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Sarawak Forestry Corporation, 2018) Faculty of Resource Science and Technology Historical Perspective, Distribution, Ecology and Population Genetics of Saltwater Crocodile (Crocodylus porosus Schneider, 1801) in Sarawak, Malaysian Borneo Mohd Izwan Zulaini bin Abdul Gani Doctor of Philosophy 2019 Historical Perspective, Distribution, Ecology and Population Genetics of Saltwater Crocodile (Crocodylus porosus Schneider, 1801) in Sarawak, Malaysian Borneo Mohd Izwan Zulaini bin Abdul Gani A thesis submitted In fulfillment of the requirements for the degree of Doctor of Philosophy (Zoology) Faculty of Resource Science and Technology UNIVERSITI MALAYSIA SARAWAK 2019 DECLARATION I hereby declare that the thesis has not been accepted for any degree and is not concurrently submitted in candidature of any other degree. …………………………………………. Mohd Izwan Zulaini bin Abdul Gani 15010191 Date: i ACKNOWLEDGEMENT First of all, I would like to thank God for always giving me the strength and blessing during all these challenging times, as well as my beloved parents, Abdul Gani Abdullah and Nasabah Ismawi, all family members and friends for giving me motivation and moral support to finish my research. I gratefully acknowledge my supervisor Associate Prof. Dr. Ruhana Hassan for her continual encouragement, enthusiasm and support throughout my degree, and for providing constructive comments on all that I have written. I would like to express heartfelt thanks to my co-supervisor Mr. Rambli Ahmad for his invaluable advices that had greatly helped me to improve my research. In addition, I also would like to convey my gratitude to Mr. Oswald Braken Tisen, Mr. Engkamat Lading, Mr. Christoper Kri, Mr. Paschal Dagang, SWAT members and other staffs of Sarawak Forestry Corporation Bhd. (SFC) and Forest Department of Sarawak (FDS) for providing supports during crocodile samplings and other technical aspects of the study. A special thanks to Dr. Rossazana for advices on currency matters and also to fellow colleagues in the Molecular Aquatic Laboratory for helping me in the field surveys as well as to the lab assistants for helping to prepare the equipment before going to the field. Finally, I would like to express my special gratitude to the University Malaysia Sarawak (UNIMAS) for providing funds for this research through Dana Pelajar Ph.D. Grant no. F07/DPP53/1282/2015(28) and also the FDS for granting permit NCCD.907.4.4(jld.12)-193 and Permit No. NPW.907.4.4(JLD.14)-149 to conduct research on the crocodiles in Sarawak. Last but not least, thanks to Ministry of Higher Education for financing my study under MyPhD Scholarship Program. ii ABSTRACT This study is designed to gather information on historical exploitation and ongoing HCC; recent distribution and ecology of crocodile and genetic relationship of crocodile population in Sarawak, to aid sustainable crocodile management and finding solutions for mitigating the HCC. Historical data saw a connection between the exploitation of crocodile with decreasing trend of HCC in Sarawak from the Rajah Brooke era (1900 – 1941) until the post- war period (1946 – 1979), and an increasing trend of HCC from 1980 until 2017 in response to the recovery of the animal populations. Since 1900, crocodile attacks had been occurred in 22 major river basins (RB) in Sarawak, suggesting that the reptile has been widely dispersed throughout all major river basins in the state. For 118 years (1900 – 2017), the highest number of crocodile attacks were recorded in Lupar RB (22.2%) and the attacks had happened up to the inland areas of Belaga and Pelagus in Rajang RB. Further analysis of incidents show crocodile attacks were associated with the human activities pattern, where more attacks involved male victims (84.4%) and adults from age 31 to 40 years old (19.3%). The data also revealed that crocodile attacks in Sarawak could happen anytime regardless of the time, month, season, lunar cycle or tidal. However, more attacks were recorded during the daylight, in the months of March and April, during the Northeast monsoon, at the nights of the first quarter of the lunar cycle and at the time of high tide. Furthermore, fishing (25.2%) and bathing (24.4%) possess the highest risk of crocodile attack in Sarawak, clearly showed that crocodiles are more likely to attack when the victim is in water. Crocodile survey in selected tributaries in Rajang RB showed the distribution of the reptiles throughout the river basin with higher crocodile density at the lower region, the highest density was in Igan River (1.37 individuals/km); while in the middle and upper regions had recorded iii relatively low density with the lowest density recorded was in Katibas River (0.06 individuals/km) and no crocodile was spotted in Kanowit River. Four out of eight surveyed rivers in Rajang RB recorded increase in the density of crocodile compare to previous survey suggesting that the crocodile population in the river basin is experiencing recovery. The presence of crocodile in different regions (lower, middle and upper) of Rajang RB indicated that C. porosus in Sarawak live in wide range of habitats; from large salt water river system and small tidal tributaries (near to estuary) in lower region into hypo-saline or fresh water non-tidal tributaries in the middle and upper regions. Variation in term of density and distribution of crocodile between the different regions are mainly influenced by the saline characteristic of the river, habitats and the abundance of food sources for crocodile. Based on the analysis of DNA microsatellite sequence data, distinctive subpopulations of C. porosus according to geographical area (river basin) could be observed. High gene flow (Nm) among the crocodile subpopulations suggests frequent movements of the reptile happen across the river basins throughout Sarawak. In general, populations of C. porosus in Sarawak are experiencing expansion as supported by the mismatch distribution and evolutionary neutrality test data, suggesting that populations of crocodile in Sarawak are panmictic population. The findings of the present study imply that increasing of crocodile attacks is associated with the recovery and increased distribution of the reptile in Sarawak, thus crocodile management should emphasis on mitigating HCC and simultaneously continue the efforts for conservation of crocodile and its habitat. Keywords: Crocodylus porosus, human-crocodile conflict, recovery, expansion. iv Perpektif Sejarah, Taburan, Ekologi dan Genetik Populasi Buaya Air Masin (Crocodylus porosus Schneider, 1801) di Sarawak, Borneo, Malaysia ABSTRAK Kajian ini direka untuk mengumpul maklumat berkaitan sejarah eksploitasi dan KMB; taburan terkini populasi dan ekologi buaya serta hubungan genetik antara populasi buaya di Sarawak, untuk membantu pengurusan buaya secara lestari serta mencari solusi untuk mengurangkan KMB. Data sejarah memperlihatkan hubungkait antara eksploitasi buaya dengan tahap penurunan bilangan kes KMB di Sarawak dari era Rajah Brooke (1900 – 1941) sehingga ke tempoh selepas perang (1946 – 1979), dan tahap peningkatan bilangan kes KMB dari 1980 sehingga 2017 hasil tindakbalas daripada pemulihan populasi buaya. Semenjak tahun 1900, serangan buaya telah berlaku di 22 sungai utama di Sarawak, menunjukkan taburan luas reptilia tersebut di semua sungai utama di negeri ini. Dalam tempoh 118 tahun (1900 – 2017), serangan buaya tertinggi dicatatkan di Lembangan Sungai Lupar (22.2%) dan serangan telah berlaku sehingga ke kawasan pedalam Belaga dan Pelagus di Lembangan Sungai Rajang. Analisis lanjut insiden menunjukkan serangan buaya berkaitan dengan corak aktiviti manusia, dimana lebih banyak serangan melibatkan mangsa lelaki (84.4%) dan individu dewasa berumur dari 31 sehingga 40 tahun (19.3%). Data juga mendedahkan bahawa serangan buaya di Sarawak boleh berlaku bila-bila masa tanpa mengira masa, bulan, musim, kitaran bulan atau pasang surut air. Walau bagaimanapun, lebih banyak serangan telah direkodkan pada waktu siang, di bulan Mac dan April, semasa musim monsun Timur Laut, pada malam suku pertama kitaran bulan dan ketika air pasang. Aktiviti memancing (25.2%) and mandi di sungai (24.4%) mempunyai risiko serangan buaya yang tertinggi menunjukkan buaya lebih suka menyerang ketika mangsa berada di air. Survei buaya di sungai-sungai terpilih di Lembangan Sungai Rajang mendapati reptilia tersebut mendiami pelbagai habitat di sepanjang sungai, dengan kepadatan lebih tinggi di v bahagian hilir, tertinggi dicatatkan di Sungai Igan (1.37 individu/km); sementara itu di bahagian tengah dan hulu mencatatkan kepadatan yang lebih rendah dengan catatan terendah di Sungai Katibas (0.06 individu/km) dan tiada buaya dijumpai di Sungai Kanowit. Empat daripada lapan sungai yang disurvei di Lembagan Sungai Rajang mencatatkan peningkatan kepadatan buaya berbanding dengan survei terdahulu, menunjukkan bahawa populasi buaya di sungai ini sedang mengalami pemulihan. Kehadiran buaya di bahagian berbeza (bahagian hilir, tengah dan hulu) di Lembagan Sungai Rajang menunjukkan C. porosus, hidup di pelbagai habitat; dari sungai besar dan anak sungai (berhampiran muara) air masin di bahagian hilir sehinggalah kepada anak sungai air tawar yang tidak dipengaruhi pasang surut di bahagian tengah dan hulu lembagan sungai tersebut. Kepelbagaian dari segi kepadatan dan taburan di antara bahagian-bahagian berbeza adalah banyak dipengaruhi oleh ciri kemasinan sungai, habitat dan kelimpahan sumber makanan untuk buaya. Berdasarkan analisis data jujukan DNA mirosatelit, subpopulasi C. porosus berdasarkan kawasan geografi
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