Philippine Journal of Science 150 (5): 1121-1129, October 2021 ISSN 0031 - 7683 Date Received: 08 May 2020

Kandelia candel (L.) Druce, a True Native Species in the Philippines

Pastor L. Malabrigo Jr.1,2,4, Gerald T. Eduarte3,4, Laarni D. Malabrigo4, Ericson E. Coracero5

1Department of Forest Biological Sciences, College of Forestry and Natural Resources 2Museum of Natural History; 3Graduate School University of the Philippines Los Baños, Laguna 4031 Philippines 4Pro-Seeds Development Association Inc. Batong Malake, Los Baños, Laguna 4031 Philippines 5Department of Forestry and Environmental Sciences Aurora State College of Technology (Baler Campus), Aurora, Philippines

Kandelia candel (L.) Druce is a true species under the family . It is distributed throughout southeast Asia to south China, the Ryukyu Island and southern Japan, Taiwan, and Hong Kong. In the Philippines, its only known area of occurrence is the province of Aurora, the central easternmost coast of Luzon island. However, there has been a long-standing debate on the presence of natural populations of Kandelia candel in the Philippines. Adding to the confusion, a number of molecular studies reported that the Kandelia populations from north of the South China Sea (SCS) are a genetically different species – – making the non-monotypic. Some mangrove biologists hypothesized that Kandelia populations in Aurora might have been introduced to the province by some fishermen from Taiwan. On the other hand, some papers suggest that Kandelia populations in the Philippines were shaped by the founders from the populations in Southern SCS. A recent population inventory conducted by the authors, complemented by taxonomic characterization, strongly suggests that Kandelia candel is native in the province of Aurora, Philippines.

Keywords: Kandelia candel, Kandelia obovata, Rhizophoraceae

INTRODUCTION Kong (Giang et al. 2006). As compared to other mangrove species, K. candel has limited global distribution and Kandelia candel is a true mangrove species that is throughout its distributional range, the species was known to produce viviparous propagules (denoting the considered to be nowhere abundant (Tomlinson 1986). special ability of seeds to germinate inside the fruit while still attached to the parent tree) that are buoyant In the Philippines, it is surprising that the species and presumably capable of long-distance dispersal distribution is restricted in the province of Aurora, the (Chiang et al. 2001). The known distribution of Kandelia central easternmost coast, despite its viviparous nature. candel includes the Ganges delta, western , Burma Rotaquio et al. (2006) in their study of the (Myanmar) through southeast Asia to south China, the of Aurora observed that K. candel populations are Ryukyu Island and southern Japan, Taiwan, and Hong present only in small stands along the muddy river of *Corresponding Author: [email protected] the municipalities of Baler and Casiguran. Given this

1121 Philippine Journal of Science Malabrigo et al.: Kandelia candel (L.) Druce, a True Vol. 150 No. 5, October 2021 Native Species in the Philippines very restricted distribution and small population sizes, This paper presents the results of the population the species is categorized as critically endangered in the distribution study, including a 100% population inventory Philippine red list for (DENR 2017). of K. candel in the province of Aurora. It provides new information on the spatial distribution, dominance, and Despite a number of publications (Primavera 2000; stand structure of the species population, which could Primavera et al. 2004; Rotaquio et al. 2006, 2007; Garcia give a better understanding of the naturality of the species et al. 2013) documenting the occurrence of the population in the Philippines. A taxonomic characterization and of the species in the country, the presence of natural nomenclature review of the species was also conducted to populations of Kandelia candel in the Philippines is still complement the findings of the population survey. unrecognized in the international scientific community (Chiang et al. 2001; Rotaquio et al. 2007). The paper of Hou (1958), which claimed no natural populations of K. candel in the Philippines, became the baseline reference MATERIALS AND METHODS that was transferred from generation to generation and still being adopted by other contemporary mangrove ecologists, i.e. Sheue et al. (2003), Chiang et al. (2001), Maxwell Population Inventory et al. (1997). Sheue et al. (2003) even hypothesized that A 100% inventory of all mature individuals was conducted Kandelia populations in Aurora might have been introduced from 12–14 Jul 2019, in known populations of Kandelia to the province by some fishermen from Taiwan. On the candel in Aurora, particularly in the municipalities of other hand, both papers of Tomlinson (1986) and Huang Baler and Casiguran (Figure 1). Matured individuals and Chen (2000) – as cited by Chiang et al. (2001) – state considered in this study were those with a diameter of that during summer, the propagules of K. candel are ≥ 3 cm measured at 1 ft above ground (diameter at foot dispersed from the Indian Ocean to the SCS and then height or dfh) or 10 cm above the highest buttress. Based through the Bashi Strait (the strait between Taiwan and on earlier observations, K. candel is a precocious species the Philippines) into the Pacific. It, therefore, indicates that that start bearing flowers and fruits as early as it reaches 3 Kandelia populations in the Philippines were shaped by the cm dfh. The use of dfh instead of diameter at breast height founders from the populations in southern SCS and should, or dbh is extremely rational since K. candel could start therefore, retain the name K. candel. Unfortunately, except flowering before reaching the height of 1.3 m. Total height for the mangrove diversity assessment study in Aurora by and dfh were measured to account for the stand structure Rotaquio et al. (2006), there has been no population study and dominance of the species. GPS coordinates of each on Kandelia candel in the country. individual were also noted to aid in the construction of an interactive distribution map for future assessment and Kandelia has long been recognized as a monotypic genus, monitoring studies. with Kandelia candel as the only species. Yet in 2003, Sheue et al. found significant morphological as well as genetic differentiation between the Kandelia populations Taxonomic Characterization from the northern part of SCS (NSCS) and populations Following the key characters used by Sheue et al. (2003) from the southern part of SCS (SSCS). They proposed that in comparing Kandelia candel vs. Kandelia obovata, the populations in each side of SCS be recognized as two specimens of Philippine Kandelia were collected and distinct species. They reported a new species, Kandelia characterized. A total of 20 herbarium specimens were obovata, and further suggested that the name be adopted deposited, 10 each at LBC and CAHUP (Malabrigo for the populations from the NSCS. This treatment was 590–597, Coracero 001–002). For the leaf characters (leaf further supported by more recent molecular studies (Giang shape/dimension and number of lateral veins), 300 leaf et al. 2006; Kado et al. 2006). Unfortunately, not one of samples collected from 30 individual trees (10 each) were these studies contains Philippine materials. observed and measured. Thirty (30) individual flowers in bloom from 10 individual trees were sampled and In 2006, the study of Rotaquio et al. reported significant characterized. Three hundred (300) matured fruits from 10 morphological differences between Kandelia populations fruiting trees were measured to determine the length of the from Aurora, Philippines with either the NSCS or SSCS hypocotyl. And lastly, the height measurement from the populations. It seems to suggest that Philippine Kandelia 100% inventory was used as the basis for the maximum is neither K. candel nor K. obovata. However, it should height of the tree. Samples were collected in such a way be cautiously noted that the morphological characters that both subpopulations in Zabali and Castillo were used by Rotaquio et al. are measurements (length and equally represented. Results of taxonomic characterization width) of leaves and fruits. These characters are expected were then compared with the two known species to to vary within species living in different geographical determine the identity of the Philippine Kandelia. environments.

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Figure 1. Locations of Kandelia candel populations in the Province of Aurora, Philippines.

RESULTS species) along the Casiguran river through a government reforestation project. The project contractor confirmed that the seedlings came from the four matured individuals Population in Casiguran, Aurora and were planted 3 yr ago. Accordingly, compared to other Available literature (Rotaquio et al. 2006, 2007; Pelser et al. mangrove species (Rhizophora apiculata, R. mucronata, 2011 onwards) consistently mentioned the municipalities R. stylosa, Avicennia marina) used/planted, K. candel of Baler and Casiguran as the native habitat of K. candel in had the highest mortality and the slowest growth. The the Philippines. A recent population survey in the province average height of the surviving K. candel seedlings was of Aurora confirmed that those few individuals growing only 58.5 cm. in Brgy. Cozo in Casiguran Bay were planted from the mother trees from Baler. The local guide who assisted the authors during the survey was the same person who Population in Baler, Aurora first planted the trees as early as 1990. Accordingly, he The first record of the occurrence of Kandelia candel in the initially planted more than 200 individuals and less than Philippines was from the municipality of Baler, Aurora. 50 trees survived. There were larger trees before that were Hence, the species is commonly known as “bakauan already cut. Only four matured living individuals from Baler.” The recent survey documented two subpopulations Casiguran, with a diameter ranging from 6–8 cm (Plate of the species in the municipality: one along the main river 1) were recorded, all of which were at the late flowering tributary in Baler known as Castillo river and another one or early fruiting stage during the time of visit. from Zabali river, a smaller inner tributary that also drains to the main river (see Figure 1). Apart from the matured individuals, a total of 148 naturally growing seedlings were recorded just under or It is important to note that both subpopulations in Baler near the vicinity of the mother trees. There were also eight were restricted along the riverbank with an average width Kandelia candel seedlings planted (with other mangrove of 5 m (Plate 2). Further, no trees were found near the

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Plate 1. Artificial population of Kandelia candel in Casiguran Aurora: a) matured individuals documented along Casiguran river; b) flowers at their early fruiting stage showing the developing fruit buds; c) one of the surviving seedlings from a government reforestation project.

Plate 2. The two subpopulations of Kandelia candel in Baler, Aurora: a) part of the Zabali population located further inland (820 m from the river mouth); b) portion of Castillo population along the main river tributary in Baler (320 m from the river opening).

1124 Philippine Journal of Science Malabrigo et al.: Kandelia candel (L.) Druce, a True Vol. 150 No. 5, October 2021 Native Species in the Philippines mouth of the river or the seaward portions. Population in Table 2. Number of Kandelia individuals per diameter class in Baler, Castillo river stretched from 320–650 m away from the Aurora, Philippines. river mouth while Zabali population is a lot further inland Diameter class (cm) Zabali Castillo Across sites at 820–1,000 m from the river opening. 3.0–5.0 89 890 979 The population in Zabali had a smaller area as compared 5.01–10.0 67 38 105 2 with Castillo, having a 180-m stretch (≈ 900 m ). But > 10 30 0 30 Zabali seemed to be the older population as it comprised larger and taller trees. A total of 186 matured individuals Total 186 928 1114 was recorded in Zabali, with an average diameter and height of 6.88 cm and 5.19 m, respectively (Table 1). Thirty (30) individuals were found to have a diameter of Morphological Characterization more than 10 cm (Table 2). The largest tree recorded had a Comparison of key morphological characters between diameter of 17 cm while the tallest tree has a height of 9 m. the two described species and the Philippine Kandelia The Castillo population had a longer stretch of about 330 revealed that the species in Aurora belongs to K. candel m (≈ 1,650 m2). It had a significantly higher density with (Table 3). The study suggests that blade characters, as well 928 matured individuals. However, the Castillo population as sepal characters, are not the best traits in separating was relatively younger with an average diameter and the two species because high variability was observed height of 3.63 cm and 3.34 m, respectively. For both even within individuals. In addition, the measurements of subpopulations, the majority of the trees belonged to the blade and sepal for the two species have a wide overlap. lower diameter class (3.0–5.0 cm) It was observed that the abaxial color of the sepals varies from light green (early bloom) to yellowish-white (late bloom). However, the rest of the parameters are more Table 1. Summary of inventory of Kandelia in Baler, Aurora, reliable that could easily distinguish one species from Philippines. the other. The number of lateral veins of Philippine Parameter Zabali Castillo Across sites Kandelia consistently showed eight or more veins for both populations. The average length of the hypocotyl (31 No. of individuals 186 928 1114 cm) is significantly longer than the length range recorded Average diameter (cm) 6.88 3.63 4.18 for K. obovata. The survey also noted that Philippine Maximum diameter (cm) 17 8 17 Kandelia is a true mangrove tree that could attain a height Average height (m) 5.19 3.34 3.57 of 9 m. This is far beyond the recorded height for K. Maximum height (m) 9 6 9 obovata, which is only 3 m, suggesting that the species is only a shrub. The matrix of flower comparison adopted

Table 3. Comparison of key morphological characters between Kandelia candel, K. obovata (extracted from Sheue et al. 2003), and the Philippine Kandelia (this study). Species K. candel K. obovata Philippine Kandelia Elliptic-oblong to oblong-lanceolate, Obovate to obovate-elliptic, very rarely Elliptic-oblong to oblong-lanceolate, 9.2 Leaf 6-16 cm x 3–6 cm obovate-oblong, 6–12 cm x 2.5–5 cm cm (4.3–13) x 3.1 cm (2.0–4.6) Lateral veins Mostly 8–11 (–13) pairs Mostly 5–8 pairs 11 (8–14) pairs Light green (early bloom) to yellowish Light green (abaxial side) when White (abaxial side) when blooming, Sepal white (late bloom) (abaxial side), 1.5 cm blooming, 1.4–1.6 cm x 1.9–2.1 mm 1.5–1.9 cm x 2.5–3.0 mm (1.2–1.7) x 2.2 mm (1.8–2.4) Deeply bifid and each half with 3–5 Partially bifid and each half with 7–8 Deeply bifid and each half with 4 threads Petal threads or bristles threads or bristles or bristles Style Distinctly shorter than the stamens Longer or subequal to stamens Distinctly shorter than the stamens Bracteole V-shaped U-shaped V-shaped Anther Curved and grooved Cone-shaped without distinct groove Curved and grooved 31 cm long (18–42), 0.8 cm (0.6–1.0) 20–40 cm long, 0.7–0.9 cm at the 15–20 (–23) cm long, 0.9–1.4 cm at the Hypocotyl at the broadest part; apex attenuate to broadest part; apex attenuate broadest part; apex acuminate acuminate Tree height Up to 7 m tall Up to 3 m tall Up to 9 m tall

1125 Philippine Journal of Science Malabrigo et al.: Kandelia candel (L.) Druce, a True Vol. 150 No. 5, October 2021 Native Species in the Philippines from the original publication of K. obovata (Sheue et al. and grooved. Comparison of actual photos of Philippine 2003) further supports the identity of Kandelia in Baler Kandelia with Kandelia obovata from a plantation in as K. candel (Plate 3). The style is distinctly shorter than Quang Ninh province, Northern Vietnam also confirms a the stamens. The petal is bifid with four threads in each lot of dissimilarities (Plate 4). half. The bracteole is V-shaped and the anther is curved

Plate 3. Matrix of comparison between Kandelia of Baler and K. obovata (adopted from Sheue et al. (2003): A) style (red arrow) distinctly shorter than the stamens in Baler Kandelia; B) style longer or subequal to stamens in K. obovata; C) petal deeply bifid and each half with four threads in Baler Kandelia; D) petal partially bifid and each half with 7–8 threads; E) V-shaped bracteole in Baler Kandelia; F) U-shaped bracteole in K. obovata; G) anther curved and grooved in Baler Kandelia; H) anther cone-shaped without distinct groove in K. obovata. Scale bars = 2 mm.

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Plate 4. Actual photos of Philippine Kandelia (a–c) from natural population in Baler, Aurora and K. obovata (d–f) from a 3-yr-old plantation in Quang Ninh province, northwest Vietnam: a) the largest tree recorded with diameter of 17 cm and height of 9 m; b) flowers (early bloom) and buds showing the light green sepals; c) fruiting twig displaying the long fruits/hypocotyl; d) the longest fruit measured in the plantation; e) a 1.2 m tall K. obovata already producing fruits, demonstrating early maturity; f) matured fruiting twig showing the short obovate leaves.

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DISCUSSION to Kwangtung, Fukien, Taiwan, the Ryukyus, and southern Japan) was followed by a generalization that The occurrence of Kandelia candel in the province of all populations from northern SCS belong to K. obovata. Aurora, Philippines was first recorded in 1996, when a Kado et al. (2006) even suggested that the border of the two mangrove specialist working as a consultant of the then Kandelia species is located between Đồng Rui (northern Aurora Integrated Area Development project discovered Vietnam) and Cần Giờ (southern Vietnam). Following this the species in Baler and Casiguran Bay (Rotaquio et al. generalization, then those Kandelia populations in central 2007). Since then, no other sighting of the species in other Vietnam and those located in between the two provinces, mangrove areas in the Philippines has been reported. To like Aurora in the Philippines, are in limbo. Hence, this date, not a single population study has been conducted generalization needs to be substantiated by population on this extremely rare mangrove species in the country. studies of all existing populations in the region. The only relevant studies are those from Rotaquio et al. (2006, 2007), which provided diversity assessment and The morphological comparison made in this study using general characterization of the mangrove forests in Aurora. the key characters developed by Sheue et al. (2003) Both papers consistently mentioned “few” individuals in strongly suggests that Philippine populations belong to Casiguran Bay. Results of the study revealed that there is the species Kandelia candel. It, therefore, dismissed the no natural population of Kandelia in Casiguran, Aurora. hypothesis that Kandelia populations in the Philippines There were only four mature individuals in Brgy. Cozo, were introduced by Taiwanese fishermen since Taiwan Casiguran, which were artificially planted from mother populations were confirmed to be Kandelia obovata. trees in Baler. The overdominance – both in sizes and abundance (1,114 matured individuals) – of Kandelia candel in Baler, The population distribution study, including the 100% Aurora, suggests that the species is native in the area. inventory, provided important baseline information on Consequently, it implies that Kandelia candel populations the abundance and stand structure of the species to better in the Philippines were shaped by the founders from the understand its population status in Aurora. The two populations in southern SCS. A more comprehensive subpopulations documented in Baler were both located phylogenetic study, including population genetics, of along the river at least 320 m away from the river mouth. Kandelia candel in the Philippines, could provide a This conforms with previous observations from other better understanding of the true phylogenetic origin of populations (Parida and Jha 2010; Popp et al. 1993) that the Philippine population. the species is an interior riverine mangrove that indicates its intolerance to high salinity. The Zabali population is the landward most part of the mangrove forest in the area and almost completely dominated by K. candel with a CONCLUSION few scattered Nypa fruticans in between. Non-mangrove vegetation immediately followed going landward. In The population study including the 100% tree inventory of the larger population along the Castillo river, almost a all Kandelia trees in the province of Aurora, Philippines pure stand of K. candel serves as a river border while revealed new information necessary to better understand Sonneratia alba and Nypa fruticans dominate at the back. the identity and naturality status of the species. It was It was confirmed that Philippine Kandelia is a major confirmed that there is no natural population of Kandelia mangrove species capable of forming pure, dense stand. in Casiguran Bay, and the few individuals growing While both populations have a relatively small extent, in the area were all planted from the mother trees of Kandelia overdominates its areas of occurrence, almost Baler. A more complete morphological characterization totally excluding other mangrove species to grow. of Philippine Kandelia was also presented, and a comparison with two known Kandelia species suggests There were also new morphological findings of Kandelia it belongs to K. candel. With the confirmation of its that were obviously missed by some characterizations identity, being different from that of Taiwan, it eliminated made by mangrove biologists in the Philippines. Firstly, the long-standing theory that Kandelia in the Philippines the species produce large buttresses for better anchorage was introduced from Taiwan. The stand structure and in waterlogged areas, unlike Rhizophora spp. that over dominance of Kandelia candel in Baler strongly develop stilt roots. And secondly, Philippine Kandelia is suggests naturality of the species in the area and, unquestionably a tree capable of attaining a diameter of therefore, conform with the papers of Tomlinson (1986) 17 cm and height of 9 m. and Huang and Chen (2000) that Kandelia populations The publication of Sheue et al. (2003) of the new species, in the Philippines were shaped by the founders from the Kandelia obovata, from populations located on the populations in southern SCS. northern part of SCS (i.e. Gulf of Tonkin northeastward

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ACKNOWLEDGMENTS KADO T, FUJIMOTO A, GIANG LH, TUAN M, HONG PN, HARADA K. 2006. Genetic structures of natural The authors are grateful to the Provincial Environment populations of three mangrove species, Avicennia and Natural Resources Office of Aurora for allowing the marina, Kandelia candel, and Lumnitzera racemosa, authors to conduct the study in the mangroves of Baler in Vietnam revealed by maturase sequences of plastid and Casiguran. Special thanks go to Foresters Manny Dela DNA. Species Biology 19(2): 91–99. Peña and Ronald Samonte of the Municipal Environment and Natural Resources Office of Baler for their invaluable MAXWELL GS, HAVANOND S, NAKAMURA T. 1997. assistance during the conduct of fieldworks. Sincere The ecogeography of Kandelia candel in Thailand, gratitude also goes to the Department of Forestry and Hongkong and Southern Islands of Japan. Biol Bull Environmental Sciences of the Aurora State College of NTNU 32(2): 89–95. Technology for generously allowing us to use the FBS PARIDA AK, JHA B. 2010. Salt tolerance mechanisms Laboratory, which allowed us to complete the survey and in mangroves: a review. Trees-Structure and Function characterization of plant specimens. Most importantly, the 24: 199–217. senior author is thankful to the Department of Science and Technology–Philippine Council for Agriculture, Aquatic, PELSER PB, BARCELONA JF, NICKRENT DL eds. and Natural Resources Research and Development for 2011 onwards. Co’s Digital Flora of the Philippines. providing the research grant to complete the project. Retrieved on 30 Apr 2020 from www.philippineplants. org POPP M, POLANIA J, WEIPER M. 1993. Physiological adaptations to different salinity levels in mangrove. REFERENCES Towards the rational use of high salinity tolerant plants. CHIANG TY, CHIANG YC, CHEN YJ, CHOU CH, Dordrecht: Kluwer Academic Publishers. HAVANOND S, HONG TN, HUANG S. 2001. PRIMAVERA JH. 2000. Development and conservation Phylogeography of Kandelia candel in East Asiatic of Philippine mangroves: institutional issues. Elsevier mangroves based on nucleotide variation of chloroplast Ecological Economics 35: 91–106. and mitochondrial DNAs. Molecular Ecology 10: 2697–2710. PRIMAVERA JH, SADABA RB, LEBATA MJH, ALTAMIRANO JP. 2004. Handbook of Mangroves [DENR] Department of Environment and Natural in the Philippines: Panay. Southeast Asian Fisheries Resources. 2017. Administrative Order No. 2017-11. Development Center (SEAFDEC), Philippines. Updated National List of Threatened Philippine Plants and Their Categories. Retrieved on 30 Apr 2020 from ROTAQUIO EL, NAKAGOSHI N, ROTAQUIO RL. http://www.bmb.gov.ph/index.php/e-library/laws-and- 2007. Species Composition of Mangrove Forests policies/denr-administrative-orders/dao-2017-2019 in Aurora, Philippines – A Special Reference to the Presence of Kandelia candel (L.) Druce. Journal of GARCIA KB, MALABRIGO PL, GEVAÑA DT. 2013. International Development and Cooperation 13(1): Philippines’ Mangrove Ecosystem: Status, Threats 61–78. and Conservation. In: Mangrove Ecosystems in Asia: Status, Challenges, and Management Strategies. ROTAQUIO EL, NAKAGOSHI N, ROTAQUIO RL. Faridah-Hanum I, Latiff A, Hakeem KR, Ozturk M 2006. Kandelia candel (L.) Druce – a mangrove eds. Springer-Verlag New York Inc. p. 81–94. population recorded in the Philippines and its leaf and fruit variations among other populations recorded GIANG LH, GEADA GL, HONG PN, TUAN MS, LIEN in Thailand, Brunei, Hong Kong and Japan. Hikobia NTH, IKEDA S, HARADA K. 2006. Genetic variation 14(4): 483–491. of two mangrove species in Kandelia (Rhizophoraceae) in Vietnam and surrounding area revealed by SHEUE CR, LIU HY, YONG JWH. 2003. Kandelia microsatellite markers. Int J Plant Sci 167: 291–298. obovata (Rhizophoraceae), a new mangrove species from Eastern Asia. Taxon 52: 287–294. HOU D. 1958. Rhizophoraceae. In: Flora Malesiana, Series I, Vol. 5. van Steenis CGGJ ed. p. 429–493. TOMLINSON PB. 1986. The botany of mangroves. Cambridge, MA: Cambridge University Press. HUANG S, CHEN YC. 2000. Patterns of genetic variation of Kandelia candel among populations around South China Sea. In: Wetland Biodiversity: Proceedings of Symposium of Biodiversity in Wetlands. Taiwan Endemic Species Research Institute, Nantou, Taiwan. p. 59–64.

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