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Dynamic Response of a Philippine Dipterocarp Forest to Typhoon Disturbance

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Dynamic Response of a Philippine Dipterocarp Forest to Typhoon Disturbance Journal of Vegetation Science 27 (2016) 133–143 Dynamic response of a Philippine dipterocarp forest to typhoon disturbance Sandra L. Yap, Stuart J. Davies & Richard Condit Keywords Abstract Biomass; Dipterocarp forest; Forest dynamics; Forest resilience; Mortality and recruitment; Questions: Natural hazards can wreak catastrophic damage to forest ecosys- Regeneration; Tree demography; Typhoon tems. Here, the effects of typhoon disturbance on forest structure and demogra- disturbance phy of the 16-ha Palanan Forest Dynamics Plot in the northeast Philippines were examined by comparing census intervals with (1998–2004) and without Nomenclature (2004–2010) a strong typhoon. Category 4 Typhoon Imbudo, with wind gusts Co et al. (2006) exceeding 210 kph, hit Palanan in July 2003. In this study, we ask: (1) was there Received 5 August 2014 an effect of the typhoon on stand structure and biomass; (2) was there an impact Accepted 5 February 2015 on species diversity; (3) did annual mortality, growth and recruitment change Co-ordinating Editor: Kerry Woods significantly between typhoon and non-typhoon periods; and (4) did the typhoon’s impact vary with local topography, from leeward to windward sides Yap, S.L. ([email protected])1, of a ridge? Davies, S.J. (corresponding author, Location: Lowland mixed dipterocarp forest, Palanan, Isabela, Philippines. [email protected])2, Condit, R. ([email protected])3 Methods: Census data from 1998, 2004 and 2010 for all trees ≥1 cm DBH in a 1Institute of Biology, University of the 16-ha permanent plot in Palanan, Isabela, were used to assess tree demography. Philippines, Diliman, Quezon City, PH 1101, Recorded in the census were species identification and measurements of DBH Philippines; and tree locations. Biomass was calculated from published allometry. 2Center for Tropical Forest Science-Forest Global Earth Observatory, Smithsonian Tropical Results: Species diversity and stand structure remained stable, except for an – Research Institute, PO Box 37012, Washington, increase in small-sized trees (1 2 cm) recorded in the census conducted within DC 20013-7012, USA; a year of the typhoon disturbance. Tree mortality was significantly higher during À 3Smithsonian Tropical Research Institute, PO the typhoon interval at 2.27%Áyr 1 and more so in windward than leeward Box 0843-03092, Balboa, Ancon, Panama habitats. Above-ground biomass loss in the typhoon interval was minimal (2.64%) and biomass exceeded pre-typhoon levels after 6 yrs. Recruitment rate during the typhoon interval was almost four times the rate in the non-typhoon interval that followed, attributed to the rapid growth of seeds and seedlings of pioneer species due to the open, defoliated canopy. Negative population growth was recorded for the early successional species in the non-typhoon interval. Sig- nificantly higher growth rates of trees in the non-typhoon interval also con- tributed to biomass gain. Conclusions: Mortality, recruitment and growth rates vary across a heteroge- neous landscape and are related to typhoon disturbances. The relatively low mortality and fast recovery of the Palanan forest demonstrates the resistance and resilience of the forest to intense episodic typhoon disturbances. during a 1938 hurricane (Foster 1988), while in Nicaragua, Introduction Hurricane Joan damaged 75% of trees in the rain forests it Typhoons and hurricanes are among the most severe natu- struck (Boucher et al. 1990). Most such studies on the ral disturbances forests ever face, leading Yih et al. (1991) impact of tropical storms have been done in the Americas, to conclude that “the appropriate metaphor for natural particularly in Puerto Rico, where major hurricanes pass ecosystems is not eternal constancy, but rather cycles of every 50–60 yrs (Zimmerman et al. 1994; Ostertag et al. death and resurrection” after studying Hurricane Joan in 2005). In the tropical western Pacific, typhoons recur Nicaragua. In New England, 70% of trees were felled much more frequently: category-five storms, with winds Journal of Vegetation Science Doi: 10.1111/jvs.12358 © 2015 International Association for Vegetation Science 133 Response of dipterocarp forest to typhoons S.L. Yap et al. >250 kph, pass over Fushan Forest in Taiwan every 12 yrs ity, growth and recruitment; and (3) both of these in rela- on average (Mabry et al. 1998), and in the northeast tion to the varying topography from leeward to windward Philippines, three category-five storms struck in the last sides of a ridge. We predicted that the typhoon disturbance 10 yrs. Yet few studies of typhoon damage on forests have would result in lower tree density across all diameter been done in Asia (Lin et al. 2011), and none in the Philip- classes, reduced species diversity and reduced biomass. We pines. This begs the question of how typhoons impact also expected that annual mortality would be higher in the Asian forests: could 70% of the trees in a Philippine forest typhoon interval, with growth and recruitment rates be toppled every decade? higher in the non-typhoon interval that followed. More- Dense and tall dipterocarp forests dominate the land- over, we anticipated increasing impacts from leeward to scape throughout the Malesian region (Maury-Lechon & windward habitat types. Curtet 1998). In the Philippines, these forests are special because local endemism is high. Philippine forests harbour Methods >9000 plant species, 50% of which are endemic (Merrill Study site 1926; Myers et al. 2000). Unfortunately, only 7% of the old-growth closed-canopy forests remain (Heaney & The research was conducted in a mixed dipterocarp forest Regalado 1998). Severe damage to these remnants during of the Northern Sierra Madre Natural Park in Isabela, typhoons could thus put species at risk of extinction. In Philippines, on the northeast coast of Luzon Island, in the order to quantify typhoon damage, we established a per- foothills of the Sierra Madre Mountains (Fig. 1). The area manent plot in the Palanan forest of northeast Luzon is characterized by an ever-wet climate with mean annual Island in 1994. With a powerful typhoon in 2003, we now rainfall of 3218 mm, with individual years ranging from have complete censuses of 100 000 trees during a 1347 to 6841 mm (Co et al. 2006). From Nov to Jun, typhoon-impacted interval (1998–2004) and a non- northeast winds off the Pacific bring rain to the eastern side typhoon interval (2004–2010), providing a comparative of the Sierra Madre, and during the other half of the year, measure of typhoon impact on forest dynamics. We inves- tropical cyclones approach from the southeast. Between tigated the effects of typhoon disturbance on: (1) stand 1948 and 2004, nine typhoons made landfall in the Philip- structure, species diversity and biomass; (2) annual mortal- pines per year (pagasa.dost.gov.ph). Among the many that Fig. 1. Map of the 16-ha Palanan FDP showing its topography, habitats and position along typhoon tracks. The Palanan FDP is located in the Pacific coast of Luzon Island exposed to the direction of most typhoons. The five habitat types represented are leestream, windstream, ridge, leeslope and windslope. Journal of Vegetation Science 134 Doi: 10.1111/jvs.12358 © 2015 International Association for Vegetation Science S.L. Yap et al. Response of dipterocarp forest to typhoons visited the area, Category-four Typhoon Imbudo (local west of the ridge the leeward, unexposed side. Directly name: Harurot) hit the Pacific coast of northeast Luzon on adjacent to ridge habitats, on both windward and leeward 22 July 2003, with maximum sustained winds of 210 kph sides, are quadrats sloping down to the streams bisecting (pagasa.dost.gov.ph). It was reported by local communities the plot, ranging from <104.4 m a.s.l. to 77 m a.s.l. (26-m to have inflicted extensive damage in and around Palanan, range), correspondingly referred to as windslope and lee- Isabela. slope habitats. Similarly, streamside quadrats are referred to as windstream and leestream habitats on the east and west of the ridge, respectively. Streams are recognized as The tree census distinct habitats as other studies have shown species–habi- A permanent, 16-ha forest census plot was established at tat associations with streamside niches (Harms et al. 2001; 17°02036 N, 122°22058 E in 1994 and named the Palanan Chuyong et al. 2011). Forest Dynamics Plot (Co et al. 2006). The Palanan Plot is a member of the network of long-term plots of the Center Taxonomy for Tropical Forest Science (CTFS, www.ctfs.si.edu), and plot establishment followed the standard CTFS protocol Every individual not identified immediately in the field by (Condit 1998). Work began in 1994, when 8 ha of forest an experienced botanist (James V. LaFrankie or Leonard L. were censused; in 1998, the plot was extended to 16 ha. Co) had leaves with or without flowers collected. The Re-censuses of the 16 ha were completed in 2004 and unknown specimens were sorted into groups thought to 2010. At each census, all trees ≥1 cm DBH were tagged, be single species, then compared to specimens at the measured, mapped and identified to species. Here we University of the Philippines Herbarium (PUH), Philippine report on the three full censuses of 16 ha, providing esti- National Herbarium (PNH) and Harvard University Her- mates of forest demography and change over 1998–2004 barium (HUH). Herbarium vouchers for each morphos- and 2004–2010. With Typhoon Imbudo passing over the pecies are currently stored at the PUH and Isabela State plot in July 2003, the former interval included the typhoon University. Those still not identified were given morphos- impact while the latter did not. The census of 2004 began pecies names, and demographic rates of those morphos- in Dec 2003, and was finished by Jun 2004, within pecies were calculated just as for the rest of the species 12 months of Imbudo passing over.
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