B I O D I V E R S I T A S ISSN: 1412-033X (printed edition)

Volume 12, Number 4, October 2011 ISSN: 2085-4722 (electronic)

Pages: 212-217 ' 2 ,

Species composition and interspecific association of plants in primary

succession of Mount Merapi, Indonesia

SUTOMO1,, DINI FARDILA2, LILY SURAYYA EKA PUTRI2

1
Bali Botanic Garden, Indonesian Institute of Sciences, Candikuning, Baturiti, Tabanan 82191, Bali, Indonesia. Tel. +62-368-21273. Fax. +62-368-
22051. email: [Link]@[Link]
2
Biology Department, Faculty of Science and Technology, Syarif Hidayatullah State Islamic University, South Tangerang 15412, Banten, Indonesia.

Manuscript received: 12 April 2011. Revision accepted: 24 August 2011.

ABSTRACT

Sutomo, Faradila D, Putri LSE (2011) Species composition and interspecific association of plants in primary succession of Mount
Merapi, Indonesia. Biodiversitas 12: 212-217. Primary succession refers to the establishment of plant species and subsequent changes in
composition following major disturbance such as volcanic activity. The study of succession may assist in recognizing the possible
effects of species interactions (i.e. facilitation or inhibition). The barren landscapes created by volcanic disturbance on Mount Merapi,
Java, Indonesia, provide excellent opportunities to study primary succession. Fifty -six species belonging to 26 families were recorded in
the five nues ardentes deposits. The highest number of species belonged to the Asteraceae, then Poaceae, followed by Fabaceae and
Rubiaceae. In Mount Merapi primary succession, the ecosystem may be developing with time as indicated by the increase in the number
of species associations. The number of positive associations was generally higher than the number of negative associations, except in the
2001 deposit where it was equal. Native and alien invasive species had different patterns of interspecific associations. This research
demonstrates that in primary succession sites on Mount Merapi, positive interspecific association increased as time progressed, which
may support the view that facilitation is more prominent in a severely disturbed habitat as compared to competition.

Key words: primary succession, interspecific association, interaction, facilitation, pioneer, Mount Merapi.

INTRODUCTION does not support any organism (Walker and del Moral
2003). Vegetation establishment on volcanic deposits has
Volcanoes has shape many of the Earth landscapes been documented in many parts of the world such as in
(Dale et al. 2005a). More than half of the active terrestrial USA, Italy and Japan and their rates have been shown to
volcanoes encircle the Pacific Ocean and are known as the vary (Eggler 1959; Tsuyuzaki 1991; Aplet et al. 1998; Dale
ring of fire. Hence, there are many parallel situations in et al. 2005c). For example, plant establishment and spread
the world where volcanic activity has become a major on the debris-avalanche deposit were slow during the first
disturbance such as in Hawaii (Mount Mauna Loa), New years after eruption of Mt St Helens in USA (Dale et al.
Zealand (Mount Ruapehu), USA (Mount St. Helens), and 2005c).
Indonesia (Mount Krakatau). Indonesia is particularly Species interactions are of central importance in the
unique because of a series of active volcanoes which stand study of succession. The study of succession may assist in
in line from the Sumatran Island to Java Island. With 130 recognizing the possible effects of species interactions (i.e.
active volcanoes lies on its region, Indonesia has become facilitation or inhibition) (Connell and Slatyer 1977; Walker
the most volcanic country on Earth (Weill 2004). et al. 2007). Facilitation promotes establishment and in the
Primary succession refers to establishment of plant context of succession, facilitation can be defined as any
species and their changes in composition following major role of plants that influences a change in species
disturbance such as volcanic disturbance (Walker and del composition to the next stage (Walker and del Moral
Moral 2003). One type of volcanic disturbance is nues 2003). Previous studies have shown that in a severely
ardentes or pyroclastic flows. Nues ardentes are hot
disturbed habitat, the role of facilitation will be more
turbulent gas and fragmented material resulting from a
prominent for species change and restoration, whereas
collapsed lava dome that rapidly moves down the volcanic
slope (Dale et al. 2005b). The accumulation of this material competition tends to be significant in a more productive
is called a nues ardentes deposit and it may be up to 10 m and established habitat (Callaway and Walker 1997;
thick (Franklin et al. 1985). Volcanic eruptions are strongly Walker et al. 2007). The barren landscapes created by
linked to depositions of volcanic materials avalanche to volcanic disturbance provide excellent opportunities to
form un-vegetated barren areas which started primary examined the role of pioneer species in facilitating or
succession. Primary succession commence on a barren inhibiting later species in succession (Morris and Wood
substrate that does not have any biological legacies and 1989; Walker and del Moral 2003). However, initial
interactions occurring during primary succession that drive
 SUTOMO et al. Species composition and plants association of Mount Merapi 213

the subsequent community composition remain studied in

only a few locations (Connell and Slatyer 1977; Climatologically, based on Schmidt and Fergussons
Bellingham et al. 2001). climate classification, the Merapi area is classified as a
The nues ardentes deposits found in Mount Merapi are type B, tropical monsoon area, which is characterized by
relatively young, with the last known eruptions occurring high intensity of rainfall in the wet season (November-
between 1994 and 2006. Here we examine whether or not April) and then the dry season (April-October). Its annual
early interaction patterns among species can be identified precipitation varies from 1,500-2,500 mm. The variation of
by examining their interspecific association and test the rainfall on Merapi slope is influenced by orographic
hypothesis that positive association will more apparent precipitation. Like in many other tropical monsoon areas,
compared with negative association over time. there are minor temperature and humidity variations.
Merapis relative humidity varies from 70 -90% with daily
average temperatures varying from 19-30 C (Forest Office
MATERIALS AND METHODS of Yogyakarta 1999).
The research sites were located in the southwest flank
Study sites forests of Mount Merapi within Merapi National Park.
Merapi is one of the most active volcanoes in Indonesia These sites are the most prone to and most often affected
which is located 30 km North of Yogyakarta Province in by volcanic disturbance due to the nues ardentes that tend
Java Island at 735 S and 11024 E (Figure 1). to flow down the hills in this direction. Using

Scale 1:25.000

Figure 1. Map of Mt. Merapi National Parks eruption deposits. Circular symbols refer to the position of sampling sites in each deposit.
The rectangle refers to the site position of an undisturbed forest in Kaliurang, Yogyakarta.
214 B I O D I V E R S I T A S 12 (4): 212-217, October 2011

chronosequence (space for time substitution) method, we Interspecific association

chose five areas that were affected by nues ardentes Interspecific association between species was measured
deposits between 1994 and 2006 (Figure 1). The five using the chi-square (2) test of the species presence/ absence
deposit sites were located in a lower montane zone data on a 2 2 contingency table (Ludwig and Reynolds
(Montagnini and Jordan 2005). The 1994 sampling site or 1988; Kent and Coker 1992; Supriyadi and Marsono 2001).
the late primary succession site is located in an area
surrounding the Kuning River, at an altitude of 1,180 m. Species B
In this late primary successional site the vegetation is Present Absent
generally composed of Eupatorium odoratum (Asteraceae) Species Present a b a+b
A Absent c d c+d
and Imperata cylindrica (Poaceae).
a+c b+d N = a+b+c+d
Sampling
Vegetation on the five nues ardentes deposits was a = the number of sampling unit (SU) where both
sampled in 2008. We sampled ten 250 m 2 circular plots in species occur
each deposit (50 plots in total), assigned at random to grid b = the number of SUs where species A occur but not B
cells on a map Dale et al. 2005c). Each plot was located in c = the number of SUs where species B occur but not A
the field with reference to a compass and a handheld Global d = the total number of SUs
Positioning System GPS (Garmin E-Trex Legend). We
measured plant abundance as density, a count of the Then a chi square test statistic is employed to test the
numbers of individuals of a species within the quadrat null hypotheses of independence in the 2 2 table:
(Kent and Coker 1992; Endo et al. 2008). We noted both
local plant name and scientific name (when known). x2= ( ad - bc )2 N
Whenever there was any doubt about species name, a ( a + b) (a + c) (b + d) (c + d)
herbarium sample was made. Drying and sample
identification were done in Laboratory of Dendrology,
Faculty of Forestry, Universitas Gadjah Mada, Yogyakarta. The significance of the chi-square test statistic is
Vascular plant nomenclature is based on Backer and determined by comparing it to the theoretical chi-square
Bakhuizen van den Brink (1963). Although homogeneity of distribution (P = 0.05, df = 1) There are two type of
the sites was taken into account when choosing sample association:
sites, differences in site conditions were likely to occur. Positive, if x2 test > x2 theoretical and observed a >
Hence, for each circular plot, site attributes (altitude and (a + b) (a + c)
expected a, Where expected a = , that is the
slope) were measured. Altitude was measured using a GPS N
and referenced against 1:25,000 topographic maps. A pair of species occurred together more often than expected.
clinometer (Suunto PM-5) was used to determine the slope
Negative, if x2 test > x2 theoretical and observed a <
(in degrees) (Le Brocque 1995). expected a, that is to say that the pair of species occurred
together less often than expected.
Data analysis
The strength level of the association was measured
Species composition using the Ochiai index, which is equal to 0 at no
Plant community composition between deposits was association and to 1 at complete/maximum
described by Curtis and McIntoshs Importance Value association(Kent and Coker 1992).
Index or IVI (1950).
Ochai Index = a
IVI = RD+RF (a + b) (a + c)

IVI = Importance Value Index

RD = Relative Density RESULTS AND DISCUSSION

Fifty-six species belonging to 26 families were

Relative Density of A species = recorded in the five nues ardentes deposits which mostly
comprise of species belonged to the Asteraceae (herbs),
Number of individual of A species x 100%
then Poaceae (grasses), followed by Fabaceae (N 2 fixing
tree seedling) and Rubiaceae (shrub). Based on vegetation
Total number individual of all species
analysis with IVI computation, we found that each deposit
has almost similar set of species composition except for the
RF = Relative Frequency latest deposits sites namely 2006 and 2001 (Figure 2).
Some species such as Anaphalis javanica, Eupatorium
Relative Frequency of A species = riparium and I. cylindrica showed consistency of their
appearance in almost all of the deposits (Figure 3). It is
Frequency value of A species x 100% interesting to see that these species have fluctuated over
Total frequency value of all species time except for invasive pioneer I. cylindrica which was
 S UTOMO et al. Species comp osition and pla nts association of Mount Merapi 21 5

d eclining in IVI index. Th is phenomeno n may reflect that Native and invasive speci es had differ ent patterns of
the abundanc e and domination of I. cylindrica decreasing a interspecific asso ciations (Table 2). Amo ng the nativ e
s the comm unity developed over time. The presence of o species, A. java nica possess ed the highe st number of
ther pioneer species such as A. java nica and inv asive nega tive associations with other species, followed by Pin us
s pecies such as E. ripari um may ha ve suppresse d the mer kusii. In con trast, Calliandra callothyrsus had th e
d omination of I. cylindrica in more devel oped sites. highest number of positive interspecific associatio n compare
to Athyrium macrocarpum. A. ma crocarpum ha d the most
tendencies to co-occu r (or to absent together) with Polygala
panicul ata as shown by their str ongest positiv e association.
Am ong the invasive specie s however, I. cylin drica is very
aggressive a nd may beco me dominant in the site as indica ted
by the absence of othe r co-occurrin g species in all deposit
sites. E. riparium was more likely to
occu r together with Melastoma affine, whereas th e
presence of Calliandra callothyrsus is mo re likely wi th
Cyp erus rotundus (Table 2).
T he nitrogen fixing legume, C. callothyrsus showed th e
highest number of positive ass ociations with other species,
Figure 2. Dominant species based on Importance Value In dex in mos tly grasses su ch as C. rotundus and Eleusine indica.
e ach deposits o f primary succession on Mount Merapi Native from Mex ico, this species is now wi dely introduce d
in many tropical regions. C. callothyrsus is a ble to grow o n a
wide range of s oils types, including the mo derately acidic
volcanic origin soils that are a common feature in th e
Southeast Asia ( Palmer et al. 1994). This species is now
naturalized in Asia including In donesia (Pal mer et al. 199 4)
T he mistflow er ( E. riparium) had a higher number of
positive associati ons as compared to negative associatio
ns (Tab le 2). This species is also the domina nt groundcov
er species in Kaliura ng, an intact forest on the southern
slop e of M ount Merapi (Sutomo 2004). This species may
hav e
indir ectly facilitated co-occ urring species such as
Gnaphalium jap onicum and M. affine by assisting in
Figure 3. Chan ges in IVI of some pioneer species of inte rest in stabi lizing and preventing er osion on the deposit site
e ach deposits o f primary succession on Mount Merapi. (Hey ne 1987) . H owever, over domination by this invasiv
e species could be a problem itself. Eupatori um is native
to South America, and this unpalatable and high ly
competitive species has become a problem e lsewhere, suc
h as in Nepal (Kun war 2003).
C ogongrass (I. cylindrica) did not exhibit an y
association with o ther species in any deposits (Table 2). I.
cylin drica is an aggressive al ien invader that has a lon g
record of colonising cleared lands in Indonesia ( A.
Ham blin, personal communi cation, 28 May 2009). I.
cylin drica domination in Mo unt Merapi nues ardent es
depo sits is presum ably due to its wide-sprea d rhizomes an d
its w ind-disperse d seeds (Jon athan and Ha riadi 1999). I.
Figure 4. Interspecific association of species in each deposit at p
cylin drica may h ave contribut ed indirectly to the increa se in
rimary success ion sites of different age.
the number of species coloni zing the depo sits, especial ly in
th e early stages, by altering the soil prop erties (Walk er and
The numb er of positive associatio ns was generally h del Moral 200 3; Collins and Jose 2009).
igher than the number of negative asso ciations, exc ept in S pecies changes are do not only occuring in response
the 2001 dep osit, where it was equal ( Figure 4). Positive to chan ges in physical environment, but can al so be the
a ssociations w ere highest in 1994, lowest in 2001 and 1998 resu lt of in teraction with another spec ies, thus spec ies
a nd intermed iate value in 2006 and 1997 deposits. interactio ns are also an important indicati ng factor in
Generally, th e number of negative associations for each succession an d ecosystem devel opment (Wal ker and del
d eposit was le ss than positive associatio ns. Less Moral 200 3; Mul ler 2005). Species co-occu rrence obser
variability o ccurred in the negative ass ociations. Negative vations may b e seen as the first attempt to detect species
associ ations were lowest in 2001 and 19 98, but almo st interaction (i.e.
similar for 2006, 1 997 and 199 4 deposits. facilitation and inhibition) and niche processes that
struc ture a com munity (Wal ker and del Moral 200 3;
Widyatmoko and Burgman 200 6).
216 B I O D I V E R S I T A S 12 (4): 212-217, October 2011

While the general explanation of why two species are competition (Callaway and Walker 1997). Primary
positively associated is because they favor the same succession on Mount St. Helen was reported to be very
environmental conditions, this explanation is not always as slow due to isolated and physically stressful habitat
apparent as might first appear and may be over-simplistic however, facilitation by nitrogen fixing species such as
(Kent and Coker 1992; Belyea and Lancaster 1999; Lupinus lepidus may have also occurs (del Moral and
Ruprecht et al. 2007). There are other factors such as plant Wood 1993). Positive interaction in plant communities is
species strategies, competition and interaction that also more common than negative interaction in high-elevation
need to be considered (Belyea and Lancaster 1999; Dukat ecosystems (Callaway 1998; Endo et al. 2008). However,
2006). Facilitation may have a more vital role in species there has been accumulating evidence that stated
change and restoration in a severely disturbed habitat, facilitation is the dominant form of interaction in many
whereas competition will tend to be important in a more ecosystems (Callaway 2007).
productive and established habitat (Callaway and Walker Plant association has also been found in other volcanic
1997; Walker et al. 2007). Furthermore, one of the most sites across the globe. Early associations comprised of
important questions in plant community assembly rules Honkenya peploides, a low-growing, sand-binding pioneer,
may be generated from this observation: which lyme grass, Elymus arenarius, and the lungwort, Mertensia
combination of species occurs together and why? (Bond maritima, have contributed to the development of a
and Wilgen 1996). relatively unstable ecosystem on Surtsey, a volcanic island
In primary succession on Mount Merapi, the primary in Iceland 30 years after eruption (Thornton 2007) . On the
succession ecosystem may be developing to later stages volcanic island of Krakatau in Indonesia, the beach-
with time as indicated by the increase in the number of creepers Ipomoea pes-caprae and Canavalia rosea, and the
species associations. Differences in the number of grasses I. cylindrica (alang-alang ) or Saccharum
occurrences of positive associations with the negative spontaneum ( glagah), have been found to form association
associations were also recorded. Generally positive related to the slowly growing sand dunes community on
association was more apparent as compared with negative the island (Thornton 2007). Furthermore, on a volcanic
association as time progressed. This observation might desert of Mount Fuji, Japan, a dwarf pioneer shrub Salix
support the view that in a severely disturbed habitat where reinii was clumped together and positively associated with
primary succession is occurring, the role of facilitation will the tree seedling Larix kaempferi and has shown its role as
have a stronger role in species change as compared to nurse-plant in primary succession (Endo et al. 2008).

2 CONCLUSION
Table 2. Association tests using chi-squared test statistic ( ) between discriminating
native and invasive pioneer species. This research has demonstrated
that in Mount Merapi primary
Species Paired species Result of chi- Type of Ochiai succession sites, positive
squared test asso- Index interspecific association increased as
ciation
time progressed, which supports
Anaphalis javanica Debregeasia longifolia Associated - 0
already establish view that
Humata repens Associated - 0
facilitation is more prominent in a
Rubus flaxinifolius Associated - 0 severely disturbed habitat as
Athyrium macrocarpum Polygala paniculata Associated + 1 compared to competition. This result
Polygonum chinense Associated - 0.4 could have important value for
Calliandra callothyrsus Crassocephalum crepidioides Associated + 0.84 restoration programs, which could
Cyperus rotundus Associated + 1 concentrate on re-planting
Polygala paniculata Associated + 0.84 subsequent species that have
Panicum reptans Associated - 0.4 positive association with native
Eleusine indica Associated + 0.77 pioneer species, perhaps preferably
Polytoca bracteata Associated + 0.70 focusing on legume species to
Polytrias amaura Associated - 0.4 enhance the barren substrates.
Eupatorium riparium* Gnaphalium japonicum Associated + 0.84
Stachytarpheta jamaicensis Associated - 0 ACKNOWLEDGEMENTS
Melastoma affine Associated + 1
Imperata cylindrica* n.a. Not n.a. n.a.
We would like to thank Dr. Viki
associated
Cramer and Prof. Richard Hobbs
Pinus merkusii Polygala paniculata Associated - 0.28
Shuteria vestita Associated - 0 from the University of Western
Note: Association is significant at 0.05 levels. Values of the Ochiai Index (strength of Australia for insightful discussion,
association) are equal to 0 at no association and to 1 at complete/maximum Soewarno HB from the Faculty of
association. An asterisk (*) indicates an invasive species. Forestry, Gadjah Mada University,
Tri Prasetyo, the head of the Merapi
 SUTOMO et al. Species composition and plants association of Mount Merapi 217

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