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Colonisation of orchids on the Krakatau Islands - Royal Botanic ...

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Partomihardjo, <str<strong>on</strong>g>Col<strong>on</strong>isati<strong>on</strong></str<strong>on</strong>g> <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>orchids</str<strong>on</strong>g> <strong>on</strong> <strong>the</strong> <strong>Krakatau</strong> <strong>Islands</strong> 305<br />

A means <str<strong>on</strong>g>of</str<strong>on</strong>g> estimating forest maturity is based <strong>on</strong> epiphytes, mostly c<strong>on</strong>sisting <str<strong>on</strong>g>of</str<strong>on</strong>g><br />

<str<strong>on</strong>g>orchids</str<strong>on</strong>g>. It has been suggested that in well-established forests epiphytic species<br />

(mostly <str<strong>on</strong>g>orchids</str<strong>on</strong>g>) are abundant, and that as <strong>the</strong> forest matures epiphytic species<br />

become proporti<strong>on</strong>ally more abundant, particularly <strong>on</strong> trees <str<strong>on</strong>g>of</str<strong>on</strong>g> > 30 cm dbh (Benzing<br />

1981). In mature dipterocarp forests at Wanariset, East Kalimantan, 50% <str<strong>on</strong>g>of</str<strong>on</strong>g> <strong>the</strong><br />

epiphytes were <str<strong>on</strong>g>orchids</str<strong>on</strong>g> (Partomihardjo 1995). It has been menti<strong>on</strong>ed that <strong>the</strong><br />

epiphytic <str<strong>on</strong>g>orchids</str<strong>on</strong>g> have a wide distributi<strong>on</strong> within <strong>the</strong> trees. For example, epiphytic<br />

<str<strong>on</strong>g>orchids</str<strong>on</strong>g> can be found to grow <strong>on</strong> <strong>the</strong> terminal branches <str<strong>on</strong>g>of</str<strong>on</strong>g> <strong>the</strong> canopy layer to <strong>the</strong> very<br />

base <str<strong>on</strong>g>of</str<strong>on</strong>g> large trees. Interestingly, <strong>the</strong> rate <str<strong>on</strong>g>of</str<strong>on</strong>g> increasing <str<strong>on</strong>g>of</str<strong>on</strong>g> epiphytic and terrestrial<br />

<str<strong>on</strong>g>orchids</str<strong>on</strong>g> is somewhat anomalous.<br />

Dispersal mechanisms and limits to col<strong>on</strong>isati<strong>on</strong>: The cumulative survey data<br />

<str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>orchids</str<strong>on</strong>g> from 1883 to 1998 shows 24 to 41 species <str<strong>on</strong>g>of</str<strong>on</strong>g> terrestrial and epiphytic<br />

respectively. On <strong>Krakatau</strong>, <str<strong>on</strong>g>orchids</str<strong>on</strong>g> are assumed to have col<strong>on</strong>ised through wind<br />

dispersal (anemochorous). Dressler (1981) suggested that most orchid seeds are wellsuited<br />

for wind dispersal. Measurements <str<strong>on</strong>g>of</str<strong>on</strong>g> <strong>the</strong> air space inside seeds in <strong>the</strong> laboratory<br />

and <strong>the</strong> time needed for <strong>the</strong>m to fall 1.5 m in a glass tube, indicate that l<strong>on</strong>g-distance<br />

travel by air is clearly possible for orchid seeds (Arditti 1992). Air transport <str<strong>on</strong>g>of</str<strong>on</strong>g> orchid<br />

seeds <strong>on</strong> <strong>the</strong> fea<strong>the</strong>rs or legs <str<strong>on</strong>g>of</str<strong>on</strong>g> birds has been suggested by Went (1990) and Ridley<br />

(1930), although Arditti (1992) argued that <strong>the</strong> chance is very small. This evidence is<br />

supported by <strong>the</strong> fact that large numbers <str<strong>on</strong>g>of</str<strong>on</strong>g> comm<strong>on</strong> species <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>orchids</str<strong>on</strong>g> have no<br />

correlati<strong>on</strong> with <strong>the</strong> bird migrati<strong>on</strong> roots. Wind dispersal seems <strong>the</strong> <strong>on</strong>ly mode <str<strong>on</strong>g>of</str<strong>on</strong>g><br />

orchid dispersal, because all species recorded <strong>on</strong> <strong>the</strong> <strong>Krakatau</strong>s have capsules that<br />

open and release <strong>the</strong> seeds free to float <strong>on</strong> <strong>the</strong> air. Dispersal by sea as suggested by<br />

Ridley (1930) seems very unlikely.<br />

Most <str<strong>on</strong>g>orchids</str<strong>on</strong>g> are very sensitive to altitude, which really means temperature and<br />

moisture regimes (Comber 1990). Because <strong>the</strong> mountain <strong>on</strong> <strong>Krakatau</strong> is relatively low,<br />

<strong>the</strong>re is <strong>on</strong>ly a narrow range <str<strong>on</strong>g>of</str<strong>on</strong>g> temperatures available to suit <strong>the</strong> needs <str<strong>on</strong>g>of</str<strong>on</strong>g> many<br />

<str<strong>on</strong>g>orchids</str<strong>on</strong>g>. Moisture c<strong>on</strong>densati<strong>on</strong> in <strong>the</strong> form <str<strong>on</strong>g>of</str<strong>on</strong>g> rain or mist <strong>on</strong> <strong>the</strong> highest point <str<strong>on</strong>g>of</str<strong>on</strong>g><br />

Rakata is much greater than at <strong>the</strong> lower altitudes, because this island is surrounded<br />

by sea, and it is c<strong>on</strong>tinuously swept by moist winds which cool as <strong>the</strong>y rise and release<br />

<strong>the</strong>ir rain water <strong>on</strong> <strong>the</strong> summit area. So it has been found that most <str<strong>on</strong>g>of</str<strong>on</strong>g> <strong>the</strong> <str<strong>on</strong>g>orchids</str<strong>on</strong>g><br />

recorded in <strong>Krakatau</strong> grow exclusively <strong>on</strong> <strong>the</strong> trunks and branches <str<strong>on</strong>g>of</str<strong>on</strong>g> trees at high<br />

altitude, where <strong>the</strong>y <strong>on</strong>ly have a few ferns as competitors.<br />

Sebesi is <strong>the</strong> closest island to <strong>the</strong> <strong>Krakatau</strong> group and is <str<strong>on</strong>g>of</str<strong>on</strong>g> similar altitude to Rakata.<br />

Sebesi was less badly damaged by <strong>the</strong> erupti<strong>on</strong>s <str<strong>on</strong>g>of</str<strong>on</strong>g> 1883. As <str<strong>on</strong>g>of</str<strong>on</strong>g> 1921 its inland forests<br />

were in a more advanced state <str<strong>on</strong>g>of</str<strong>on</strong>g> recovery than <strong>on</strong> <strong>Krakatau</strong>, and possessed a larger<br />

number <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>orchids</str<strong>on</strong>g> (Docters van Leeuwen 1936). At least nine o<strong>the</strong>r species occurring<br />

<strong>the</strong>re have not yet been recorded for <strong>Krakatau</strong>. This simple comparis<strong>on</strong> suggests that<br />

fur<strong>the</strong>r additi<strong>on</strong>s to <strong>the</strong> <strong>Krakatau</strong> orchid assemblage can be expected.<br />

Altitudinal distributi<strong>on</strong> and species richness <str<strong>on</strong>g>of</str<strong>on</strong>g> epiphytic <str<strong>on</strong>g>orchids</str<strong>on</strong>g>: At higher<br />

altitudes <strong>on</strong> Rakata a greater diversity <str<strong>on</strong>g>of</str<strong>on</strong>g> epiphytic <str<strong>on</strong>g>orchids</str<strong>on</strong>g> is present than in lower<br />

areas. To investigate <strong>the</strong> altitudinal variati<strong>on</strong> <str<strong>on</strong>g>of</str<strong>on</strong>g> <strong>the</strong> orchid species, plots were set up<br />

at 10, 100, 300, 500 and 700 m a.s.l. (Table 3). The total number <str<strong>on</strong>g>of</str<strong>on</strong>g> species counted<br />

within each plot were 2, 2, 7, 9 and 14 species respectively. The populati<strong>on</strong> was<br />

highest in <strong>the</strong> 500 m plot, with <strong>the</strong> number <str<strong>on</strong>g>of</str<strong>on</strong>g> individuals declining to 50.75 per<br />

100 m2 at <strong>the</strong> summit. In <strong>the</strong> 300 m plot <strong>the</strong>re were <strong>on</strong>ly 8.5 individuals per 100 m2 .<br />

An altitudinal preference was evident in some species <str<strong>on</strong>g>of</str<strong>on</strong>g> <str<strong>on</strong>g>orchids</str<strong>on</strong>g>. The epiphytic<br />

species Appendicula reflexa and Pholidota articulata were recorded in <strong>the</strong> highest plot<br />

<strong>on</strong>ly. The comm<strong>on</strong> species Dendrobium crumenatum tends to be more abundant in <strong>the</strong><br />

lower areas, while D. mutabile grows at higher altitude.

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