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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 They colonize diverse habitats from high mountains to


deep forests (Shaw and Goffinet 2000); and even in hot
Mosses are remarkably successful plants that thrive deserts though sporadic (Flowers 1973). They make up
alongside more conspicuous vascular plants. They the largest and most familiar group of bryophytes.
originated during the Upper Ordovician-Lower Silurian
border (~440 million years) together with the first land The Philippines, being a tropical country, is conducive
plants (Shaw and Goffinet 2000). They are currently to rich growth of bryophytes. However, very few scientists
represented by approximately 10,000 species worldwide. and researchers have taken interest in this very rich
bryological vegetation (del Rosario 1979). An aspect of
*Corresponding author: udangya@hotmail.com Philippine bryology where there is even more of a lack of
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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

information, is the relationship of these organisms with the defined these as sets of co-adapted traits designed,
unique set of environmental conditions prevailing in the by natural selection, to solve particular ecological
different ecological zones of the country. Mosses are the problems. Bryophyte life strategy categories have been
perfect indicator species of the stature of the environment refined in successive papers by During (1979, 1992). He
because the possibility that they are artificially introduced proposed several categories of bryophyte life strategies
in an ecosystem is practically nil. As such, their and correlated these to environmental conditions where
existence and current state truly reflect what the ambient such a tactic is observed. This classification was based
environment really is. on 3 major trade-offs: (1) few large spores to many small
spores, (2) survival of the difficult season as spores only,
Distinct environmental zones with increasing altitude discarding the gametophyte (avoidance) versus survival
on Mt. Pulag, Benguet are reflected by the occurrence of the gametophyte (tolerance), and (3) for the tolerance
of different vegetation zones on its slopes recently group, potential life span of the gametophyte, which
elaborated by Buot and Okitsu in 1998. The first zone is negatively correlated with reproductive effort. This
they described was the pine forest (Pinus kesiya). Merill resulted to the establishment of different life strategy
and Meritt (1910) describe this vegetation as occupying tactics namely: fugitives, colonists, perennials, annual
the main bulk of the mountain slopes up to an elevation shuttles, short lived shuttles and long lived shuttles. In
of 2,600 m. Jacobs (1972) explains that this vegetation 1992, during modifies these categories emphasizing the
type is both a regeneration type of vegetation preparing heterogeneity within the colonists and perennial life
the ground for the primary forest (mossy forest), and a strategies. As such, he created sub-categories under the
destruction stage vegetation because of the recurring fire Colonist strategy namely: ephemeral colonists, colonists
that prevents the mossy forest to be next in the succession. sensuo stricto and pioneers. Under the perennial life
The upper limit of the pine forest is the mixed pine and strategy, he distinguished competitive perennials from
broad-leafed forest at the boundary of the pine and mossy stress tolerant perennials. Table 1 and 2 summarizes the
forest (Pinus-Deutzia-Schefflera).Above this vegetation characteristics of the different life strategies.
type is the mossy forest (Lithocarpus-Dacrycarpus-
Syzygium-Leptospermum). Merill and Meritt (1910) These aspects of moss research, especially in our
vividly describe this forest being made up of dense stand country have not been dealt with satisfactorily in the past.
of small, irregularly shaped trees, comprising numerous Expressions of this response in the form of diverse life
species, the ground, and trunks and branches of trees being strategies are facets of moss biology that is the interest
covered with a profusion of mosses, scale mosses, lichens, of this paper.
epiphytic ferns and orchids. The highest vegetation
type is the grassland (e.g. Yushania, Calamagrostis,
MATERIALS AND METHODS
Deschampsia) at the peak of the mountain. Merill and
Four collection sites were identified in the different
Meritt (1910) mention that this is the only mountain in
vegetation zones:
the Philippines that has a well defined grass vegetation
above the mossy forest. pine forest at ~2440 m
Bryophyte (mosses in particular) responses among mossy forest (lower altitude) at ~2590 m
these zoneswould be expected to differ as these organisms mossy forest (higher altitude) at ~2725 m
are generally considered to be strongly sensitive to grassland at ~2850 m.
microclimatic (Frahm 2002). Currently, there is much
interest within ecology in integrated sets of plant traits Field work was carried out in the first weeks of May
that accompany particular sets of ecological conditions and September 2003, in the first weeks of February and
(During 1979,1992). Grime (1979) named these in April 2004. Sampling was done within the different
combinations of characters strategies. Stearns (1976) vegetation zones recognized by Merill and Merritt (1910)

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


Potential Life Reproductive
Life strategy
span (yr) Effort
Spores numerous, very light (<20 m) Spores few, large (>20 m)
<1 fugitives Annual shuttle high
few Colonists Short lived shuttle high
many Perennial stayers Perennial Shuttle/Long lived shuttle low

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Philippine Journal of Science Hipol et al.: Life Strategies of Mosses in
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Table 2: Life strategy categories according to During (1992)


Potential Life span (yr) Life strategy Reproductive Effort
Spores numerous, very light (<20 m) Spores few, large (>20 m)
<1 fugitives Annual shuttle high
Colonists Medium
- Ephemeral Colonists Short lived shuttle
few high
- Colonists sensuo stricto Long-lived shuttle
- Pioneers
Perennial stayers
many - Competitive Perennial Dominants low
- Stress Tolerant Perennial

through the multiple or multi-stage stratified sampling Buot and Okitsu (1998) determined that the annual
method. In this method, the population is subdivided mean temperature at 2000m is 16C. At the upper limit
or partitioned into strata, and each stratum is sampled of the pine forest (2300m), the temperature is 14C. At
separately. Partitioning is typically done so as to make elevations around 2800m the annual mean temperature
each stratum more homogeneous than the overall is around 12C. The mountain is also characterized by
population. For this research, the primary strata were having a vegetation zonation pattern unique from other
the different collection sites as determined by altitude mountains in that it does not have a dipterocarp flora in
and the identified vegetation types of Merill and Meritt its lower slopes. In the publication of Merrill and Merritt
(1910). Collection at the pine forest was at ~2440m; at (1910), they recorded a total of 57 bryophyte species, 91
the blower edge of the mossy forest at ~2725m; at the fern species, 3 gymnosperms and 377 angiosperms for
upper boundary of the mossy forest at ~2590m; and at Mt. Pulag.
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 RESULTS
cm2 quadrat (0.25 x 0.25 m). The presence or absence of
The moss collections from Mt. Pulag totaled 58 species
each species in each habitat type and in each vegetation
that belong to 28 families bringing overall inventory to
zone was noted. Identification was done using Bartrams
144 species, an increase of 27 species from the previous
(1939) Mosses of the Philippines and Eddys (1996)
count of 117 (from those documented by Bartram in 1939
Handbook on Malesian Mosses. Validation of the
and Tan and Iwatsuki in 1991). Table 3 summarizes the
scientific names and identification of difficult taxa were
collected specimens.
done in the Cryptogamic Laboratory of the National
University of Singapore under the supervision of Dr.
Benito Tan. Voucher specimens of the collected mosses Life Strategy and Vegetation Type
are also deposited at this facility. Five out of the six life strategies of During (1979) have
been observed to occur in the moss populations of Mt.
The classification of bryophyte life strategies followed Pulag. The only strategy that was not seen was the
that proposed by During (1979). short lived shuttle. As can be observed, the frequency
of colonists in all of the collection sites is quite high,
Site Description however lesser in mossy forest 1 (~2725 m). Fugitives
Mt. Pulag (163036 N; 1205020E) is the highest and annual shuttle species are observed to occur only in
peak in Luzon island at 2,924m (Buot and Okitsu 1998). the pine forest. The frequency of perennials and long
Together with Mt. Tabayoc and Mt. Panotoan, it forms lived shuttle species vary between the different collection
the Mt. Pulag National Park. The park covers an area of sites. For perennials, the mossy forests are the habitats
about 11,560 ha lying on the north to south spine of the that they prefer basing on the relatively high frequency of
Gran Cordillera Central. The municipalities of Bokod, occurrence of this strategy in the two collection sites. The
Kabayan, and Buguias of the province of Benguet, Tinoc long lived shuttle dominate the grassland. Abundance of
of Ifugao, and Kayapa of Nueva Vizcaya bound it. 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
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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 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


(Campylopus spp) and long lived shuttle strategies
Overall affinities of specific life strategies to the different (Calyptrochaeta remotifolia), being the life strategies
altitudinal vegetation zones have been found. The fit for this kind of environment, are dominant. Annual
following discuss the observed moss based vegetation shuttle strategy described by During (1979) is strongly
pattern correlated with the dominant vegetation pattern determined by seasonal fluctuations and a severe stress
mentioned by Merill and Meritt (1910). period which is avoided by being in the spore stage only.
During (1979) also describes the colonists as a strategy
Grassland Moss Flora that is adapted to environments that are unpredictable, but
The absence of a tree canopy that could protect against are more or less predictably last for some years. Colonists
intense light and high wind velocity makes the grassland also often appear early in the succession series. However,
environment particularly desiccating for such organisms, the idea that the presence of colonists in the grasslands
and may be the prime reason why the grassland is the may signify succession may not be correct. Because of

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Philippine Journal of Science Hipol et al.: Life Strategies of Mosses in
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the prevalent drying conditions, this situation may be CONCLUSION


considered as auto-succession (Longton 1992). Longton
adopts the definition of auto-succession as a succession Mt. Pulag bryoflora is one of the more diverse communities
that consists of a single stage, in which pioneer and climax of bryophytes especially of mosses. It is here that many
species are the same. It occurs particularly where climatic of the mosses and other organisms find their last refuge
severity so restricts the number of species that competition in view of the continuous and large-scale land conversion
is minimized and displacement fails to occur. In this happening throughout the country. It is also where life
particular situation, the colonists, by themselves are also strategy distribution has never been investigated. In this
the climax species of the grassland. 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
Mossy Forest Moss Flora of Merill and Merrits (1910) 3 distinct vegetation zones.
This vegetation type is the richest habitat for bryophytes. The following moss-life-strategy based zonation would be:
The abundance of long span life strategies (long lived (1) Mixed strategy Zone where those with short life spans
shuttles and perennials) demonstrate that the environment are present (Fugitives and Annual Shuttle species) at the
is stable and humid enough for long periods of moss pine forest, (2) Perennial Stayers and Long Lived Shuttle
growth. During (1979) also adds that these strategies exist Zone within the mossy forest, and (3) Colonists-Annual
in environments where even if there are fluctuations in the Shuttle Species Zone at the grassland. The dominance of
environment, these changes are within the tolerance range these life strategies unique to each of the vegetation zones
of these organisms. Gradstein and Pocs (1989) explain illustrates the distinctly different environmental conditions
this abundance as a result of more favorable moisture offered by each of these ecosystems to which the mosses
conditions due to clouds and fog and the prevailing have to adapt to. It would be very interesting to determine
lower temperatures. Gonzales-Mancebo and Hernandez- what these factors are that specifically favor the abundance
Garcia (1996) also found this out in their research on of specific strategies. In the words of During (1992), results
the life strategies of mosses in the Canary Islands. of these types of researches are a valuable guide in the further
Ecophysiologically, temperature and insulation may have investigation regarding environmental key factors that define
been optimal in this environment that assimilation exceeds specific environments or regions.
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 ACKNOWLEDGEMENTS
is the climax community.
We would like to thank Dr. Benito Tan of the National
University of Singapore for his invaluable help in the
Pine Forest Moss Flora identification of the moss collections, and Dr. Inocencio Buot
Bryophyte flora in this vegetation zone is unique in that and Prof. Maria Fe Sangalang of UP Los Baos for the help in
it is only here that the short life span strategies are found the preparation of this research. Our gratitude also to the Tan
(annual shuttle species and fugitives) together with Kin Chee Foundation for a small grants award, SEARCA-
colonists. Perennial strategists were conspicuously scarce. SEAMEO for the thesis grant, and UP Baguio for the Local
The presence of short lived strategy types suggests that Faculty Fellowship grant to the corresponding author. The
there are strong seasonal fluctuations and severe stress assistance of Mr. Orlando Apostol, Ronnel Almazan, and Rexel
periods that are purposely avoided by the bryophytes being Almazan during field work is also gratefully acknowledged.
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
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