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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
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

They colonize diverse habitats from high mountains to


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
Philippine bryology where there is even more of a lack of
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Vol. 136 No. 1, June 2007

Hipol et al.: Life Strategies of Mosses in


Mt. Pulag, Philippines

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-DacrycarpusSyzygium-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)

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)

Table 1: Life strategy categories according to During (1979)


Potential Life
span (yr)
<1

12

Reproductive
Effort

Life strategy
Spores numerous, very light (<20 m)

Spores few, large (>20 m)

fugitives

Annual shuttle

high

few

Colonists

Short lived shuttle

high

many

Perennial stayers

Perennial Shuttle/Long lived shuttle

low

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Hipol et al.: Life Strategies of Mosses in


Mt. Pulag, Philippines

Table 2: Life strategy categories according to During (1992)


Potential Life span (yr)
<1

few

Life strategy

Reproductive Effort

Spores numerous, very light (<20 m)

Spores few, large (>20 m)

fugitives

Annual shuttle

Colonists

Medium

- Ephemeral Colonists

Short lived shuttle

- Colonists sensuo stricto

Long-lived shuttle

high

high

- Pioneers
Perennial stayers
many

- Competitive Perennial

Dominants

low

- 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.

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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Hipol et al.: Life Strategies of Mosses in


Mt. Pulag, Philippines

80%
60%

Pine Forest (~2440 masl)


Mossy Forest 2 (~2590 masl)
Mossy Forest 1 (~2725 masl)

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

Grassland (~2850 masl)

ni s
t

% occurrence

100%

Life Strategy

long lived

Pine Forest (~2440 masl)

Mossy Forest 2 (~2590


masl)

unknown
perennial

Mossy Forest 1 (~2725


masl)

100%
90%
80%
70%
60%
50%
40%
30%
20%
10%
0%

Grassland (~2850 masl)

% 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

see next page for continuation . . . .

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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

Bartramiaceae

m1

Long lived shuttle

Leiomela javanica
see next page for continuation . . . .

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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
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
16

most inhospitable of the zones for mosses. Colonist


(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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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 HernandezGarcia (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, 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.

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