 2,600 m. Jacobs (1972) explains that this vegetation
type is both a regeneration type of vegetation preparing
the ground for the primary forest (mossy forest), and a
destruction stage vegetation because of the recurring fire
that prevents the mossy forest to be next in the succession.
The upper limit of the pine forest is the mixed pine and
broad-leafed forest at the boundary of the pine and mossy
forest (Pinus-Deutzia-Schefflera).Above this vegetation
type is the mossy forest (Lithocarpus-Dacrycarpus-
Syzygium-Leptospermum). Merill and Meritt (1910)
vividly describe this forest being made up of dense stand
of small, irregularly shaped trees, comprising numerous
species, the ground, and trunks and branches of trees being
covered with a profusion of mosses, scale mosses, lichens,
epiphytic ferns and orchids. The highest vegetation
type is the grassland (e.g. Yushania, Calamagrostis,
Deschampsia) at the peak of the mountain. Merill and
Meritt (1910) mention that this is the only mountain in
the Philippines that has a well defined grass vegetation
above the mossy forest.
Bryophyte (mosses in particular) responses among
these zoneswould be expected to differ as these organisms
are generally considered to be strongly sensitive to
microclimatic (Frahm 2002). Currently, there is much
interest within ecology in integrated sets of plant traits
that accompany particular sets of ecological conditions
(During 1979,1992). Grime (1979) named these
combinations of characters strategies. Stearns (1976)
Table 1: Life strategy categories according to During (1979)
Potential Life
Life strategy
span (yr)
Spores numerous, very light (<20 m)
<1 fugitives
few Colonists
many Perennial stayers
12
Hipol et al.: Life Strategies of Mosses in
Mt. Pulag, Philippines
defined these as sets of co-adapted traits designed,
by natural selection, to solve particular ecological
problems. Bryophyte life strategy categories have been
refined in successive papers by During (1979, 1992). He
proposed several categories of bryophyte life strategies
and correlated these to environmental conditions where
such a tactic is observed. This classification was based
on 3 major trade-offs: (1) few large spores to many small
spores, (2) survival of the difficult season as spores only,
discarding the gametophyte (avoidance) versus survival
of the gametophyte (tolerance), and (3) for the tolerance
group, potential life span of the gametophyte, which
is negatively correlated with reproductive effort. This
resulted to the establishment of different life strategy
tactics namely: fugitives, colonists, perennials, annual
shuttles, short lived shuttles and long lived shuttles. In
1992, during modifies these categories emphasizing the
heterogeneity within the colonists and perennial life
strategies. As such, he created sub-categories under the
Colonist strategy namely: ephemeral colonists, colonists
sensuo stricto and pioneers. Under the perennial life
strategy, he distinguished competitive perennials from
stress tolerant perennials. Table 1 and 2 summarizes the
characteristics of the different life strategies.
These aspects of moss research, especially in our
country have not been dealt with satisfactorily in the past.
Expressions of this response in the form of diverse life
strategies are facets of moss biology that is the interest
of this paper.
MATERIALS AND METHODS
Four collection sites were identified in the different
vegetation zones:
pine forest at ~2440 m
mossy forest (lower altitude) at ~2590 m
mossy forest (higher altitude) at ~2725 m
grassland at ~2850 m.
Field work was carried out in the first weeks of May
and September 2003, in the first weeks of February and
in April 2004. Sampling was done within the different
vegetation zones recognized by Merill and Merritt (1910)
Reproductive
Effort
Spores few, large (>20 m)
Annual shuttle high
Short lived shuttle high
Perennial Shuttle/Long lived shuttle low
Philippine Journal of Science
Vol. 136 No. 1, June 2007
Table 2: Life strategy categories according to During (1992)
Potential Life span (yr)
Spores numerous, very light (<20 m)
<1 fugitives
Colonists
- Ephemeral Colonists
few
- Colonists sensuo stricto
- Pioneers
Perennial stayers
many - Competitive Perennial
- Stress Tolerant Perennial
through the multiple or multi-stage stratified sampling
method. In this method, the population is subdivided
or partitioned into strata, and each stratum is sampled
separately. Partitioning is typically done so as to make
each stratum more homogeneous than the overall
population. For this research, the primary strata were
the different collection sites as determined by altitude
and the identified vegetation types of Merill and Meritt
(1910). Collection at the pine forest was at ~2440m; at
the blower edge of the mossy forest at ~2725m; at the
upper boundary of the mossy forest at ~2590m; and at
the grassland at ~2850m. The second strata were the
different substrate preferences (tree, rock, soil) in each
of the identified elevation. Mosses in these substrates
were scored (species occurrence and cover) using a 625
cm2 quadrat (0.25 x 0.25 m). The presence or absence of
each species in each habitat type and in each vegetation
zone was noted. Identification was done using Bartrams
(1939) Mosses of the Philippines and Eddys (1996)
Handbook on Malesian Mosses. Validation of the
scientific names and identification of difficult taxa were
done in the Cryptogamic Laboratory of the National
University of Singapore under the supervision of Dr.
Benito Tan. Voucher specimens of the collected mosses
are also deposited at this facility.
The classification of bryophyte life strategies followed
that proposed by During (1979).
Site Description
Mt. Pulag (163036 N; 1205020E) is the highest
peak in Luzon island at 2,924m (Buot and Okitsu 1998).
Together with Mt. Tabayoc and Mt. Panotoan, it forms
the Mt. Pulag National Park. The park covers an area of
about 11,560 ha lying on the north to south spine of the
Gran Cordillera Central. The municipalities of Bokod,
Kabayan, and Buguias of the province of Benguet, Tinoc
of Ifugao, and Kayapa of Nueva Vizcaya bound it.
Hipol et al.: Life Strategies of Mosses in
Mt. Pulag, Philippines
Life strategy Reproductive Effort
Spores few, large (>20 m)
Annual shuttle high
Medium
Short lived shuttle
high
Long-lived shuttle
Dominants low
Buot and Okitsu (1998) determined that the annual
mean temperature at 2000m is 16C. At the upper limit
of the pine forest (2300m), the temperature is 14C. At
elevations around 2800m the annual mean temperature
is around 12C. The mountain is also characterized by
having a vegetation zonation pattern unique from other
mountains in that it does not have a dipterocarp flora in
its lower slopes. In the publication of Merrill and Merritt
(1910), they recorded a total of 57 bryophyte species, 91
fern species, 3 gymnosperms and 377 angiosperms for
Mt. Pulag.
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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Philippine Journal of Science Hipol et al.: Life Strategies of Mosses in
Vol. 136 No. 1, June 2007 Mt. Pulag, Philippines

100%
% occurrence 80% Pine Forest (~2440 masl)
60% Mossy Forest 2 (~2590 masl)
40% Mossy Forest 1 (~2725 masl)
20% Grassland (~2850 masl)
0%

n
d
al

l
e
t

nia
ni s

ow
ive
ive
itiv
nu

ren
lo

kn
gl
rt l
an

fug
co

un
lon
o

pe
sh

Life Strategy
Figure 1. Abundance of each moss life strategy in each of the vegetation zones in %s.

100%
90%
80%
% occurrence

70%
60%
50%
40%
30% unknown
20%
perennial
10%
0% long lived
Grassland (~2850 masl)

Pine Forest (~2440 masl)

Mossy Forest 1 (~2725

Mossy Forest 2 (~2590

short lived
fugitive
annual
masl)

masl)

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
see next page for continuation . . . .

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Philippine Journal of Science Hipol et al.: Life Strategies of Mosses in
Vol. 136 No. 1, June 2007 Mt. Pulag, Philippines

Bryum ramosum Bryaceae pine Colonist

Calyptrochaeta remotifolia Hookeriaceae m1 Long lived shuttle
Calyptrochaeta remotifolia Hookeriaceae grassland Long lived shuttle
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 Long lived shuttle
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 Long lived shuttle
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 Long lived shuttle
Isothecium trichocladon Lembophyllaceae m2 Perennial
Leiomela javanica Bartramiaceae m1 Long lived shuttle
see next page for continuation . . . .

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Philippine Journal of Science Hipol et al.: Life Strategies of Mosses in
Vol. 136 No. 1, June 2007 Mt. Pulag, Philippines

Leucobryum aduncum var. scalare Leucobryaceae pine Perennial

Macromitrium reinwardtii Orthotrichaceae pine Long lived shuttle
Macromitrium reinwardtii Orthotrichaceae grassland Long lived shuttle
Macromitrium reinwardtii Orthotrichaceae m2 Long lived shuttle
Macromitrium sulcatum Orthotrichaceae pine Long lived shuttle
Meteoriopsis reclinata Meteoriaceae pine Long lived shuttle
Miothecium microcarpum Sematophyllaceae pine Long lived shuttle
Mnium laevinerve Mniaceae m1 unknown
Mnium laevinerve Mniaceae m1 unknown
Papillaria fuscescens Meteoriaceae m2 Long lived shuttle
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 Long lived shuttle
Syrrhopodon tjibodensis Calymperaceae pine Colonist
Taxiphyllum taxirameum Hypnaceae m1 Perennial
var. recurvifolium
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 most inhospitable of the zones for mosses. Colonist

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
(Campylopus spp) and long lived shuttle strategies
(Calyptrochaeta remotifolia), being the life strategies
fit for this kind of environment, are dominant. Annual
shuttle strategy described by During (1979) is strongly
determined by seasonal fluctuations and a severe stress
period which is avoided by being in the spore stage only.
During (1979) also describes the colonists as a strategy
that is adapted to environments that are unpredictable, but
are more or less predictably last for some years. Colonists
also often appear early in the succession series. However,
the idea that the presence of colonists in the grasslands
may signify succession may not be correct. Because of

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Philippine Journal of Science
Vol. 136 No. 1, June 2007
the prevalent drying conditions, this situation may be
considered as auto-succession (Longton 1992). Longton
adopts the definition of auto-succession as a succession
that consists of a single stage, in which pioneer and climax
species are the same. It occurs particularly where climatic
severity so restricts the number of species that competition
is minimized and displacement fails to occur. In this
particular situation, the colonists, by themselves are also
the climax species of the grassland.
Mossy Forest Moss Flora
This vegetation type is the richest habitat for bryophytes.
The abundance of long span life strategies (long lived
shuttles and perennials) demonstrate that the environment
is stable and humid enough for long periods of moss
growth. During (1979) also adds that these strategies exist
in environments where even if there are fluctuations in the
environment, these changes are within the tolerance range
of these organisms. Gradstein and Pocs (1989) explain
this abundance as a result of more favorable moisture
conditions due to clouds and fog and the prevailing
lower temperatures. Gonzales-Mancebo and Hernandez-
Garcia (1996) also found this out in their research on
the life strategies of mosses in the Canary Islands.
Ecophysiologically, temperature and insulation may have
been optimal in this environment that assimilation exceeds
that of respiration as evidenced by the high bryophyte
biomass estimated at 1053.9 kg/ha (Frahm 1990). The
dominance of perennial life strategies facilitates the
conclusion that the bryophyte flora in the mossy forests
is the climax community.
Pine Forest Moss Flora
Bryophyte flora in this vegetation zone is unique in that
it is only here that the short life span strategies are found
(annual shuttle species and fugitives) together with
colonists. Perennial strategists were conspicuously scarce.
The presence of short lived strategy types suggests that
there are strong seasonal fluctuations and severe stress
periods that are purposely avoided by the bryophytes being
present in the spore stage only. The colonists in this zone
may be in the category described by During (1992) as
sensuo stricto because they colonize available productive
habitats that resulted from disturbance. The seasonal
fires within the pine forest explain the predominance of
these types of strategies. The low density of pine trees,
which results to an opened canopy is dictated by the pines
ecophysiological character of being light demanding
and germinating only on bare soil (Jacobs 1972). This
condition makes this habitat less humid and dry for
perennial strategists.
Hipol et al.: Life Strategies of Mosses in
Mt. Pulag, Philippines
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, SEARCA-
SEAMEO 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.
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