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Fish communities as related to substrate
characteristics in the coral reefs of Kepulauan Seribu
Marine National Park, Indonesia, five years after
stopping blast fishing practices
Dissertation zur Erlangung des Doktorgrades
der Naturwissenschaften – Dr.rer.nat.
im Fachbereich 2 (Biologie/Chemie)
der Universität Bremen
vorgelegt von
Unggul Aktani
angefertigt am
Zentrum für Marine Tropenökologie
Bremen 2003
Gutachter der Dissertation :
1. Gutachter: Prof. Dr. Matthias Wolff 2. Gutachter: Dr. Andreas Kunzmann
Tag des öffentlichen Kolloqiums : 15 Mai 2003
Erratum
Erratum to: “AKTANI, U. 2003. Fish communities as related to substrate characteristics in the coral reefs of Kepulauan Seribu Marine National Park, Indonesia, five years after stopping blast fishing practices” A list of corrections follows: Page iv. Line 6-7 from above should be:
Chaetodon octofasciatus was abundant in areas dominated by Acropora corals. Chromis analis was abundant in areas dominated by sub-massive corals and other fauna.
Page iv. Line 11 from above should be:
… the current zoning management can not be considered an adequate tool to achieve this purpose.
Page 84. Line 11-12 from above should be:
C. octofasciatus is more abundant in area dominated by Acropora corals. C. analis is more abundant in areas dominated by sub-massive corals and other fauna.
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SUMMARY AKTANI, U. 2003. Fish communities as related to substrate characteristics in the coral reefs of Kepulauan Seribu Marine National Park, Indonesia, five years after stopping blast fishing practices
Kepulauan Seribu (“Thousand Islands”) is an archipelago of 110 small islands in the
southwest Java Sea. The archipelago is currently used for traditional fishing area,
tourism, sand mining, off shore oil exploration, sailing, and conservation. The major
problem in Kepulauan Seribu was blast fishing since the 1970’s, which had caused
extensive coral destruction. Blast fishing stopped since 1995 when the Kepulauan
Seribu Marine National Park was founded (since 1982 there was a nature reserve).
Six islands were chosen, each with three permanent transects (at 4-5 m depth) on the
northeast parts of each island, covering three management zones: Bira and Putri
(Sanctuary Zone), Genteng and Melinjo (Intensive Utilization Zone), and Pandan and
Opak (Traditional Utilization Zone). From October 2000 until August 2001,
underwater visual censuses were carried out within 45 day-intervals. The fish
transects were 50 × 5 m. Within the fish transects, underwater sequential photographs
were taken (50 × 1 m) to assess benthic groups and coral reef coverage. Classification
of the substrate type was based on benthic groups and life form categories.
Hard coral coverage was 43, 29, 25, 20, 18 and 7 % in Genteng, Pandan, Melinjo,
Bira, Opak and Putri, respectively. Dead corals were the dominant cover in all islands
surveyed (range: 52 to 83 %). The long-lasting impact of blast fishing on the
substrate was reflected by the presence of extensive fields of dead coral rubble (range:
31 to 59 %). In contrast to the zoning allocation, the percent hard coral cover in the
Sanctuary Zone was lowest and percent cover of dead coral was highest. The highest
cover of hard coral was found in the Intensive Utilization Zone.
A total of 119 fish species belonging to 25 families (32 863 fishes) were determined.
Pomacentridae was the most abundant family (range: 53 to 62 %), followed by
Labridae (27 to 33 %). Planktivore (28 to 40 %) and omnivore (27 to 37 %) fish were
the two most abundant trophic groups. The composition of the fish community
changed seasonally according to the alteration of west and east monsoon; with
seasonal shifts in both the fish species composition and fish abundances. During the
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west monsoon, Chromis atripectoralis and Halichoeres argus, while during the east
monsoon Pomacentrus lepidogenys, P. alexanderae and Cirrhilabrus cyanopleura
were abundant, respectively. The fish community was more related to the presence of
benthic groups and life form categories than to the coverage of hard corals.
Pomacentrus lepidogenys was abundant at encrusting corals. Pomacentrus
alexanderae was abundant at mushroom and dead corals. Chaetodon octofasciatus
and Chromis analis were abundant in areas dominated by Acropora corals. Benthic
feeders and omnivores preferred substrate with high cover of dead corals.
Planktivores preferred foliose corals.
Since the goal of the national park management is maintenance of a high coverage of
hard coral and a high diversity fish community, the current zoning management can
be considered an adequate tool to achieve this purpose. The results highly suggest a
re-zoning of the national park and should encourage the management to intensify both
surveillance frequency and law enforcement for the entire national park.
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ZUSAMMENFASSUNG
AKTANI, U. 2003. Fischgemeinschaften und ihr Bezug zu Substrat-Charakter-istika in den Korallenriffen vom Kepulauan Seribu Marine National Park, Indonesien, fünf Jahre nach dem Einstellen der Dynamitfischerei
Kepulauan Seribu (“Tausend Inseln”) ist ein Archipel mit 110 kleinen Inseln in der
südwestlichen Javasee. Das Archipel wird zur Zeit genutzt für traditionelle
Fischereigebiet, Tourismus, Sandabbau, Off-shore Ölförderung sowie den
Naturschutz. Das Hauptproblem in Kepulauan Seribu war seit den siebziger Jahren
die Dynamitfischerei, die in grossen Bereichen zur Zerstörung der Korallenriffe
geführt hatte. Die Dynamitfischerei ist seit 1995 eingestellt, als der Kepulauan Seribu
National Park gegründet wurde.
Sechs Inseln wurden ausgewählt, die in drei Managementzonen liegen: Bira und Putri
(Kernzone), Gentang und Melinjo (Intensive Nutzungszone), sowie Pandan und Opak
(Traditionelle Nutzungszone). An der Nordostseite jeder Insel wurden drei
Dauertransekte in 4-5 m Wassertiefe festgelegt. Von Oktober 2000 bis August 2001
wurden dort alle 45 Tage visuelle Fischzählungen durchgeführt. Die Fischtransekte
maßen 50 × 5 m. Innerhalb der Fischtransekte wurde das Substrat fotografiert (50 × 1
m), um den Deckungsgrad an benthischen Gruppen und an Korallen zu quantifizieren.
Die Substrattyp-Klassifizierung basierte auf benthischen Gruppen und „life form
categories“.
Der Deckungsgrad mit Hartkorallen in Gentang, Pandan, Melinjo, Bira, Opak und
Putri betrug jeweils 43, 29, 25, 20, 18 und 7 %. Tote Korallen waren die dominante
Bedeckung auf allen untersuchten Inseln (zwischen 52 und 83 %). Weite Flächen mit
Korallenschutt (31 bis 59 %) spiegeln den bleibenden Einfluss der Dynamitfischerei
auf das Substratgefüge wieder. Im Widerspruch zum höchsten Schutzstatus der
Kernzone wurde dort der geringste Deckungsgrad an Hartkorallen und der höchste
Grad an Bedeckung mit toten Korallen gefunden.
Insgesamt wurden 119 Fischarten aus 25 Familien nachgewiesen (32 863 Fische).
Pomacentridae stellten die häufigste Familie (53 bis 62 %), gefolgt von Labridae (27
bis 33 %). Planktivore (28 bis 40 %) und omnivore (27 bis 37 %) Fischarten waren
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die beiden häufigsten trophischen Gruppen. Die Zusammensetzung der
Fischgemeinschaft veränderte sich saisonal entsprechend dem Wechsel zwischen
West- und Ostmonsun, mit Verschiebungen sowohl in der
Fischartenzusammensetzung als auch in den Fischabundanzen. Während des
Westmonsuns waren Chromis atripectoralis und Halichoeres argus und während des
Ostmonsuns Pomacentrus lepidogenys, P. alexanderae und Cirrhilabrus cyanopleura
häufige Arten. Die Fischgemeinschaft stand eher im Bezug zu der Anwesenheit
benthischer Gruppen und „life form categories“ als zum Deckungsgrad mit
Hartkorallen. Pomacentrus lepidogenys war häufig mit Krustenkorallen
vergesellschaftet. Pomacentrus alexanderae wurde häufig an pilzförmigen Korallen
und an toten Korallen angetroffen. Chaetodon octofasciatus und Chromis analis
waren in Bereichen häufig, die von Acropora-Korallen dominiert wurden. Fische, die
ihre Nahrung am Boden finden und omnivore Fische bevorzugten Substrat mit einem
hohen Anteil an toten Korallen. Planktivore bevorzugten den Aufenhalt in der Nähe
von trichterförmigen Korallen.
Da der Erhalt eines hohen Deckungsgrades mit Hartkorallen und einer
Fischgemeinschaft mit grosser Diversität erklärte Aufgabe des Nationalpark-
Managements ist, kann die aktuelle Zonierung nicht als ein adäquates Instrument zum
Erreichen dieser Ziele angesehen werden. Die Ergebnisse weisen deutlich auf die
Notwendigkeit einer Re-Zonierung des Nationalparkes hin und sollten das
Management dazu ermutigen, sowohl die Überwachung vor Ort als auch die
Vollstreckung geltender Gesetze für den gesamten Nationalpark zu verstärken.
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RINGKASAN
AKTANI, U. 2003. Komunitas ikan dan keadaan substrat terumbu karang di Taman Nasional Laut Kepulauan Seribu, Indonesia, setelah lima tahun tidak terjadi penangkapan ikan dengan bahan peledak
Kepulauan Seribu terdiri dari 110 buah pulau kecil di Laut Jawa bagian barat daya.
Di kepulauan ini terdapat kegiatan wilayah penangkapan ikan tradisional, pariwisata,
pengambilan karang/pasir, penambangan minyak lepas pantai, pelayaran dan
perlindungan alam. Sejak tahun 1970-an permasalahan utama di Kepulauan Seribu
adalah penangkapan ikan dengan menggunakan bahan peledak yang mengakibatkan
kerusakan hebat terumbu karang. Penangkapan ikan dengan bahan peledak tidak
terjadi lagi sejak 1995 ketika kawasan tersebut dijadikan Taman Nasional (sejak 1982
sudah menjadi kawasan cagar alam).
Enam pulau di tiga zona pengelolaan yang berbeda dipilih sebagai lokasi penelitian,
masing-masing dengan tiga tempat pengamatan tetap (di kedalaman 4-5 m) pada
bagian timur laut pulau: Bira dan Putri (Zona Inti), Genteng dan Melinjo (Zona
Pemanfaatan Tradisional), Pandan dan Opak (Zona Pemanfaatan Trdisional). Sejak
bulan October 2000 sampai Agustus 2001, dilakukan pencacahan bawah air terhadap
komunitas ikan. Transek untuk pencacahan ikan berukuran 50 × 5 m. Di dalam
transek tersebut dilakukan pemotretan substrat terumbu karang secara
berkesinambungan sepanjang 50 × 1 m. Pengelompokan substrat terumbu didasarkan
pada jenis substrat dan bentuk terumbu karang.
Luas penutupan karang hidup di Genteng, Pandan, Melinjo, Bira, Opak dan Putri
berturut-turut adalah: 43, 29, 25, 20 dan 7 %. Jumlah penutupan karang mati adalah
paling luas diantara jenis substrat yang lain (berkisar dari 52 – 83 %). Dampak jangka
panjang kegiatan pengangkapan ikan dengan bahan peledak ditandai oleh banyaknya
luasan puing terumbu yang masih tampak (31 – 59 %). Hasil yang mengejutkan
adalah rendahnya luas penutupan karang hidup dan tingginya luas penutupan karang
mati di Zona Inti. Luas penutupan karang hidup tertinggi terdapat di Zona
Pemanfaatan Intensif.
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Jumlah ikan yang tercatat sebanyak 32863 ekor yang termasuk kedalam 119 species
dan 25 family. Pomacentridae merupakan family yang paling melimpah (53 – 62 %),
diikuti oleh Labridae (27 – 33 %). Planktivora (28 – 40 %) dan omnivora (27 – 37 %)
merupakan kelompok pemakan yang terbanyak. Komposisi komunitas ikan, dalam
hal ini species dan kelimpahan, berubah sesuai perubahan musim barat dan timur.
Selama musim barat species yang melimpah adalah Chromis atripectoralis dan
Halichoeres argus, sedangkan pada musim timur yang paling melimpah adalah
Pomacentrus lepidogenys dan Cirrhilabrus cyanopleura. Komunitas ikan lebih
terkait terhadap jenis substrat dan bentuk karang dibandingkan dengan luas penutupan
karang hidup. Pomacentrus lepidogenys melimpah pada karang ‘encrusting’.
Chaetodon octofasciatus dan Cromis analis melimpah di tempat yang banyak terdapat
koral jenis Acropora. Ikan pemakan hewan dasar dan omnivora menyukai substrat
karang mati. Planktivora menyukai karang ‘foliose’.
Tujuan pengelolaan taman nasional adalah menjaga tingginya penutupan terumbu
karang dan tingginya keragaman ikan, namun berdasarkan hasil peneltian
memperlihatkan bahwa pengelolaan yang ada belum bisa mencapai tujuan tersebut.
Saran yang bisa diajukan adalah melakukan penataan kembali zonasi yang ada dan
pihak pengelola melakukan peningkatan pengawasan di lapang dan penegakan
hukum.
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ACKNOWLEDGMENTS I would like to thank to German Academic Exchange Service (DAAD). This study would not have been possible without the scholarship from DAAD. Many people made this work possible and I would like to express my gratitude to all of them. I am most grateful to Prof. Dr. Matthias Wolff, who supervised and supported my work, for suggestions and the critical revision to work out essential results of my work. I am most thankful to Dr. Andreas Kunzmann who supervised me, for many fruitful discussions, and who has been patients, understanding and supportive through the whole period of my work. The late Prof. Dr. H.M. Eidman, who has gave supports and many valuable criticisms during preparation of my work. I deeply appreciate to Dr. Iris Kötter, who gave valuable comments and corrections in the earlier versions of my manuscript and also for her friendship and encouragements. Deeply thankful to Uwe Krumme for many fruitful discussions, valuable comments and who many times helps me to translate many letter to German, including the summary of this dissertation and suggestion to rethink about the title. I wish my gratitude to Center for Tropical Marine Ecology (ZMT) that gave best facilities to do my work. Special thanks for Prof. G. Hempel, Prof. V. Ittekkot, Prof. U. Saint-Paul and Dr. Werner Ekau. My grateful also for Tilman Appermann and Dr. Marc Kochcius for valuable comments. I thank to authority of the Kepulauan Seribu Marine National Park who gave me permission to do the field study, especially Drs. Achmad Abdullah, Ir. Andi Rusandi and Ibu Nena. The field work would have been impossible without help from Rangers of the marine park, especially: Pak Teguh and his family for preparing the food and accommodation; Pak Sairan, Pak Nelson and Pak Henry as diving buddy and for their enthusiasms and help me on all field surveys; Pak Riyad and Pak Daeng for accompanying me in many surveys; Pak Zakaria for diving equipments; Pak Salim, Pak Syarif, Pak Sokeh, Pendi, and Sigit. Thank you to Pak Mujar for the boat. Thank you Kak Jony for driving me to Muara Angke. My special thanks go to Dr. Mark Wunch, Dr. Claudio Richter, Gaby Boehme, Christa Müller, Sabine Kadler, Dr. Sabine Dittmann, Silke Meyerholz, Andreas Hanning, Matthias Birkicht, Dr. Carlos Jimenez, Dr. Gesche Krause, Iris Freytag, Jenny, Fernano Porto, Inga Nordhause, Kerstin Kober, Dr. Uta Berger, Dr. Marion Glasser, Kai Bergmann, Dr. Chriastiane Snack, Dr. Daniela Unger, Dieter Peterke, Dr. Tim Jennerjahn, Dr. Petra Westhaus-Ekau, Dr. Joko Samiaji, Dr. Rubén Lara, Natalie Loick, Uschi Stoll, Uschi Werner, Cristiane Hueerkamp, Ario, Auck, Eugene, and Mukhlis for encouragements and friendships over the last years. Many thanks for Jochen Scheuer who lent us many things. Many thank also to Wazir for maintenance of our underwater camera. For Dini and Yus Rustandi, thank you very much for the map. I am very thankful for my parents who gave me a chance to have a good education even in many social and economic difficulties. And also thanks for our big family that supports my education. The last but not least, many thanks for my wife, Mia, for the patient and who give me many supports, encourage and love. And for our children, Lala and Dhika, who gave me inspirations.
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TABLE OF CONTENTS
SUMMARY iii ZUSAMMENFASSUNG v RINGKASAN vii ACKNOWLEDGMENTS ix LIST OF FIGURES xii LIST OF TABLES xv LIST OF APPENDICES xv
1. INTRODUCTION .............................................................................................. 1
1.1. CORAL REEFS AND FISHES ...................................................................... 1
1.2. REEF FISHERIES ...................................……........................................... 3
1.3. KEPULAUAN SERIBU (THOUSAND ISLANDS) – STUDY SITE .................... 6
1.4. HYPOTHESES ....................................................................…………….. 11
1.5. OBJECTIVES OF THE STUDY..................................................................... 11
1.6. METHODOLOGICAL APPROACH.……………………………………….. 12
2. MATERIAL AND METHODS .......................................................................…. 14
2.1. THE STUDY AREA .................................................................................... 14
2.2. SAMPLING SITES ..................................................................................... 15
2.3. TIME FRAME OF STUDY ......…................................................................. 17
2.4. PERMANENT TRANSECTS ....................................................................... 17
2.5. CORAL SAMPLING ................................................................................... 18
2.6. REEF FISH SAMPLING .............................................................................. 20
2.7. DATA ANALYSES .………….................................................................. 21
3. RESULTS …………………………………………………………………... 31
3.1. FEATURES OF THE BENTHIC HABITAT.……….…….………………….. 31
3.2. PATTERN OF MAJOR BENTHIC GROUPS AND LIFE FORM CATEGORIES.… 35
3.3. REEF FISH COMMUNITY.….…………………………………………… 37
3.4. FISH DIVERSITY ………………………………………………………. 47
3.5. FISH SPECIES-ABUNDANCE RELATIONSHIP MODEL.……………………. 49
3.6. FISH COMMUNITY STRUCTURE.……………………………………….. 57
3.7. RELATING BENTHIC HABITAT WITH FISH COMMUNITY STRUCTURE …… 61
4. DISCUSSION ……………………………………………………………….. 68
4.1. VARIATION IN CORAL REEF COVERAGE ALONG THE GRADIENT OF BLAST FISHING IMPACT ………………………………………………
68
4.2. VARIATION IN FISH COMMUNITY ALONG THE GRADIENT OF BLAST FISHING IMPACT ……………………………………………………..
72
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4.3. SEASONAL CHANGES IN THE FISH COMMUNITY STRUCTURE.………… 79
4.4. VARIATION IN FISH DIVERSITY WITHIN THE ZONING MANAGEMENT…... 80
4.5. METHODOLOGICAL ASPECTS ………………………………………... 81
4.5.1. ASSESSMENT OF LIFE FORM CATEGORIES AND BENTHIC GROUPS …… 81
4.5.2. FISH VISUAL CENSUS.……………………………………………….. 82
5. CONCLUSIONS AND OUTLOOK……………………………….……………... 84
5.1. CONCLUSIONS ………………………………………………………... 84
5.2. OUTLOOK……………………………………………………………... 85
6. REFERENCES ………………………………………………………………. 86
APPENDICES.……………………………………………………………….. 94
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LIST OF FIGURES
FIGURE 1.1. Kepulauan Seribu Marine National Park (bordered by dash line) – the study area. 7 FIGURE 1.2. The zoning management of Kepulauan Seribu Marine National Park. ………….. 10 FIGURE 2.1. The study sites were located in Kepulauan Seribu Marine National Park. Insets
are the six selected islands with three sampling sites at each island. ………….….
16 FIGURE 2.2. The tetra-pod frame for photography coral coverage (modification from English
et al. 1994). ……………………………………………………………………….
19 FIGURE 2.3. Observer swims along the 50-m permanent transect at 0.5 m above the
substratum to visually census the reef fish (English et al. 1994). ………………..
21 FIGURE 2.4. The process of the multivariate analysis (modified from Field et al. 1982). (CA:
cluster analysis, PCA: principle component analysis, NMDS: non-metric multidimensional scaling). ………………………………………………………..
30 FIGURE 3.1. Percent cover of the benthic groups: hard corals, dead corals, other fauna and
algae. (The sequence of the islands was based on the zoning management: the Sanctuary Zone (Bira and Putri), the Intensive Utilization Zone (Melinjo and Genteng) and the Traditional Utilization Zone (Opak and Pandan)). ……...….….
32 FIGURE 3.2. Percent cover of Acropora life form categories: Acropora Branching (ACB),
Acropora Digitate (ACD) and Acropora Tabulate (ACT). .....................................
33 FIGURE 3.3. Percent cover of Non-Acropora life form categories, consisting of: Coral Sub-
massive (CS), Coral Foliose (CF), Coral Branching (CB), and Coral Encrusting (CE). ………………………………………….......................................................
33 FIGURE 3.4. Percent cover of dead coral in each island, consisting of: rubble dead corals
(DCR), massive dead corals (DCM), and dead corals with algae (DCA). .............
34 FIGURE 3.5. Average percentage of the number of colonies for all hard coral categories. ......... 34 FIGURE 3.6. The hierarchal dendrogram of all components of benthic groups and life form
categories produced by group average linkage displayed a tendency to separate the islands into three groups at 77 % similarity level (dash and dot line) and into two groups of geographical position: West and East side of the islands (solid line), without Opak (dash line). .............................................................................
35 FIGURE 3.7. NMDS plot of all components of benthic groups and life form categories. ........... 36 FIGURE 3.8. The PCA-biplot of benthic and life form categories. .............................................. 37 FIGURE 3.9. Abundance of the most abundant fish families at the different study sites during
the study time: October 2000 (a), March 2001 (b), April 2001 (c), June 2001 (d), August 2001 (e). Data were pooled from all sites in each island. .........................
40 FIGURE 3.10. Abundance of the most abundant fish families during the time of the study. Data
were pooled from all sites in each island. ...............................................................
41 FIGURE 3.11. Abundance of the different trophic groups at each study sites during the time of
study: October 2000 (a), March 2001 (b), April 2001 (c), June 2001 (d), and August 2001 (e). ......................................................................................................
42 FIGURE 3.12. Abundance of different trophic fish groups during the time of the study. Data
were pooled from all islands. ..................................................................................
43
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FIGURE 3.13. Number of fish species censused from Pandan (a), Opak (b), Bira (c), Putri (d),
Melinjo (e) and Genteng (f) with three sites each from October 2000 - August 2001. Solid triangle with solid line indicates the pooled (from 3 sites per island) number of species. Solid circle with dash line indicates the mean number of species (n = 3 sites per island, ± SE). ...................................................................
44 FIGURE 3.14. Rank abundance plot (a) and linear regression analysis of fish abundance (b) in
Pandan (the linear relationship is highly significant, P<0.01). Sampling time was in October 2000, March 2001, April 2001, June 2001 and August 2001. The table shows the ten most abundant species. R2 = coefficient of determination. .....
50
FIGURE 3.15. Rank abundance plot (a) and linear regression analysis of fish abundance (b) in
Opak (the linear relationship is highly significant, P<0.01). Sampling time was in October 2000, March 2001, April 2001, June 2001 and August 2001. The table shows the ten most abundant species. R2 = coefficient of determination. ..............
51
FIGURE 3.16. Rank abundance plot (a) and linear regression analysis of fish abundance (b) in
Bira (the linear relationship is highly significant, P<0.01). Sampling time was in October2000, March 2001, April 2001, June 2001 and August 2001. The table shows the ten most abundant species. R2 = coefficient of determination. ..............
52 FIGURE 3.17. Rank abundance plot (a) and linear regression analysis of fish abundance (b) in
Putri (the linear relationship is highly significant, P<0.01). Sampling time was in October 2000, March 2001, April 2001, June 2001 and August 2001. The table shows the ten most abundant species. R2 = coefficient of determination. ..............
53 FIGURE 3.18. Rank abundance plot (a) and linear regression analysis of fish abundance (b) in
Melinjo (the linear relationship is highly significant, P<0.01). Sampling time was in October 2000, March 2001, April 2001, June 2001 and August 2001. The table shows the ten most abundant species. R2 = coefficient of determination. .....
54 FIGURE 3.19. Rank abundance plot (a) and linear regression analysis of fish abundance (b) in
Genteng (the linear relationship is highly significant, P<0.01). Sampling time was in October 2000, March 2001, April 2001, June 2001 and August 2001. The table shows the ten most abundant species. R2 = coefficient of determination. .....
55 FIGURE 3.20. Rank abundance plot (a) and linear regression analysis of fish abundance (b) in
all islands (the linear relationship is highly significant, P<0.01). Sampling time was in October 2000, March 2001, April 2001, June 2001 and August 2001. The table shows the ten most abundant species. R2 = coefficient of determination. .....
56 FIGURE 3.21. Dendrogram of hierarchical clustering with group linkage methods of the fish
community, based on species abundance. Three replicate samples were made from each island at each sampling. (A=Pandan, B=Opak, C=Bira, D=Putri, E=Melinjo, F=Genteng). .........................................................................................
58 FIGURE 3.22. Non-metric multidimensional scaling ordination of the fish community based on
species abundance. Three replicate samples were made for each island at each sampling. (A=Pandan, B=Opak, C=Bira, D=Putri, E=Melinjo, F=Genteng). …...
58 FIGURE 3.23. PCA-plot of the fish community based on species abundance. Three replicate
samples were made for each island at each sampling. (A=Pandan, B=Opak, C=Bira, D=Putri, E=Melinjo, F=Genteng). ………………………………………
59 FIGURE 3.24. PCA-biplot of trophic group of fish produced by SVD method. The sampling
times were October 2000 and March, April, June, and August 2001. (A=Pandan, B=Opak, C=Bira, D=Putri, E=Melinjo, F=Genteng). …………………………….
60
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FIGURE 3.25. CCA-triplot of the distribution of selected fish-species found during October 2000-August 2001 in six islands: fish species (solid circle), life form and benthic variables (hollow circle), and the islands (solid square). The benthic variables were: Acropora Branching (ACB), Acropora Digitate (ACD), Acropora Tabulate (ACT), Coral Branching (CB), Coral Encrusting (CE), Coral Foliose (CF), Coral Massive (CM), Coral Sub-massive (CS), Mushroom Coral (CMR), Millepora (CME), Heliopora (CHL), Other Fauna (OT), Algae (AL), and Dead Coral (DC). The fish species were Chaetodon octofasciatus (Ctoc), Chromis analis (Cran), Pomacentrus alexanderae (Pmal), and Pomacentrus lepidogenys (Pmle). The sampling times were October 2000 (Oc), and March (Ma), April (Ap), June (Ju) and August 2001 (Au). …………………………………………...
62 FIGURE 3.26. CCA-triplot of the distribution of selected fish-species found during October
2000-August 2001 in six islands: fish species (solid circle), life form and benthic variables (hollow circle), and the islands (solid square). An arrow (dash line) was projected along the Acropora Branching variable that indicating a gradient; the perpendicular dash line in the arrow indicated the position of the islands along this gradient. (Refer to Figure 3.25 for abbreviations).…………………….
63 FIGURE 3.27. CCA-triplot of most abundant of fish-families from October 2000-August 2001
in six islands: fish families (solid circle), life form and benthic variables (hollow circle), and the islands (solid triangle). The benthic variables were: Acropora Branching (ACB), Acropora Digitate (ACD), Acropora Tabulate (ACT), Coral Branching (CB), Coral Encrusting (CE), Coral Foliose (CF), Coral Massive (CM), Coral Sub-massive (CS), Mushroom Coral (CMR), Millepora (CME), Heliopora (CHL), Other Fauna (OT), Algae (AL), and Dead Coral (DC). The fish families were: Pomacentridae (Poc), Labridae (Pmal), Scaridae (Sca), Chaetodontidae (Cha) and Nemipteridae (Nem). The sampling times were October 2000 (Oc), and March (Ma), April (Ap), June (Ju) and August 2001 (Au). ……………………………………………………………………………...
65 FIGURE 3.28. CCA-triplot of trophic groups of fish found from October 2000-August 2001 in
six islands: fish families (solid circle), life form and benthic variables (hollow circle), and the islands (solid triangle). The benthic variables were: Acropora Branching (ACB), Acropora Digitate (ACD), Acropora Tabulate (ACT), Coral Branching (CB), Coral Encrusting (CE), Coral Foliose (CF), Coral Massive (CM), Coral Sub-massive (CS), Mushroom Coral (CMR), Millepora (CME), Heliopora (CHL), Other Fauna (OT), Algae (AL), and Dead Coral (DC). Trophic groups of fish: herbivore (H), omnivore (O), planktivore (P), detritivore (D), benthic feeder (B), coralivore (C) and piscivore (Pi). The sampling times were October 2000 (Oc), and March (Ma), April (Ap), June (Ju) and August 2001 (Au). . ……………………………………………………………………….
67
xv
LIST OF TABLES
TABLE 2.1. The geographical-position of sampling sites in each island. .........………………… 17 TABLE 3.1. Complete list of fish families and species according to systematic order produced
by visual census method in all surveyed islands. …………………………………
39 TABLE 3.2. The diversity of fishes calculated by using some diversity formulas (A), and the
distribution model of fish species abundance in each island and for all islands together (B). The χ2 test is used to describe the goodness-of-fit of the distribution model with P<0.05. The percent value in brackets indicates the probability of the observed data to be the same as the expected distribution model. …..……………..
46 TABLE 3.3. The Comparison of the Shannon diversity index (H') between the islands in the
core zone (P. KA Bira and P. Putri Timur) and outside the core zone from each sampling time. The t-test was run at a significance level of P<0.001 (n.s.= Not significantly different; s. = Significantly different). …………………...…………...
47 TABLE 3.4. Comparison of Shannon diversity index (H') between the sampling times in all
islands. The t-test was performed at a significance level of P<0.001 (n.s. = Not significantly different; s. = Significantly different). . ………………………….…...
48
LIST OF APPENDICES
Appendix 1. Complete list of the percent cover of the major benthic groups and life form
categories (%) at the different study sites. ………………………………………...
94 Appendix 2. Number of coral colonies differentiated by their growth form at the study sites
assuming that coral growth is 2.4 mm per month and in circular direction, S=small (< 651 cm2; growth during five years), M=medium (651 - 940 cm2; growth during six years) and L=large (> 940 cm2; growth during seven or more years) (van Moorsel 1988). ……………………………………………………….
95 Appendix 3. Complete list of fish species according to their systematic order and their
abundances at each site throughout the study period. …………………………….
96 Appendix 4. Trophic group of all fish species observed (Sources: Lieske & Myers 1997; Fish
Base www.fishbase.org).…………………………………………………………..
100
1
1. INTRODUCTION
1.1. CORAL REEFS AND REEF FISHES
At least 794 species of scleractinian corals are known to build coral reefs
(Spalding et al. 2001). As biogenic structures, coral reefs constitute highly
fragmented habitats that are defined both by physical structure and the organisms
associated with them, including fishes and many invertebrates (Rosen 1984, Hubbard
1988, Choat & Bellwood 1991, Spalding et al. 2001).
The distribution and abundance of the coral reef fish community is strongly
influenced by biological and physical factors like wave exposure, sediment loads,
water depth as well as topographical complexity (rugosity) of the coral reef substrate
(e.g. Risk 1972, Luckhurst & Luckhurst 1978a, Carpenter et al. 1981, Williams 1982,
Bell & Galzin 1984, Sano et al. 1984, Galzin et al. 1994, McClanahan 1994,
Chabanet et al. 1997). Additionally, weather and currents were found to influence
reef fish community composition (Walsh 1983). Within a given family of reef fish,
ecological parameters like the coverage of living Scleractinian corals, food diversity
and reproductive behavior seem to affect the diversity of reef fishes (Galzin et al.
1994). However, according to Jennings & Polunin (1997), a single dominant process
rarely governs the structure of reef fish communities. Therefore, the general opinion
is that reef fish abundance and diversity are correlated with the complexity and health
of the coral reef habitat.
More than 4,000 species of teleost fish, representing about 18 % of the total
number of fishes, can be found in coral reefs (Choat & Bellwood 1991, Lieske &
Myers 1994, Spalding et al. 2001). According to Bellwood (1996 & 1998) it is hard
to define the fish living on coral reefs as “reef fish”, since “reef fish” families are
2
characteristic for coral reefs but their distribution are usually not restricted to them.
Therefore, it is not surprising that there are no fish families that are only restricted to
the coral rich region (Robertson 1998).
Choat & Bellwood (1991), however, found a number of fishes with a
characteristic appearance and morphology that are almost always associated with
coral reefs and achieve their highest abundance on them. The assemblages and
distribution of fishes on coral reefs vary greatly among habitat patches and the
complex architecture of the reef building corals (Choat & Bellwood 1991). As
biogenic structures corals depend on their physical and biological environment and
the interaction between biological and geological processes (Choat & Bellwood 1991,
Sale 1991, Williams 1991). Hence, Bellwood (1998) defines reef fishes as those
species that live on coral reefs and Robertson (1998) as the fish species that live on
consolidated substrata that form coral and inorganic reefs.
Two different theories about reef fish assemblages have been proposed:
according to the “order/deterministic” theory, reef fish have evolved specific habitat
requirements that reduce competition for limited resources and thereby enables the
coexistence of a great number of specialized species (Smith 1977). By contrast the
“chaos/stochastic” theory or “lottery” hypothesis postulates that reef fish assemblages
are highly variable and unpredictable over time (Sale 1974). Studies supporting one
or the other theory can be found. For example Greene & Shenker (1993) found that
the fish assemblages appeared to be extremely stable over the two-year period of their
investigation. The series studies from Sale (1974, 1975, 1976, 1982) supported the
chaos theory, albeit he derived the theory from coexistence in territorial behavior of
pomacentrid fishes in which each individual defended a small permanent territory for
3
food, shelter and nests sites. However, according to Bohnsack (1983) both theories
are valid for coral reef fish communities.
1.2. REEF FISHERIES
Coral reef fishes are mainly small and sedentary throughout most parts of their
lives (Sale 1991). However, they are important resources on coral reefs (Russ 1991),
contributing about 9 % of the total fish biomass in the World Oceans (Sorokin 1995)
or 7 % of the marine fish captured worldwide (Russ 1991). Coral reef fishery is an
important livelihood, particularly in developing countries (Munro & Williams 1985,
McManus 1997) and is typically a multi-species and multi-gear fishery (Spalding et al
2001).
Russ (1991) gave a comprehensive review about the effects of fishing on coral
reefs. According to his findings (that is hoped to be reinforced by the proposed
thesis), fishing activities cause habitat modification, thereby affecting fish populations
and communities’ level of reef fishes. Intensive fishing can cause large-scale and
long-term damage of coral coverage or structural heterogeneity of the benthic
substratum, and hence significantly affects reef fish communities. Destructive fishing
techniques have clearly negative impacts on reef fish communities (Russ & Alcala
1989, Saila et al. 1993). Munro & Williams (1985) stated that significant fishing
pressure can change the age and size structure of fish populations, decrease the stock
sizes and may change the community structure within a coral reef.
The blast fishing technique was introduced in the Indonesian Archipelago after
World War II as an easy way to catch schooling fish (Pet-Soede & Erdmann 1998).
The explosives are usually home made; often using glass bottles filled with a mixture
of agricultural fertilizer and kerosene, although dynamite is sometimes used as well
(Pet-Soede & Erdmann 1998; Spalding et al. 2001). The fishermen throw the bomb
4
by hand toward the reef, where it explodes on the water surface or within the water
body (Pet-Soede & Erdmann 1998). Even though Indonesian law prohibits blast
fishing, it is still common throughout the archipelago, particularly in remote areas
where the law enforcement is weak (Pet-Soede et al. 2000, UNESCO 2000). Besides
the ecological damage blast fishing also caused considerable economic losses to the
Indonesian society (Pet-Soede et al. 2000). This method kills both targeted (such as
dense schools of Siganids and Caesionids) and non-targeted fish, as well as
invertebrates (Pet-Soede & Erdmann 1998). However, the taxonomic and yield
composition of blast fishing varied highly (Fox & Erdmann 2000). Blast fishing also
damaged or destroyed the reef habitat and caused fields of coral rubble when the same
reef area was bombed several times (Pet-Soede & Erdmann 1998).
Blast fishing is considered one of the most destructive anthropogenic threats to
coral reefs, as not only the target fish, but also almost all organisms within the blast
radius get killed (McManus 1997, Pet-Soede et al. 1999, Fox et al. 2001). Destructive
fishing practices have reduced the productivity of coral reefs around the world
(Spalding et al. 2001) and led to a substantial reduction in cover of live coral and an
increase of dead coral rubble (Russ & Alcala 1989). This increase may attract fish
species, which are specialized in feeding on or settling onto coral rubble or both, e.g.
Labridae (Russ & Alcala 1989, Aktani 1990).
Recovery of the reef structure from a single blast may take years or decades
(Spalding et al. 2001). McManus et al. (1997) predicted that every year
approximately 1.4 % of the coral cover in the Philippines is lost due to blast fishing
and calculated that a reduction of fishing effort by approximately 60 % is required to
gain an optimal resource use and to solve the over-fishing problem due to blast fishing
activities. Riegl & Luke (1998) also found significant changes in coral and fish
5
community composition of blasted sites. Bombed or anchor-damaged coral reefs in
Indonesia are around 50% less diverse in shallow water as compared to undamaged
areas (Edinger et al. 1998).
Although coral reefs are of great ecological and economic importance, little is
known how coral reefs respond to human destructive fishing activities. Particularly
the process of recovery and natural regeneration of the coral reef itself and associated
animals lacks detailed studies (Saila et al. 1993, Riegl & Luke 1998, Hodgson 1999,
Fox et al. 2001). Furthermore, the understanding of the diversity of live, the
complexity of ecological interactions and the structures and patterns within coral reefs
is still limited (Sale 1976, Smith 1977, Hodgson 1999, Spalding et al. 2001).
Kaufman (1983) found that the destruction of reef fish habitats was followed by
changes in predator abundance, herbivore feeding behavior, and the distribution of
territorial damselfishes. Sano et al. (1984) observed that the destruction of
hermatypic corals led to changes in fish community structure resulting from a change
of food resources and the decrease in structural complexity of coral colonies.
Herbivore fishes, zooplankton feeders and omnivores were significantly more
abundant and of higher species richness on the living coral colonies than on damaged
coral colonies; or vice versa: when the structural complexity of the coral reef
decreased due to bio- and physical-erosion, the diversity and abundance of resident
reef fishes decreased as well. Bell & Galzin (1984) stated that the presence and
amount of live coral cover may be more important in structuring fish communities
than previously thought.
6
1.3. KEPULAUAN SERIBU (THOUSAND ISLANDS) – STUDY SITE
Kepulauan Seribu (Thousand Islands) is an archipelago that is located in the
southwest Java Sea or just northwest of Jakarta Bay (Fig. 1.1). It consists of 110
vegetated islands that stretch around 80 km from northwest to southeast and 30 km
from east to west. The southernmost reefs are located around 25 km northwest of
Jakarta Bay and are separated by a deep channel from Java Island (Ongkosongo &
Sukarno 1986, Tomascik et al. 1997). The islands are generally smaller than 10 ha
and their altitude is less than 3 m above sea level. The archipelago is used for
tourism, sand mining, off shore oil exploration, sailing, and conservation (UNESCO
2000). For many years, the major problem in Kepulauan Seribu was blast fishing,
which caused coral degradation (Hutomo 1987, Sukarno 1987).
Most of the ecological studies from Kepulauan Seribu are about the coral reefs,
but only few deals with reef fishes. According to Suharsono et al. (1998) at least 132
fish species belonging to 24 families can be found in Kepulauan Seribu. Hutomo &
Adrim (1986) observed that the diversity and abundance of fishes in Kepulauan
Seribu were higher on the reef slope than on the reef edge. Pomacentridae and
Labridae were the dominant fish families at the reef of Kepulauan Seribu (Hutomo
1987, Suharsono et al. 1998).
According to Moll & Suharsono (1986), Kepulauan Seribu has 193 coral species
belonging to 56 genera. The genera Acropora and Montipora dominate most of the
coral communities in the reef flat and the upper reef crest (Hutomo 1987). Moll &
Suharsono (1986) found a high coral diversity in many reefs in Kepulauan Seribu.
The coral cover, average colony size and diversity indicated a gradual increase with
distance from the mainland of Java. 88 species of scleractinian corals were described
in the southern reefs and 190 species in the north of Kepulauan Seribu (Spalding et al.
7
2001). However, the species composition of the upper reef slope is dependent on
environmental factors (Tomascik 1997).
FIGURE 1.1. Kepulauan Seribu Marine National Park (bordered by dash line) – the study area.
Since the 1920s the coral reefs in Jakarta Bay and some of the Seribu Islands
have been studied. In the past they were generally in good condition, though human
disturbance was already present (Moll & Suharsono 1986). Between 1985–1995,
8
most of these reefs were rapidly degrading (Moll & Suharsono 1986, UNESCO
2000). Reefs within Jakarta Bay were in dramatic decline, although they had already
been in poor condition in 1985. Most of these reefs can be considered functionally
dead (Ongkosongo & Sukarno 1986, Stoddart 1986). Three islands in this region
disappeared below sea level during this time and several others were eroding,
probably caused by a combination of dredging for landfill and natural loss of
sediments (UNESCO 2000). A decline in coral reef cover was also observed 15 km
to 50 km offshore from Java Island in 1995. However, several reefs had increased in
coral cover. In this region, the major problems were natural and human disturbances.
The natural disturbances became apparent when outbreaks of the crown-of-thorns
starfish occurred and water temperature increased due to the El Niño Southern
Oscillation Phenomenon (ENSO) (UNEP/IUCN 1988, Brown & Suharsono 1990,
Warwick et al. 1990). The human disturbances were identified as the poison fishing
method, pollution from the Jakarta coastal area and the muro-ami coral breakage in
the 1980s and 1990s (UNESCO 2000). Muro-ami is a fishing technique that uses a
drive-in net and a line to scare the fish and drive them out of the reef toward a bag net
(often cause the breaking of live corals) (Erdmann 1998). Most reefs beyond 50 km
off Java Island also indicated a decline in coral cover during 1985-1995. Destructive
fishing practices like blast and cyanide fishing were the major problems in the outer-
region (Brown 1986, UNEP/IUCN 1988; UNESCO 2000). However, the outer reefs
of Kepulauan Seribu showed relatively high coral cover and diversity compared with
reefs in Jakarta Bay.
According to Erdmann (1998) and UNESCO (2000), there was no evidence of
blast fishing in Kepulauan Seribu from 1995 until the field research of this study
started in 2000 (pers. com. with the Rangers of Kepulauan Seribu Marine National
9
Park). The disappearance of blast fishing may be related to the absence of target
fishes and the establishment of a Marine National Park in this area (Ministry of
Forestry Decree No. 162/Kpts-II/1995, 21 March 1995) (UNEP/IUCN 1998).
However, Kepulauan Seribu was already declared as a reserve since 1982 (Ministry of
Agricultural Decree No. 257/Kpts/7/82, 21 July 1982) (BAPEDALDA 2000). But,
unfortunately small-scale sodium cyanide fishing and illegal coral rock mining were
still occurring in Kepulauan Seribu (Alder et al. 1994) until now. In contrast to blast
fishing, the targets of cyanide fishing are ornamental fishes and invertebrates for the
aquarium trade.
The area of Kepulauan Seribu Marine National Park is divided into four
management-zones (Fig. 1.2) (KSMNP 2000). The first, Sanctuary (Core) Zone is a
strict nature reserve, consisting of three areas: Sanctuary Zone I is set aside as a
hawksbill turtle habitat, Sanctuary Zone II as a hawksbill nesting area, and Sanctuary
Zone III for the coral reef ecosystem. The second, Protection Zone, is purposed for
protection of the Sanctuary Zone. The third, Intensive Utilization Zone is purposed
for tourism activities, such as snorkeling, SCUBA diving, beach based activities and
boating without conflict or environmental damage. The forth, Traditional Utilization
Zone is designated for traditional fishing methods using trap, net, and hand line
fishing.
The anthropogenic impact on the coral reef ecosystem in the Sanctuary Zone
and the Protection Zone was expected to be more obviously visible, when compared
to the other zones, since this zone was designated for the protection and preservation
of plants and animals. Entering this zone was strictly limited to research and
educational activities. The anthropogenic impact in the Intensive Utilization Zone
was expected to be moderate, due to its use for tourism activities. Considering all
10
zones in Kepulauan Seribu, the anthropogenic impact on the coral reef was predicted
to be highest in the Traditional Utilization Zone.
FIGURE 1.2. The zones of Kepulauan Seribu Marine National Park.
11
The former blast fishing activities (in all zones) were indicated clearly by the
large fields of coral rubble and subsequent new coral growth on rubble (although the
author had not directly witnessed the former blast fishing activities). With this in
mind, it seemed interesting to study how the reef fish-community has recovered from
blast fishing practices in the past.
1.4. HYPOTHESES
It is hypothesized that coral reef fishes are more diverse and abundant in the
Sanctuary Zone. A second hypothesis postulates that reef fish communities have
developed a clear pattern of relationship with the heterogeneity of benthic substrates.
This relationship corresponds to the degree of recovery of the coral reef habitat after
five years of no blast fishing.
1.5. OBJECTIVES OF THE STUDY
The overall goal was to find information on reef fish assemblages associated
with the recovery of coral reefs that had suffered from blast fishing activities several
years ago.
The following specific questions were addressed in this study:
- Are impacts of blast fishing on a coral reef fish community still visible after five
years?
- What degree of relationship between varying heterogeneity of benthic substrates
can be found and is the reef fish community structure different in the sites/areas
now?
- Which environmental factors determine the structure of reef fish communities?
- Are reef fishes more diverse in the Sanctuary Zone than outside?
12
The study results are expected (1) to allow for the prediction of the succession
of a reef fish community after blast fishing, (2) to contribute to the solution of
maintaining the biodiversity, and (3) to provide information for evaluating the zoning
management of the national park.
1.6. METHODOLOGICAL APPROACH
Several approaches were used for this study. The study was based on the
following facts and assumptions: since 1995 until 2000 no blast fishing had occurred
in all islands within the park (UNESCO 2000), so the coral reefs had already
recovered at least partly. According to personal communication with the marine park
rangers, there was no fishing in the Sanctuary Zone, and five years were enough time
for fish communities to recover from blast fishing impact.
The coral reef coverage was assessed by taking underwater sequential
photographs. This technique has the advantage, that it takes relatively little time in
the field and provides a permanent record (Done 1981). However, it has also some
disadvantages, like ineffectiveness in sampling small and hidden colonies, a very
limited perception of depth (Done 1981), and it is very time-consuming to evaluate
the pictures on the computer.
Underwater visual census (UVC) was used in this study to assess the reef fish
community. UVC by SCUBA divers has been an important tool for fish ecologists in
enumerating the abundance and composition of reef fish assemblages on coral reefs
(Sale & Sharp 1983, Bell et al. 1985, Harvey et al. 2002). The underestimation of
reef fish densities is already known from this method (Sale & Sharp 1983, Bell et al.
1985, Harvey et al. 2001). However, trained observers showed consistent results in
estimating the same population (Bell et al. 1985, Polunin & Roberts 1993).
13
Univariate and multivariate methods were applied to analyse the benthic
substrate composition and the fish community pattern (Clarke & Green 1988). The
fish communities were also assessed using species richness indices, Shannon diversity
index and Pielou’s evenness index, and several species-abundance distribution
models.
14
2. MATERIAL AND METHODS
2.1. THE STUDY AREA
Kepulauan Seribu is an island chain that consists of patch reef complexes and
fringing reefs (Hutomo 1987, Tomascik 1997). The geographical position is between
5o24’ - 5o47’ south latitude and 106o23’ – 106o37’ east longitude. Around 25 km
north from Java Island a deep channel (-88 m depth) separates the southernmost reefs
from Java Island. Pari Island is the southernmost and the only platform of the
Kepulauan Seribu patch reef complex located on the southern side of the channel
(Ongkosongo & Sukarno 1986, Tomascik et al. 1997).
In Kepulauan Seribu, the extension of many islands and channels show a strong
east-west orientation. In addition, the lateral reef growth is characteristically along an
east-west axis (Tomascik et al. 1997). This phenomenon reflects the dominant east-
west direction of winds and currents in the Java Sea (Ongkosongo & Sukarno 1986).
During the west monsoon (in general from December through March, dominant wind
direction from the northwest; sometimes September-November is a transition to the
west monsoon), however, currents in the southwest Java Sea are mostly in a southeast
direction, sweeping across the Kepulauan Seribu at velocities generally not exceeding
40 cm.s-1 (Soegiarto 1981, Tomascik et al. 1997). During the east monsoon (in
general from April through November) currents in the southwest Java Sea run in a
southwest direction, with velocities exceeding sometimes 50 cm.s-1, generating a net
flow into the Indian Ocean through the Sunda Strait (Tomascik et al. 1997). The east
monsoon has a much larger impact on the islands geomorphology than the west
monsoon (Ongkosongo & Sukarno 1986). During the west monsoon, the wind blows
eastward and carries heavy rainfall throughout the region (Sukarno 1987). Then, the
15
city of Jakarta is becoming an increasingly important source for siltation and pollution
in Jakarta Bay and Kepulauan Seribu, since many small rivers drain from this city
(Ongkosongo & Sukarno 1986, Willoughby 1986, Uneputty & Evans 1997,
Willoughby et al. 1997, Rees et al. 1999, Williams et al. 2000).
The reversing monsoon system is also the primary environmental factor
structuring the coral-reef communities in the region (Hutomo 1987, Tomascik 1997).
The seaward reef slopes of Kepulauan Seribu have a relatively moderate angle,
usually between 30o – 60o, with corals growing down to 20 m depth (Hutomo 1987,
Tomascik et al. 1997). Most of the islands have a narrow sandy shore and a wide reef
flat (Hutomo 1987). However, the islands are considered to be located in a relatively
sheltered environment, protected from severe storms and ocean swell (Tomascik et al.
1997). The low-amplitude (microtidal) diurnal-tide (i.e. one high, one low per day) in
the Java Sea has a subordinate role in shaping current velocities that are predominant
of monsoonal character (Tomascik et al. 1997).
2.2. SAMPLING SITES
Before the permanent sampling sites were chosen, a pre-survey was done in 30
islands. Based on the results of this survey, six islands with three sampling sites each
(on the northeast parts) were chosen in three management zones (for detailed position
see Table 2.1). For the Sanctuary Zone two islands (Indonesian: Pulau) were selected:
Pulau (P.) Kayu Angin Bira and P. Putri Timur (for convenience they will be called as
Bira and Putri, respectively). In the Intensive Utilization Zone, P. Kayu Angin
Genteng and P. Melinjo were taken. P. Pandan and P. Opak Besar (Opak) were
chosen as the Traditional Utilization Zone (Fig. 2.1).
16
FIGURE 2.1. The study sites were located in Kepulauan Seribu Marine National Park. Insets are the six selected islands with three sampling sites at each island.
17
TABLE 2.1. The geographical-position of sampling sites in each island.
Island (Code) Sites
P. Pandan (A) A-1: 05° 42.388' S 106° 34.164' E
A-2: 05° 42.403' S 106° 34.114' E
A-3: 05° 42.590' S 106° 33.869' E
P. Opak Besar (B) B-1: 05° 40.008' S 106° 35.188' E
B-2: 05° 40.016' S 106° 35.148' E
B-3: 05° 40.024' S 106° 35.137' E
P. Kayu Angin Bira (C) C-1: 05° 36.329' S 106° 34.117' E
C-2: 05° 36.327' S 106° 34.076' E
C-3: 05° 36.337' S 106° 34.053' E
P. Putri Timur (D) D-1: 05° 35.326' S 106° 34.074' E
D-2: 05° 35.455' S 106° 34.411' E
D-3: 05° 35.531' S 106° 4.417' E
P. Melinjo (E) E-1: 05° 34.198' S 106° 32.552' E
E-2: 05° 34.196' S 106° 32.597' E
E-3: 05° 34.191' S 106° 32.645' E
P. Kayu Angin Genteng (F) F-1: 05° 37.281' S 106° 33.764' E
F-2: 05° 37.147' S 106° 33.779' E
F-3: 05° 37.149' E 106° 33.796' S
Abbreviation: S = Latitude South; E = Longitude East.
2.3. TIME FRAME OF STUDY
The pre-survey and preparation of this study was done between August and
September 2000. The main study was carried out from October 2000 until August
2001, with 45 day intervals between the sampling times. Unfortunately, the data from
December 2000 and January 2001 were lost (due to robbery). Thus only data of
October 2000, March, April, June, and August 2001 were available for the analysis.
Depending on the weather condition, between 9 to 12 days were needed each
time to survey all the sampling sites. Each island was observed at least during one
day. Underwater visual census along transects (see section 2.4.) was done first,
followed by coral photography. It was always tried to minimize frightening the reef
fish community.
2.4. PERMANENT TRANSECTS
Permanent transects (of 50 m length) for fish and corals were installed at fixed
locations (using the same line transects) at 4 – 5 m depth, depending on the
occurrence of new coral growth on fields of coral rubble. The transect lines were
straight, following the depth contour and were laid down parallel to the reef front.
18
Both edges of the transect were demarcated by a cemented sinker into the reef
pavement. Tape measures were laid out again between both marks at each survey,
and then removed after each census. A Global Positioning System (GPS)-receiver
was used to relocate the permanent transects.
2.5. CORAL SAMPLING
For fish and coral assessment belt transects were used. For coral assessment, it
was a combination of line intercept transect (LIT) (English et al. 1994) and
photogrammetry (Done 1981). Whereas the fish transect was 50 m x 5 m and the
coral transect was one meter wide and 50 m long (English et al. 1994).
During the time of the survey period the percent cover of corals was assessed
twice: at the beginning and at the end of the study. Therefore photographic methods
were combined with the line intercept transect. The entire length of each 50 m
transect was photographed using a Nikonos V camera with a 35-mm lens and a tetra-
pod frame (Fig. 2.2) whereby the base of the rectangular frame served as reference
bar. Continuous sequential photographs with 200-ASA negative films were taken
along 50-m transect (the coverage of the camera lens was 1 m x 1.4 m when using the
tetra-pod). A total of 1,292 photographs were scanned and the areas of the reef life-
form categories measured using ImageJ V 1.14c (NIH) software (McCook (2001)
used the same software) and converted to percent cover of benthic groups and life
form categories.
The life-form categories used in this study were based on English et al. (1994):
Acropora Branching (ACB), Acropora Digitate (ACD), Acropora Tabulate (ACT),
Coral Branching (Non- Acropora) (CB), Coral Encrusting (CE), Coral Foliose (CF),
Coral Massive (CM), Coral Sub-massive (CS), Mushroom Coral (CMR), Millepora
(CME), Heliopora (CHL), Other Fauna (OT) (including: Soft Corals, Sponges,
19
Zoanthids, others benthic organisms), Algae (AL) (consisting of: Macro Algae,
Halimeda), Dead Coral (DC) (consisting of: dead coral with Algae, rubble and
massive dead coral).
FIGURE 2.2. The tetra-pod frame for photography coral coverage (modification from English et al. 1994).
The classification of coral colony size (related to the age) was based on several
assumptions: the average coral growth in a linear direction was 24 mm per month (as
the radius, r) (van Moorsel 1998). Another assumption was that the starting time of
coral growth was 1995 to 2000, when there were no more blast fishing activities until
the study was conducted. The third assumption was, that coral growth was in circular
direction (van Moorsel 1988). The following equation was used to calculate the
colony size:
2area Size r×=π where r = radius of coral colony (time-dependent, growth rate 24 mm/month),
π = a constant, 3.14
1 m
1 m
1.685 m
0.2 m
20
2.6. REEF FISH SAMPLING
Though obtaining accurate assessment of reef fish abundance with underwater
visual census (UVC) was not perfect and not a simple matter, UVC was the most
practical non-destructive way and still permitted to estimate the abundance of reef fish
species, with relatively quick time in the field, repeatable and inexpensive (Sale &
Sharp 1983, Bell et al. 1985, English et al. 1994, Samoilys & Carlos 2000).
The reef fish community was studied with the daytime underwater visual census
method, recording the fish species and their abundance. The fish census was carried
out between 10.00 a.m. and 3.00 p.m. to avoid possible diurnal-nocturnal behavioral
changes (Carpenter et al. 1981, Helfman 1993). A census took about two hours,
including the waiting time after laying out the measuring tape. Census was done only
once per site.
Fish were generally identified to species level, but due to difficulties of getting
fish samples for closer taxonomic inspection of specimen some taxa were identified
only to genus level. Within genera every unidentified species was tentatively given a
number to name as ‘species’. Identification of fish species was based on Burgess &
Axelrod (1972), Masuda et al. (1984), Allen & Steene (1987), Kuiter (1992), Lieske
& Myers (1997) and Allen (1999).
The following procedure was used (modified from Russ 1985; Greene &
Shenker 1993; English et al. 1994):
1. A species list of reef fishes was developed for the studied area (pre-survey result).
2. A 50-m measure tape was laid out followed by a waiting period of 45-60 minutes.
3. Two SCUBA divers swam very slowly (35-50 minutes) at 0.5 m above the
substratum along the 50-m transect. A single observer recorded the fish species
21
and its abundance on an underwater slate (Fig. 2.3), while the other served as a
dive buddy swimming behind the observer.
FIGURE 2.3. Observer swims along the 50-m permanent transect at 0.5 m above the substratum to visually census the reef fish (English et al. 1994).
2.7. DATA ANALYSES
Univariate methods were used to measure the percentage of coral cover and
various diversity and evenness indices for both, coral and fishes. According to
Warwick et al. (1990), with adequate sample replication, the statistical significance of
changes in the univariate indices can be assessed using a standard test. Multivariate
analyses were used to visualize the species abundance matrix and the composition of
benthic groups and life form categories (Clarke & Green 1988). Warwick et al.
(1990) found that low level perturbation in a community might be detected with
greater sensitivity using multivariate rather than univariate analysis. Two multivariate
methods used in the study were the ordination and clustering technique. The
ordination technique was used to visualize the relationship between the samples
(Clarke & Green 1988). The cluster technique was used to form discrete groupings of
22
samples (Clarke & Green 1988). Clarke & Green (1988) suggested that combining
both techniques is a good strategy, although descriptive multivariate analyses make no
parametric assumption at all.
The data used for the analysis were pooled from three replicate sites in each
island. The pooled fish abundance data for each island were analyzed using diversity
indices. Both fish and benthic groups of fish were also analyzed by multivariate
statistical methods.
Taylor (1978) stated that ‘diversity’ was seen as a property of the multi-species
population that is equivalent to ‘density’ in a single-species population. According to
Magurran (1988) species diversity measurement can be divided into three categories:
The first are the species richness indices, which are essentially a measure of the
number of species in a defined sampling unit. They instantly provide a
comprehensive expression of diversity. In this category, number of species and
Margalef’s diversity index were used for this study. The second are the diversity
indices based on the proportional abundance of species that seek to take richness and
evenness into a single figure. This category includes the Shannon diversity index and
Pielou’s evenness index that were used in this study. The third are the species
abundance models that describe the distribution of species abundances, whereby the
relative abundance is considered to represent the basic pattern of niche utilization in
the community or area (Southwood 1978). Four species abundance distribution
models were examined in this study for the fish data: the log series (logarithmic series
distribution), the log normal distribution (truncated log normal), the geometric series
and MacArthur’s broken stick distribution model.
23
2.7.1. SPECIES RICHNESS
A simple measure of species diversity is the species number recorded (S) (Poole
1974). Margalef’s index (d) is an alternative measure of diversity to incorporate both
the total number of individuals (N) and the species numbers. However, both S and d
indices ignore the distribution of individuals among the species. Margalef’s index (d)
(Clarke & Warwick 1994) is calculated as:
( )N
Sdlog
1−=
2.7.2. DIVERSITY INDICES BASED ON THE PROPORTIONAL ABUNDANCE OF SPECIES
The Shannon diversity index (H’) is based on the proportional abundance of
species assuming that individuals are randomly sampled from an ‘indefinitely-large’
community (Magurran 1988). The Shannon diversity index was used to measure the
diversity:
( )∑=
−=′s
iii ppH
1ln
SiNnp i
i ,...,3,2,1 ; ==
where S = the number of species,
in = the number of individuals of the ith species, N = the total number of individuals for all S species, and
ip = the proportional abundance of the ith species. The variance of Shannon diversity index (Var H’) was calculated using the
formula (Poole 1974, Magurran 1988):
( )
( )21
2
1
2
21
)lnln Var
NS
N
ppppH
s
i
s
iiiii −
−
−
=′∑ ∑= =
24
To compare two Shannon diversity indices, a t-test was applied (Magurran
1988):
2/121
21
)Var Var ( HHHHt
′+′′−′
=
where H´1 is the Shannon diversity index in the first community and H´2 in the second
community.
The degree of freedom was calculated according to (Magurran 1988):
( )
′−
′′+′
=
2
22
1
21
221
)Var ()Var (Var Var
NH
NH
HHdf
where N1 and N2 were the number of individuals in the first and second sample,
respectively. A t-table was used to look up the results.
The homogeneity of the reef fish community was measured by Pielou’s
evenness index:
maxHHJ′
=′
where Hmax is the maximum possible diversity, which would be achieved if all species
were equally abundant (= ln S) (Clarke & Warwick 1994).
2.7.3. FISH SPECIES-ABUNDANCE DISTRIBUTION MODELS
The equitability of the species-abundance relationship will reflect the
underlying distribution (Southwood 1978). The rank of relative species abundance
can be used to construct community models that are a characteristic pattern of the
community (Fisher et al. 1943, May 1975, Pielou 1975, Southwood 1978, Magurran
1988). The log series distribution predicts that species arrive at an unsaturated habitat
at random intervals of time and then occupy the remaining niche (with one or few
dominant environment factors) (Magurran 1988). Theoretically, the community
25
consists of a small number of abundant species and many species with low abundance
(Magurran 1988). The log normal distribution of relative abundance indicates a large,
mature and natural community with a large number of species fulfilling diverse
ecological roles (niche) (May 1975, Magurran 1988). By contrast the geometric
series distribution or the ‘nice-preemption’ hypothesis predicts that species arrive at
an unsaturated habitat at regular intervals of time and occupy the remaining niche
fraction (May 1975, Magurran 1988). The broken stick model describes a more
equitable state of affairs than the three previous models, because it discusses more in
rank-abundance form than in species abundance (May 1975, Magurran 1988).
2.7.3.1. THE LOG SERIES DISTRIBUTION
The general formula for log series distribution is calculated according to (Fisher
et al. 1943):
+=
αα NS 1ln
and the distribution follows (Fisher et al. 1943, Poole 1974, Magurran 1988):
nxxxx
nαααα , ... ,3
,2
,32
where α is a constant known as Fisher’s diversity index, x is a sampling parameter or
a constant related to the average number of individuals per species and n is the
abundance class. The total number of species (S) is obtained by:
( )[ ]xS −−= 1lnα
To calculate the expected frequencies in each abundance class, x is estimated by
iterative solution:
( ) ( )[ ]xx
xNS
−−
−
= 1ln1
where N is the total number of individuals in the community.
26
The α was calculated as:
( )x
xN −=
1α
The octaves or doublings of species abundance class was chosen for calculation.
To compare the observed species and abundance data with the expected value, a Chi
squared (χ2)-test was done with (number of classes – 1) degrees of freedom (Sokal &
Rohlf 1995). Each class was calculated by:
( )Expected
Expected - Observed 22 =χ
2.7.3.2. THE LOG NORMAL DISTRIBUTION
The log normal distribution can be written as (May 1975, Magurran 1988):
( )220 exp)( R-aSRS =
where S(R) = the number of species in the R-th octave,
S0 = the number of species in the modal octave,
( ) 2/122σ=a = the inverse width of the distribution
However, a truncated of log normal was used since most of log normal species
abundance data are the truncated variety (May 1975, Pielou 1975, Magurran 1988).
The procedure is:
1. Each species abundance was converted into log10 ( )inx 10log= and then the
mean
= ∑
Sx
x and the variance ( )
−= ∑
Sxx 2
2σ were calculated.
2. 5.0log100 =x and rxr 10log= ; where r is the observed variate.
3. γ was calculated using: ( )20
2
xx −=
σγ ; where γ is a measure of the relationship
between the mode of the individuals curve and the upper limit of the species
curve.
27
4. The auxiliary estimation function ( )θ , which corresponds to the γ value, was
found in Cohen’s table (Magurran 1988).
5. The estimation of mean ( )xµ and variance ( )xV of x were calculated using:
( )0ˆˆ xxxx −−= θµ and ( )20
2 ˆˆ xxVx −+= θσ
6. The standardized normal variate ( )0z , which corresponds to the truncation
point of 0x , was calculated by ( )x
x
V
xzˆ
ˆ00
µ−=
7. The area under the tail of a standard normal curve to the left of 0z or
( )00 Pr zZp ≤= was found from tables of normal distribution.
8. The total number of species in the community ( )*S was determined as
01*ˆ
psS−
=
9. The octaves or doublings of species abundance class was chosen for
calculation. And Chi squared (χ2)-test was used with (number of classes – 3)
degrees of freedom.
2.7.3.3. THE GEOMETRIC SERIES DISTRIBUTION
The species abundance rank in geometric series was sequenced from most to
least abundant (May 1975, Magurran 1988):
( ) 11 −−= iki kkNCn
where in = the number of individuals in the ith species,
N = the total number of individuals,
( )[ ] 111
−−−= s
k kC and is a constant which ensures that Nni =∑
The constant (k) was calculated by iterating the following formula:
( )( )( )
−−−
−
= s
s
kk
kk
NN
111
1min
where minN is the number of individuals in the least abundant species.
28
The value of the constant Ck was calculated as ( )[ ] 111
−−−= s
k kC
A Chi squared (χ2)-test was used to find the goodness of fit with (number of species –
1) degrees of freedom.
2.7.3.4. THE BROKEN STICK DISTRIBUTION
In the broken stick distribution, the octaves or doublings of species abundance
class was also used (Magurran 1988). The expected number of species was calculated
by (May 1975):
( ) 211)(−
−
−
=S
Nn
NSSnS
where S(n) = the number of species in the abundance class with n individuals
N = total number of individuals,
S = total number of species
The observed and the expected number of species were used to calculate Chi
squared (χ2)-test with degrees of freedom (number of classes – 1).
2.7.4. MULTIVARIATE ANALYSIS
Cluster analysis (CA) was used to group entities of the benthic groups and also
the fish abundance into a dendrogram according to their similarities (Ludwig &
Reynolds 1988, Clarke & Warwick 1994, Legendre & Legendre 1998). For fish, the
cluster analysis was based on the Bray-Curtis Similarity index with the group average
linkage method. Data was transformed with square root without standardizing. For
the benthic data the same method was used but data was not transformed.
Non-metric multidimensional scaling (NMDS) was used to construct an
ordination of the benthic groups and the fish abundances in a 2D-map that plots
dissimilar objects far apart and similar objects close to each other in the ordination
space (Clarke & Warwick 1994, Legendre & Legendre 1998). The NMDS ordination
29
technique was based on Bray-Curtis similarity. The stress value that indicates how
well that configuration represents the multidimensional similarity between the
samples based on the classification from Kruskal (1964):
Stress Goodness of fit
20 % Poor 10 % Fair 5 % Good
2.5 % Excellent 0 % Perfect
Principal component analysis (PCA) was used to place the samples into a map
that reflects their similarity like in NMDS (Clarke & Warwick 1994, Legendre &
Legendre 1998). PCA appeal was based on its apparent mathematical elegance
Ludwig & Reynolds 1988). In this study, PCA-ordination of two-way interaction
(with rows and columns centered) was used (Lipkovich & Smith 2002).
Canonical correspondence analysis (CCA) was employed to relate fish
community compositions to variations of the benthic groups in the environment in a
simultaneous two-dimensional plot (ter Braak 1986, Legendre & Legendre 1998,
Lipkovich & Smith 2002). CCA was calculated using the singular value
decomposition (SVD) method of two-way matrix data. The CCA-plot was displayed
with rows and columns centered and symmetric biplot scaling (Lipkovich & Smith
2002). The process of the multivariate computation (CA, NMDS and PCA) is
summarized in Fig. 2.4.
30
FIGURE 2.4. The process of the multivariate analysis (modified from Field et al. 1982). (CA: cluster analysis, PCA: principle component analysis, NMDS: non-metric multidimensional scaling).
Multivariate computations were performed with PRIMER 5 (Plymouth Routines
in Multivariate Ecological Research) software (Clarke & Warwick 1994) and Biplot
display was performed by Biplot and Singular Value Decomposition Macro for
Excel© developed by Lipkovich & Smith (2002). The diversity indices were
calculated by PRIMER and manually. The distribution models were manually
calculated by using Excel software. The linear regression was calculated
automatically by Excel software.
PCA
CA
NMDS
31
3. RESULTS
3.1. FEATURES OF THE BENTHIC HABITAT
The percent cover of the major benthic groups (sum of all animals, plants and
dead corals) was highly variable among the surveyed islands (Fig. 3.1). Dead corals
were the most dominant cover in all surveyed islands: it was lowest at Genteng (51.6
%) and highest at Putri (83.4 %) (Appendix 1).
By contrast percent cover of hard corals was highest at Genteng (42.7 %) and
lowest at Putri (7.6 %). The group of hard corals was divided into the two life form
categories Acropora and Non-Acropora (Fig. 3.2 and 3.3). The Acropora life forms
were further subdivided into three categories and Non-Acropora life forms were sub-
divided into eight categories (see Appendix 1). The average percent cover of both
Acropora and Non-Acropora life form categories were highly variable among the
islands (Fig. 3.2).
The “other-fauna” (OT) group was present in all surveyed islands, but the cover
never exceeded 7 % at any island (Fig. 3.1). This group consisted of 12 sub-
categories, including soft corals. The algae group that consisted of three components,
covered between 1.2 % and 6.9 % (Fig. 3.1, Appendix 1).
The islands in the Sanctuary Zone (Bira) and in the border of the Sanctuary
Zone (Putri) were characterized by a high number of dead corals (Fig. 3.1). The
coverage of dead corals in Bira was 3.6 times and in Putri 11 times higher than the
cover of hard corals.
Melinjo and Genteng, which are located in the Intensive Utilization Zone, were
in better condition compared with the two islands of the Sanctuary Zone. In Melinjo
the cover of dead corals was 2.6 times higher than the live coral cover. Genteng had
32
the lowest value of dead coral cover divided by live coral (only 1.2). The hard coral
cover in Melinjo and Genteng amounted to 25 % and 42.8 %, respectively.
Pandan and Opak (Traditional Utilization Zone) had 29.1 % and 18.2 % of
hard coral cover, respectively. The comparison of dead corals to hard coral cover in
Pandan was 2.2 and in Opak 2.9.
Among the other dead coral components rubble had the highest cover in all
islands (Fig. 3.4). The coverage of rubble was between 30.6 % and 58.6 %, being
highest in Putri, followed by Bira.
Small coral colonies were dominant in all areas surveyed (Fig. 3.5, Appendix 2).
Their cover was lowest in Genteng (79.4 %) and the highest in Bira (90.4 %).
������������������������
��������������
�����������������������������������
��������������������������������������������������������
����������������������������
������������������������������������������
��������������
��������������
��������������
��������������
�������������� ������
��������������
��������������
�������������� ������
������������
��������������
0
10
20
30
40
50
60
70
80
90
P. KA Bira P. Putri Timur P. Melinjo P. KA Genteng P. Opak Besar P. Pandan
Cov
er (%
)
Dead Coral����
Hard Coral���
Other Fauna���
Algae
FIGURE 3.1. Percent cover of the benthic groups: hard corals, dead corals, other fauna and algae. (The sequence of the islands was based on the zoning management: the Sanctuary Zone (Bira and Putri), the Intensive Utilization Zone (Melinjo and Genteng) and the Traditional Utilization Zone (Opak and Pandan)).
33
�������� �������� �������� ������������������������ ��������
0
1
2
3
4
5
6
7
8
9
10
P. KA Bira P. Putri Timur P. Melinjo P. KA Genteng P. Opak Besar P. Pandan
Cov
er (%
)
ACB ACT
������ ACD
FIGURE 3.2. Percent cover of Acropora life form categories: Acropora Branching (ACB), Acropora Digitate (ACD) and Acropora Tabulate (ACT).
������������ ������
������������������������������������������������������
������������������������������������������������������������ ������
������������������������������������������������������
������������ ������
������������
���������������
������������
������������������������
������������ ������ ����� ������
������������
������������������ ������ ������
������������ ������
������������
0
5
10
15
20
25
P. KA Bira P. Putri Timur P. Melinjo P. KA Genteng P. Opak Besar P. Pandan
Cov
er (%
)
CS������ CF
������ CB
������ CM
������ CE
FIGURE 3.3. Percent cover of Non-Acropora life form categories, consisting of: Coral Sub-massive (CS), Coral Foliose (CF), Coral Branching (CB), and Coral Encrusting (CE).
34
���������������� �������� ��������
������������������������������������������������0
10
20
30
40
50
60
70
P. KA Bira P. Putri Timur P. Melinjo P. KA Genteng P. Opak Besar P. Pandan
Cov
er (%
)
DCR DCM
��������DCA
FIGURE 3.4. Percent cover of dead coral in each island, consisting of: rubble dead corals (DCR), massive dead corals (DCM), and dead corals with algae (DCA).
������������������������
����������������
������������������������
����������������
����������������
0
10
20
30
40
50
60
70
80
90
100
P. KA Bira P. Putri Timur P. Melinjo P. KA Genteng P. Opak Besar P. Pandan
Num
ber o
f col
ony
(%)
< 651 cm2 651 - 940 cm2���
> 940 cm2
FIGURE 3.5. Average percentage of the number of colonies for all hard coral categories.
35
3.2. PATTERN OF MAJOR BENTHIC GROUPS AND LIFE FORM CATEGORIES Between the surveyed islands the composition of the benthic habitat was highly
variable. Multivariate analyses were used to find linkages among them. Figure 3.6
shows a hierarchical dendrogram. At a similarity level of 77 % three different benthic
and life form groups were distinguished. The first group consisted of the islands Bira
and Putri (at 83.7 % similarity level) that are both located in the Sanctuary Zone of the
national park. The second group was Pandan, Melinjo and Genteng (at 77.6 %
similarity level). The third group consisted of Opak only.
P. P
anda
n (A
)
P. M
elin
jo (E
)
P. K
A G
ente
ng (F
)
P. P
utri
Tim
ur (D
)
P. K
A Bi
ra (C
)
P. O
pak
Besa
r (B)
100
90
80
70
60
Sim
ilarit
y
FIGURE 3.6. The hierarchal dendrogram of all components of benthic groups and life form categories produced by group average linkage displayed a tendency to separate the islands into three groups at 77 % similarity level (dash and dot line) and into two groups of geographical position: West and East side of the islands (solid line), without Opak (dash line).
West side East side
(II) (I) (III)
36
The NMDS-plot of all components of benthic groups and life form categories
showed a slightly different trend in the separation of the islands than the dendrogram
(Fig. 3.7). In the NMDS-plot Bira and Putri, both are located in the Sanctuary Zone,
were grouped together. Genteng, Melinjo and Pandan built another group. Opak was
markedly separated from these two groups.
P. Pandan (A) P. Opak Besar (B)
P. KA Bira (C)
P. Putri Timur (D)
P. Melinjo (E) P. KA Genteng (F)
Stress: 0.0
East side
West side
Intensive Utilization Zone
Traditional Utilization Zone
Sanctuary Zone
(II)
(I)
(III)
FIGURE 3.7. NMDS plot of all components of benthic groups and life form categories.
PCA-ordination showed a different trend in the grouping of the islands, when
compared to the two previous methods (Fig. 3.8). The islands of the Sanctuary Zone,
Bira and Putri, were grouped together. Genteng and Pandan built another group.
Melinjo and Opak were clearly separated from the two former groups.
37
P. KA Genteng (F)
P. Melinjo (E)
P. Putri Timur (D)
P. KA Bira ( C)
P. Opak Besar (B)
P. Pandan (A)
AL-M
AL-H
AL-C
OT-all categories
DCA
DCT
DCR
DCM
DCB
CHLCME
CS
CMRCM
CF
CE
CB
ACT
ACD
ACB
-5
-4
-3
-2
-1
0
1
2
3
4
-6 -4 -2 0 2 4 6PC-1: 51.2 %
PC-2: 35.7 %
FIGURE 3.8. The PCA-biplot of benthic groups and life form categories.
3.3. REEF FISH COMMUNITY
A total of 32,863 fishes were censused from 18 permanent sites during the study
period, but the data does not include small pelagic fishes (Appendix 3). Altogether
119 fish species belonging to 25 families were observed (Table 3.1.). The minimum
and maximum abundances per island and per observation ranged between 651 and
1,006 individuals (Table 3.2.). In general Pomacentridae was the most abundant
family at all times in all islands. Two families (Pomacentridae and Labridae) tended
to be the most abundant in each island, followed by Scaridae, Chaetodontidae,
Nemipteridae and Apogonidae (Fig. 3.9 & 3.10). Further families were of low
abundance.
Planktivore and omnivore fish were the two most abundant trophic groups in
each island and the whole surveyed area (Fig. 3.11 & 3.12; Appendix 4) followed by
benthic feeders, herbivores, coralivores, piscivores and detritivores. In the
Traditional Utilization Zone planktivores were obviously the most abundant trophic
group during the study, except in Opak where the omnivores were most abundant in
38
March and April 2001. In the Intensive Utilization Zone omnivores were generally
the most abundant trophic group (Fig. 3.11 & 3.12). However with exception, in
Melinjo benthic feeders were abundant in October 2000 and in Genteng planktivores
were the most abundant in March 2001. In the Sanctuary Zone (Bira), omnivores
were most abundant from April – August 2001 (during the east monsoon), whereas in
October 2000 during the west monsoon), benthic feeders were the most abundant
group.
At the beginning of the study (October 2000), the number of fish species was
generally lower compared to all other censuses in all observation sites (Fig. 3.13).
However, the fish abundance at the beginning of the study was not always lower when
compared to the following observations (Table 3.2.). The fish diversity index (H’)
also changed during the study (ranged 2.36 to 3.19) (Table 3.2.). Fish evenness is
given in Table 3.2.
39
TABLE 3.1. Complete list of fish families and species according to the systematic order produced by the visual census method in all surveyed islands. Muraenidae Pomacanthidae Labridae
1 Gymnothorax sp. 37 Centropyge bicolor 82 Anampses sp. Holocentridae 38 Chaetodontoplus mesoleucus 83 Cheilinus chlorourus
2 Myripristis adusta 39 Pomacanthus sp. 84 Cheilinus fasciatus 3 Myripristis violacea Pomacentridae 85 Cheilinus undulatus 4 Myripristis sp. 40 Abudefduf vaigiensis 86 Choerodon anchorago 5 Sargocentron praslin 41 Abudefduf bengalensis 87 Cirrhilabrus cyanopleura
Synodontidae 42 Abudefduf sexfasciatus 88 Diproctacanthus xanthurus 6 Synodus sp. 43 Amblyglyphidodon curacao 89 Epibulus insidiator
Aulostomidae 44 Amblyglyphidodon leucogaster 90 Gomphosus varius 7 Aulostomus chinensis 45 Amblyglyphidodon ternatensis 91 Halichoeres argus
Fistulariidae 46 Amphiprion frenatus 92 Halichoeres chloropterus 8 Fistularia commersonii 47 Amphiprion ocellaris 93 Halichoeres hortulanus
Tetrarogidae 48 Amphiprion percula 94 Halichoeres melanurus 9 Ablabys taenianotus 49 Amphiprion sandaracinos 95 Halichoeres purpurescens
Scorpaenidae 50 Amphiprion sp. 96 Halichoeres vrolikii 10 Pterois volitans 51 Cheiloprion labiatus 97 Hemigymnus melapterus Serranidae 52 Chromis analis 98 Labroides dimidiatus 11 Cephalopholis argus 53 Chromis atripectoralis 99 Macropharyngodon ornatus 12 Cephalopholis boenak 54 Chromis flavipectoralis 100 Pteragogus sp. 13 Cephalopholis sp. 1 55 Chromis viridis 101 Stethojulis strigiventer 14 Cephalopholis sp. 2 56 Chromis weberi 102 Thalassoma hardwicke 15 Epinephelus sp. 1 57 Chromis xanthura 103 Thalassoma lunare 16 Epinephelus sp. 2 58 Chromis sp. 104 Thalassoma lutescens 17 Epinephelus sp. 3 59 Chrysiptera rollandi 105 Thalassoma purpureum Apogonidae 60 Chrysiptera sp. Scaridae 18 Apogon compressus 61 Dascyllus aruanus 106 Scarus ghobban 19 Cheilodipterus macrodon 62 Dascyllus trimaculatus 107 Scarus niger 20 Cheilodipterus quinquelineatus 63 Dischistodus melanotus 108 Chlorurus sordidus 21 Sphaeramia nematoptera 64 Dischistodus prosopotaenia 109 Scarus viridifucatus Lutjanidae 65 Neoglyphidodon bonang 110 Scarus sp. 1 22 Lutjanus biguttatus 66 Neoglyphidodon melas 111 Scarus sp. 2 23 Lutjanus decussatus 67 Neoglyphidodon nigroris Blenniidae 24 Lutjanus fulviflammus 68 Neopglyphidodon oxyodon 112 Meiacanthus smithi Haemulidae 69 Neopomacentrus anabatoides Microdesmidae 25 Plectorhinchus chaetodonoides 70 Neopomacentrus azysron 113 Ptereleotris evides Nemipteridae 71 Plectroglyphidodon lacrymatus Acanthuridae 26 Pentapodus trivittatus 72 Pomacentrus alexanderae 114 Acanthurus lineatus 27 Scolopsis bilineata 73 Pomacentrus amboinensis Siganidae 28 Scolopsis lineatus 74 Pomacentrus grammorhyncus 115 Siganus canaliculatus 29 Scolopsis margaritifer 75 Pomacentrus lepidogenys 116 Siganus corallinus Mullidae 76 Pomacentrus philippinus 117 Siganus vulpinus 30 Parupeneus barberinus 77 Pomacentrus sp. 1 Ostraciidae Ephippidae 78 Pomacentrus sp. 2 118 Ostracion cubicus 31 Platax sp. 79 Pomacentrus sp. 3 Tetraodontidae Chaetodontidae 80 Pomacentrus taeniometopon 119 Arothron sp. 32 Chaetodon auriga 81 Stegastes fasciolatus 33 Chaetodon octofasciatus 34 Chaetodon vagabundus 35 Chelmon rostratus 36 Heniochus sp.
40
����� ����� ����� ��������� ����� ����� ����� ����� ���������� ����� ����� ����� ����� ���������� ����� ����� ����� ����� �����0
10
20
30
40
50
60
70
80
90
P. KA Bira P. Putri Timur P. Melinjo P. KA Genteng P. Opak Besar P. Pandan
Abu
ndan
ce (%
)
October 2000
���������� ����� �����
���������� �����
�������� ����� �����
����������
���������� ���������� �����
���������� �����
���������� ���������� �����
���������� �����
����������
0
10
20
30
40
50
60
70
80
90
P. KA Bira P. Putri Timur P. Melinjo P. KA Genteng P. Opak Besar P. Pandan
Abu
ndan
ce (%
)
March 2001
���������������
���������� �����
����������
�������������� ����� �����
����������
���������� ���������� ����� ����� �����
���������� ���������� ����� ����� ����� �����
0
10
20
30
40
50
60
70
80
90
P. KA Bira P. Putri Timur P. Melinjo P. KA Genteng P. Opak Besar P. Pandan
Abu
ndan
ce (%
)
Pomacentridae Labridae
������ Scaridae
��������Chaetodontidae
������ Nemipteridae
��������Apogonidae
April 2001
FIGURE 3.9. Abundance of the most abundant fish families at the different study sites during the study time: October 2000 (a), March 2001 (b), April 2001 (c), June 2001 (d), August 2001 (e). Data were pooled from all sites in each island.
(a)
(b)
(c)
41
����������
��������������� ����� ����� ��������� ����� �����
���������� ����� ���������� ����� ����� ����� ����� ���������� ����� ����� ����� �����
0
10
20
30
40
50
60
70
80
90
P. KA Bira P. Putri Timur P. Melinjo P. KA Genteng P. Opak Besar P. Pandan
Abu
ndan
ce (%
)
June 2001
��������������� �����
����������
��������
����������
����������
����������
����������
����������
����������
����������
���������� �����
����������
����������
���������� ����� ����� ����� ����� �����0
10
20
30
40
50
60
70
80
90
P. KA Bira P. Putri Timur P. Melinjo P. KA Genteng P. Opak Besar P. Pandan
Abu
ndan
ce (%
)
Pomacentridae Labridae���
Scaridae����
Chaetodontidae���
Nemipteridae����
Apogonidae
August 2001
FIGURE 3.9. Continued.
����� ����� ������ ������������������������ ������ ������ ������ ������������ ������ ������ ������ ����������� ������ ����� ����� ������
0
10
20
30
40
50
60
70
80
90
Oct. 00 Mar. 01 Apr. 01 Jun. 01 Aug. 01
Abu
ndan
ce (%
)
Pomacentridae Labridae���
Scaridae����
Chaetodontidae���
Nemipteridae����
Apogonidae
FIGURE 3.10. Abundance of the most abundant fish families during the time of the study. Data were pooled from all sites in each island.
(e)
(d)
42
�������������������������
����������������
������������������������
���������������������������������������������
����������������
����������
���������� ����
���������� ���� ����
��������������� ���� ����� ���� ��������� ����� ���� ����� ���� ����
272
241
116
482
572
455
262
161
299
503
160
69
306
278
384
278
209
81
49
15
46
15
40 44
0 0 0 0 0 07 2 2 13 0 14 2 4 4 2 1
0
100
200
300
400
500
600
700
800
900
P. KA Bira P. Putri Timur P. Melinjo P. KA Genteng P. Opak Besar P. Pandan
Indi
vidu
al n
umbe
r
October 2000
���������������������������������������������
��������������������
����������������������������������������
�����������������������������������
������������������������������������
������������������������������
����������
��������
����������
��������
����������������
������������������� ���� ����� ��������� ���� ����� ���� ���� ��������� ����� ���� ����� ���� ����
110
368
365 38
8
383 40
1
488
235
532
363
453
291
255
158 18
4
329
368
178
71
39 46 38
163
119
10 0 12 0 2 48 1 2 5 8 45 10 8 10 7 100
100
200
300
400
500
600
700
800
900
P. KA Bira P. Putri Timur P. Melinjo P. KA Genteng P. Opak Besar P. Pandan
Indi
vidu
al n
umbe
r
March 2001
����������������������������������������
������������������������
����������������������������������������������������
���������������������������������������������
��������������������
������������������������������
����������
������������
���������������
��������
����������������
������������������������ ����� ��������� ���� ����� ���� ���� ��������� ����� ���� ����� ���� ����
288
354
243
450
269
555
463
320
772
562
277
354
289
278
183
292
385
234
68
164
154
63
229
211
1 0 0 0
15 163 10 2 14 2 510 7 7 20 16 17
0
100
200
300
400
500
600
700
800
900
P. KA Bira P. Putri Timur P. Melinjo P. KA Genteng P. Opak Besar P. Pandan
Indi
vidu
al n
umbe
r
Planktivore
������ Omnivore Benthic feeder
������Herbivore
��������Detritivore
������ Corallivore
������Piscivore
April 2001
FIGURE 3.11. Abundance of the different trophic groups at each study site during the time of study: October 2000 (a), March 2001 (b), April 2001 (c), June 2001 (d), and August 2001 (e).
(a)
(b)
(c)
43
�������������������������������������������������������
����������������
��������������������������������
���������������������������������������������
��������������������
����������������������������������������
���������������
����������������
����������
��������
��������
������������������� ����� ���� ��������� ���� ����� ���� ���� ��������� ����� ���� ����� ����
337
278 31
0
504 53
3
680
609
208
410
511
242
399
302
233
208
213
173
255
154
198
46 56
72
142
2 1 1 0 0 2
16 5 9
27
4 125 7 3 9 0
14
0
100
200
300
400
500
600
700
800
900
P. KA Bira P. Putri Timur P. Melinjo P. KA Genteng P. Opak Besar P. Pandan
Indi
vidu
al n
umbe
r
June 2001
�����������������������������������
������������������������
������������������������
�������������������������
��������������������
�����������������������������������
��������������������
������������
���������� ����
����������������
�������������������������
��������������� ���� �����
�������� ���� ��������� ����� ���� ����� ���� ����
86
245
350
510
428
710
372
284 31
5
254
220
391
175
152
263
204
184
140
206
137
32
15
198
263
0 0 0 0
30
011 3 11 23 12 101 1 8 5 5 70
100
200
300
400
500
600
700
800
900
P. KA Bira P. Putri Timur P. Melinjo P. KA Genteng P. Opak Besar P. Pandan
Indi
vidu
al n
umbe
r
Planktivore���
Omnivore Benthic feeder���
Herbivore����
Detritivore���
Corallivore���
Piscivore
August 2001
FIGURE 3.11. Continued.
����������������������������������������
���������������������������������������������
����������������������������������������
������������������������������������������������������������
����������������������������������������
����������
������������
�������������������������
����������������
������������������������� ���� ����� ��������� ����� ����� ����� ��������� ���� ����� ���� �����
1454
2362
2748
2379
1836
2138
2015
2159
2642
2329
1536
1472
1661
1384
1118
209
476
889
668
851
0 28 32 6 3025 28 36 73 70
17 50 77 38 27
0
500
1000
1500
2000
2500
3000
Oct. 00 Mar. 01 Apr. 01 Jun. 01 Aug. 01
Indi
vidu
al n
umbe
r
Omnivore���
Planktivore Benthic feeder���
Herbivore���
Detritivore���
Corallivore���
Piscivore
FIGURE 3.12. Abundance of different trophic fish groups during the time of the study. Data were pooled from all islands.
(e)
(d)
44
0
20
40
60
Oct. '00 Dec. '00 Jan. '01
Mar. '01 Apr. '01 Jun. '01 Aug. '01
Num
ber o
f spe
cies
P. Pandan
0
20
40
60
Oct. '00 Dec. '00 Jan. '01
Mar. '01 Apr. '01 Jun. '01 Aug. '01
Num
ber o
f spe
cies
P. Opak Besar
0
20
40
60
Oct. '00 Dec. '00 Jan. '01
Mar. '01 Apr. '01 Jun. '01 Aug. '01
Num
ber o
f spe
cies
P. KA Bira
FIGURE 3.13. Number of fish species censused from Pandan (a), Opak (b), Bira (c), Putri (d), Melinjo (e) and Genteng (f) with three sites each from October 2000 - August 2001. Solid triangle with solid line indicates the pooled (from 3 sites per island) number of species. Solid circle with dash line indicates the mean number of species (n = 3 sites per island, ± SE).
(a)
(b)
(c)
45
0
20
40
60
Oct. '00 Dec. '00 Jan. '01
Mar. '01 Apr. '01 Jun. '01 Aug. '01
Num
ber o
f spe
cies
P. Putri Timur
0
20
40
60
Oct. '00 Dec. '00 Jan. '01
Mar. '01 Apr. '01 Jun. '01 Aug. '01
Num
ber o
f spe
cies
P. Melinjo
0
20
40
60
Oct. '00 Dec. '00 Jan. '01
Mar. '01 Apr. '01 Jun. '01 Aug. '01
Num
ber o
f spe
cies
P. KA Genteng
FIGURE 3.13. Continued.
(d)
(e)
(f)
46
TA
BL
E 3
.2.
The
div
ersi
ty o
f fis
hes c
alcu
late
d by
usi
ng so
me
dive
rsity
form
ulas
(A),
and
the
dist
ribut
ion
mod
el o
f fis
h sp
ecie
s abu
ndan
ce in
eac
h is
land
and
fo
r all
isla
nds t
oget
her (
B).
The
χ2 te
st is
use
d to
des
crib
e th
e go
odne
ss-o
f-fit
of t
he d
istri
butio
n m
odel
with
P<0
.05.
The
per
cent
val
ue in
bra
cket
s ind
icat
es
the
prob
abili
ty o
f the
obs
erve
d da
ta to
be
the
sam
e as
the
expe
cted
dis
tribu
tion
mod
el.
O
ctob
er 2
000
Mar
ch 2
001
April
200
1
Pand
an
Opa
k Bi
ra
Putri
M
elin
jo
Gen
teng
Pa
ndan
O
pak
Bira
Pu
tri
Mel
injo
G
ente
ng
Pand
an
Opa
k Bi
ra
Putri
M
elin
jo
Gen
teng
A.
Div
ersi
ty in
dice
s S
(Tot
al s
peci
es)
37
33
33
33
44
40
49
50
55
45
49
44
54
49
55
48
56
50
d (S
peci
es
richn
ess)
5.
56
4.64
4.
70
4.89
6.
37
5.44
6.
94
6.77
7.
88
6.57
6.
81
6.11
7.
32
6.78
7.
69
6.68
7.
62
6.76
N (T
otal
in
divi
dual
s)
651
983
900
699
851
1295
10
07
1384
94
7 81
1 11
49
1133
13
92
1193
11
22
1133
13
61
1401
J' (E
venn
ess)
0.
65
0.72
0.
73
0.81
0.
75
0.77
0.
70
0.80
0.
71
0.75
0.
69
0.79
0.
74
0.82
0.
77
0.80
0.
73
0.78
H
' (lo
g e)
2.36
2.
52
2.56
2.
84
2.84
2.
84
2.74
3.
15
2.85
2.
85
2.68
3.
01
2.94
3.
19
3.10
3.
11
2.93
3.
05
α D
iver
sity
(Fis
her)
8.50
6.
58
6.73
7.
20
9.84
7.
82
10.7
7 10
.16
12.7
2 10
.27
10.3
9 9.
11
11.1
7 10
.29
12.1
2 10
.16
11.7
7 10
.13
B.
Fit o
f Dis
trib
utio
n M
odel
Loga
rithm
ic s
erie
s Ye
s (2
4.9%
) Ye
s (9
9.4%
) Ye
s (6
8.5%
) Ye
s (2
9.3%
) Ye
s (7
6.6%
) Ye
s (1
4.2%
) Ye
s (7
9.7%
) Ye
s (7
9.1%
) Ye
s (3
8.4%
) Ye
s (9
6.8%
) Ye
s (7
2.3%
) Ye
s (3
1.6%
) Ye
s (2
2.1%
) Ye
s (3
5.5%
) Ye
s (4
3.7%
) Ye
s (2
9.2%
) Ye
s (9
5.9%
) Ye
s (9
2.4%
)
Log
norm
al
Yes
(28.
5%)
Yes
(89.
5%)
Yes
(52.
2%)
Yes
(35.
8%)
Yes
(30.
6%)
No
(0%
) Ye
s (7
7.2%
) Ye
s (1
5.9%
) Ye
s (3
0.8%
) Ye
s (9
5.1%
) Ye
s (4
6.4%
) Ye
s (5
1.1%
) Ye
s (6
3.5%
) Ye
s (1
5.6%
) Ye
s (3
1.8%
) Ye
s (7
7.7%
) Ye
s (8
6.5%
) Ye
s (4
9.3%
)
Geo
met
ric s
erie
s N
o (0
%)
No
(0%
) N
o (0
%)
Yes
(28.
8%)
No
(0%
) N
o (0
%)
No
(0%
) Ye
s (9
9.8%
) N
o (0
%)
No
(0%
) N
o (0
%)
No
(0%
) N
o (0
%)
No
(0%
) N
o (0
%)
No
(0%
) N
o (0
%)
No
(0%
)
Brok
en s
tick
No
(0%
) N
o (0
%)
No
(0%
) N
o (0
%)
No
(0%
) N
o (0
%)
No
(0%
) N
o (0
%)
No
(0%
) N
o (0
%)
No
(0%
) N
o (0
%)
No
(0%
) N
o (0
%)
No
(0%
) N
o (0
%)
No
(0%
) N
o (0
%)
Ju
ne 2
001
Augu
st 2
001
All i
slan
ds
Pa
ndan
O
pak
Bira
Pu
tri
Mel
injo
G
ente
ng
Pand
an
Opa
k Bi
ra
Putri
M
elin
jo
Gen
teng
O
ct. 0
0 M
ar. 0
1 Ap
r. 01
Ju
n. 0
1 Au
g. 0
1 A.
Div
ersi
ty in
dice
s S
(Tot
al s
peci
es)
48
36
48
51
48
49
49
45
45
48
44
41
82
84
84
84
81
d (S
peci
es
richn
ess)
6.
42
5.05
6.
47
7.32
6.
82
6.68
6.
55
6.30
6.
52
7.00
6.
24
5.78
9.
44
9.47
9.
29
9.35
9.
15
N (T
otal
in
divi
dual
s)
1504
10
24
1425
93
0 98
7 13
20
1521
10
77
851
822
979
1011
53
10
6431
76
02
7190
62
61
J' (E
venn
ess)
0.
72
0.73
0.
71
0.79
0.
71
0.75
0.
69
0.77
0.
73
0.75
0.
72
0.77
0.
73
0.74
0.
77
0.70
0.
71
H' (
log e
) 2.
80
2.60
2.
74
3.10
2.
73
2.94
2.
67
2.94
2.
77
2.90
2.
72
2.85
3.
23
3.27
3.
40
3.09
3.
11
α D
iver
sity
(Fis
her)
9.46
7.
26
9.58
11
.60
10.5
5 10
.02
9.68
9.
49
10.1
3 11
.12
9.47
8.
58
13.7
6 13
.64
13.2
1 13
.35
13.1
3 B
. Fi
t of D
istr
ibut
ion
Mod
el
Loga
rithm
ic s
erie
s Ye
s (1
8.7%
) Ye
s (5
4.3%
) Ye
s (9
1.5%
) Ye
s (8
3.4%
) Ye
s (1
9.2%
) Ye
s (9
.2%
) Ye
s (1
9.8%
) Ye
s (4
0.2%
) Ye
s (6
8.9%
) Ye
s (9
8.2%
) Ye
s (5
1.2%
) Ye
s (9
7.6%
) Ye
s (1
5.3%
) Ye
s (1
.8%
) N
o (0
%)
Yes
(17.
5%)
Yes
(31.
9%)
Log
norm
al
Yes
(18.
8%)
Yes
(45.
3%)
Yes
(94.
0%)
Yes
(59.
9%)
Yes
(12.
0%)
No
(0%
) Ye
s (2
1.0%
) Ye
s (5
4.4%
) Ye
s (4
5.4%
) Ye
s (7
8.0%
) Ye
s (2
8.8%
) Ye
s (8
4.1%
) Ye
s (3
5.4%
) Ye
s (3
4.9%
) Ye
s (1
8.2%
) Ye
s (7
3.2%
) Ye
s (9
3.2%
) G
eom
etric
ser
ies
No
(0%
) N
o (0
%)
No
(0%
) N
o (0
%)
No
(0%
) N
o (0
%)
No
(0%
) N
o (0
%)
No
(0%
) N
o (0
%)
No
(0%
) N
o (0
%)
No
(0%
) N
o (0
%)
No
(0%
) N
o (0
%)
No
(0%
) Br
oken
stic
k N
o (0
%)
No
(0%
) N
o (0
%)
No
(0%
) N
o (0
%)
No
(0%
) N
o (0
%)
No
(0%
) N
o (0
%)
No
(0%
) N
o (0
%)
No
(0%
) N
o (0
%)
No
(0%
) N
o (0
%)
No
(0%
) N
o (0
%)
47
3.4. FISH DIVERSITY
The Shannon diversity index (H’) (Table 3.2.) was used for the comparison of
fish communities in the Sanctuary Zone (Bira and Putri), the Intensive Utilization
Zone (Melinjo and Genteng), and the Traditional Utilization Zone (Pandan and Opak)
(see Table 3.3 and Table 3.4). Most of the comparison carried out for Bira showed no
significant differences to the other islands (only 8 of 25 comparisons showed
significant differences) (Table 3.3). Though the results of the significance tests do not
allow stating a clear difference between Bira and the other islands in terms of
diversity (measured as H’). Bira seemed to be slightly lower in fish diversity than the
other islands except in April 2001 if compared to Melinjo. Putri seemed to be the
most diverse island in comparison to the others, which can be seen in a significantly
higher value of H’ (Table 3.3). The Shannon diversity index in each island in October
2000 seemed to be lower compared to the following months (Table 3.4). The other
diversity indices were also calculated for fish community (Table 3.2).
TABLE 3.3. The Comparison of the Shannon diversity index (H') between the islands in the core zone (P. KA Bira and P. Putri Timur) and outside the core zone from each sampling time. The t-test was run at a significance level of P<0.001 (n.s.= Not significantly different; s. = Significantly different).
P. Putri Timur (D)
P. Melinjo (E)
P. KA Genteng (F)
P. Pandan (A)
P. Opak Besar (B)
October 2000 s. (C<D) s. (C<E) s. (C<F) s. (C<A) n.s. March 2001 n.s. n.s. n.s. n.s. s. (C<B) April 2001 n.s. s. (C>E) n.s. n.s. n.s. June 2001 s. (C<D) n.s. s. (C<F) n.s. n.s.
P. KA Bira (C)
August 2001 n.s. n.s. n.s. n.s. n.s.
P. Melinjo (E)
P. KA Genteng (F)
P. Pandan (A)
P. Opak Besar (B)
October 2000 n.s. n.s. s. (D>A) s. (D>B) March 2001 n.s. n.s. n.s. s. (D<B) April 2001 s. (D>E) n.s. s. (D>A) n.s. June 2001 s. ((D>E) s. (D>F) s. (D>A) s. (D>B)
P. Putri Timur (D)
August 2001 n.s. n.s. s. (D>A) n.s.
48
TABLE 3.4. Comparison of Shannon diversity index (H') between the sampling times in all islands. The t-test was performed at a significance level of P<0.001 (n.s. = Not significantly different; s. = Significantly different).
P. Pandan (A) (2) March 2001 (3) April 2001 (4) June 2001 (5) August 2001 (1) October 2000 s. (1<2) s. (1<3) s. (1<4) s. (1<5) (2) March 2001 - s. (2<3) n.s. n.s. (3) April 2001 - - n.s. s. (3>5) (4) June 2001 - - - n.s.
P. Opak Besar (B) (2) March 2001 (3) April 2001 (4) June 2001 (5) August 2001 (1) October 2000 s. (1<2) s. (1<3) n.s. s. (1<5) (2) March 2001 - n.s. s. (2>4) s. (2>5) (3) April 2001 - - s. (3>4) s. (3>5) (4) June 2001 - - - s. (4<5)
P. KA Bira (C) (2) March 2001 (3) April 2001 (4) June 2001 (5) August 2001 (1) October 2000 s. (1<2) s. (1<3) s. (1<4) s. (1<5) (2) March 2001 - s. (2<3) n.s. n.s. (3) April 2001 - - s. (3>4) s. (3>5) (4) June 2001 - - - n.s.
P. Putri Timur (D) (2) March 2001 (3) April 2001 (4) June 2001 (5) August 2001 (1) October 2000 n.s. s. (1<3) s. (1<4) n.s. (2) March 2001 - s. (2<3) s. (2<4) n.s. (3) April 2001 - - n.s. s. (3>5) (4) June 2001 - - - s. (4>5)
P. Melinjo (E) (2) March 2001 (3) April 2001 (4) June 2001 (5) August 2001 (1) October 2000 n.s. n.s. n.s. n.s. (2) March 2001 - n.s. n.s. n.s. (3) April 2001 - - s. (3>4) s. (3>5) (4) June 2001 - - - n.s.
P. KA Genteng (F) (2) March 2001 (3) April 2001 (4) June 2001 (5) August 2001 (1) October 2000 s. (1<2) s. (1<3) n.s. n.s. (2) March 2001 - n.s. n.s. n.s. (3) April 2001 - - n.s. s. (3>5) (4) June 2001 - - - n.s.
All islands (2) March 2001 (3) April 2001 (4) June 2001 (5) August 2001 (1) October 2000 n.s. s. (1<3) s. (1>4) s. (1>5) (2) March 2001 - s. (2<3) s. (2>4) s. (2>5) (3) April 2001 - - s. (3>4) s. (3>5) (4) June 2001 - - - n.s.
49
3.5. FISH SPECIES-ABUNDANCE RELATIONSHIP MODEL
The species rank order (sequence) based on their abundances at each island and
all islands combined can be seen in Fig. 3.14 – 3.20. The most abundant species
belong to the families Pomacentridae and Labridae, and in some islands also to the
families Scaridae and Chaetodontidae.
Four main models were examined for the fish species abundance data: the log
series (logarithmic series distribution), the log normal distribution (truncated log
normal), the geometric series and MacArthur’s broken stick distribution model (Table
3.2). All data on fish species abundance was fitted to the log series distribution.
However, all of the data also fitted the log normal distribution except two data sets
(Table 3.2) that only fitted to a geometric series distribution. There was no fish
species data set that fitted to the broken stick distribution model (Table 3.2.).
Most of the data on species abundance fitted the log series and the log normal
distribution. Only two data sets fitted to the log series, the log normal and the broken
stick (with χ2 test, P>0.05). The χ2 value of each species abundance data set was used
to find a higher probability being the same with the model, the higher percentage was
more appropriate to the model (Table 3.2.).
50
0
1
2
3
4
5
6
0 5 10 15 20 25 30 35 40 45 50 55 60
Rank of abundance
Abun
danc
e (ln
)Oct '00Mar '01Apr '01Jun '01Aug '01
Oct '00y = -0.13x + 3.96R2 = 0.85
Mar '01y = -0.10x + 4.38R2 = 0.95
Apr '01y = -0.09x + 4.57R2 = 0.9523
Jun '01y = -0.11x + 4.97R2 = 0.9662
Aug '01y = -0.11x + 4.72R2 = 0.93
0
1
2
3
4
5
6
0 5 10 15 20 25 30 35 40 45 50 55 60
Rank of abundance
Abun
danc
e (ln
)
Oct '00Mar '01Apr '01Jun '01Aug '01
P. PANDAN (A) Time Rank
October 2000 March 2001 April 2001 June 2001 August 2001 1 C. atripectoralis P. lepidogenys P. lepidogenys P. lepidogenys P. lepidogenys 2 C. weberi P. alexanderae C. cyanopleura C. cyanopleura C. cyanopleura 3 N. anabatoides C. cyanopleura P. alexanderae P. alexanderae P. alexanderae 4 T. lunare A. curacao Scarus sp. 1 C. analis Scarus sp. 1 5 A. curacao P. lacrymatus C. octofasciatus C. octofasciatus C. analis 6 P. lacrymatus T. lunare C. analis T. lunare Pomacentrus sp. 1 7 C. analis A. compressus P. lacrymatus A. curacao A. curacao 8 C. octofasciatus P. grammorhyncus A. curacao P. lacrymatus P. lacrymatus 9 Chromis sp. 1 C. analis T. lunare N. nigroris P. grammorhyncus
10 P. grammorhyncus H. melanurus Pomacentrus sp. 1 Pomacentrus sp. 1 C. octofasciatus FIGURE 3.14. Rank abundance plot (a) and linear regression analysis of fish abundance (b) in Pandan (the linear relationship is highly significant, P<0.01). Sampling time was in October 2000, March 2001, April 2001, June 2001 and August 2001. The table shows the ten most abundant species. R2 = coefficient of determination.
(a)
(b)
51
0
1
2
3
4
5
6
0 5 10 15 20 25 30 35 40 45 50 55 60
Rank of abundance
Abun
danc
e (ln
)Oct '00Mar '01Apr '01Jun '01Aug '01
Oct '00y = -0.17x + 5.02R2 = 0.97
Mar '01y = -0.11x + 5.08R2 = 0.99
Apr '01y = -0.10x + 4.88R2 = 0.98
Jun '01y = -0.15x + 4.94R2 = 0.97
Aug '01y = -0.11x + 4.66R2 = 0.97
0
1
2
3
4
5
6
0 5 10 15 20 25 30 35 40 45 50 55 60
Rank of abundance
Abun
danc
e (ln
)
Oct '00Mar '01Apr '01Jun '01Aug '01
P. OPAK BESAR (B)
Time Rank October 2000 March 2001 April 2001 June 2001 August 2001
1 P. lepidogenys P. lepidogenys P. lepidogenys C. cyanopleura P. lepidogenys 2 C. atripectoralis P. alexanderae P. amboinensis P. lepidogenys C. cyanopleura 3 H. argus C. cyanopleura C. octofasciatus C. analis Scarus sp. 1 4 C. cyanopleura A. vaigiensis P. grammorhyncus A. curacao C. analis 5 A. curacao Scarus sp. 1 A. curacao P. alexanderae P. grammorhyncus 6 C. analis C. analis N. nigroris P. grammorhyncus A. curacao 7 T. lunare A. curacao C. analis C. octofasciatus C. octofasciatus 8 C. octofasciatus C. octofasciatus T. lunare A. leucogaster N. nigroris 9 A. vaigiensis H. argus C. fasciatus N. nigroris S. sordidus
10 N. oxyodon P. grammorhyncus Scarus sp. 1 A. compressus H. melanurus FIGURE 3.15. Rank abundance plot (a) and linear regression analysis of fish abundance (b) in Opak (the linear relationship is highly significant, P<0.01). Sampling time was in October 2000, March 2001, April 2001, June 2001 and August 2001. The table shows the ten most abundant species. R2 = coefficient of determination.
(a)
(b)
52
0
1
2
3
4
5
6
0 5 10 15 20 25 30 35 40 45 50 55 60
Rank of abundance
Abun
danc
e (ln
)
Oct '00Mar '01Apr '01Jun '01Aug '01
Oct '00y = -0.16x + 4.82R2 = 0.97
Mar '01y = -0.09x + 4.10R2 = 0.93
Apr '01y = -0.09x + 4.51R2 = 0.97
Jun '01y = -0.11x + 4.83R2 = 0.96
Aug '01y = -0.11x + 4.33R2 = 0.94
0
1
2
3
4
5
6
0 5 10 15 20 25 30 35 40 45 50 55 60
Rank of abundance
Abun
danc
e (ln
)
Oct '00Mar '01Apr '01Jun '01Aug '01
P. KA BIRA (C)
Time Rank October 2000 March 2001 April 2001 June 2001 August 2001
1 H. argus P. alexanderae P. alexanderae P. alexanderae P. alexanderae 2 A. curacao A. curacao C. cyanopleura C. cyanopleura Scarus sp. 1 3 P. lepidogenys A. leucogaster A. curacao A. curacao A. curacao 4 C. atripectoralis N. nigroris P. grammorhyncus Scarus sp. 1 N. nigroris 5 C. analis C. analis N. nigroris A. leucogaster P. grammorhyncus 6 P. alexanderae P. lepidogenys P. lepidogenys C. analis C. analis 7 P. grammorhyncus C. octofasciatus A. leucogaster P. lepidogenys A. leucogaster 8 C. octofasciatus C. cyanopleura T. lunare N. nigroris T. lunare 9 T. lunare Scarus sp. 1 C. analis P. grammorhyncus C. cyanopleura
10 N. melas P. grammorhyncus C. quinquelineatus H. purpurascens C. octofasciatus FIGURE 3.16. Rank abundance plot (a) and linear regression analysis of fish abundance (b) in Bira (the linear relationship is highly significant, P<0.01). Sampling time was in October2000, March 2001, April 2001, June 2001 and August 2001. The table shows the ten most abundant species. R2 = coefficient of determination.
(a)
(b)
53
0
1
2
3
4
5
6
0 5 10 15 20 25 30 35 40 45 50 55 60
Rank of abundance
Abun
danc
e (ln
)
Oct '00Mar '01Apr '01Jun '01Aug '01
Oct '00y = -0.14x + 4.62R2 = 0.97
Mar '01y = -0.11x + 4.31R2 = 0.96
Apr '01y = -0.10x + 4.66R2 = 0.97
Jun '01y = -0.09x + 4.37R2 = 0.9758
Aug '01y = -0.11x + 4.29R2 = 0.95
0
1
2
3
4
5
6
0 5 10 15 20 25 30 35 40 45 50 55 60
Rank of abundance
Abun
danc
e (ln
)
Oct '00Mar '01Apr '01Jun '01Aug '01
P. PUTRI TIMUR (D)
Time Rank October 2000 March 2001 April 2001 June 2001 August 2001
1 C. atripectoralis C. cyanopleura C. cyanopleura C. cyanopleura C. cyanopleura 2 C. cyanopleura N. anabatoides P. alexanderae Scarus sp. 1 P. alexanderae 3 H. argus P. alexanderae N. anabatoides N. nigroris N. nigroris 4 T. lunare A. curacao Scarus sp. 1 P. alexanderae Pomacentrus sp. 1 5 A. curacao C. analis A. leucogaster Pomacentrus sp. 1 Scarus sp. 1 6 C. fasciatus N. nigroris N. nigroris P. lacrymatus A. curacao 7 C. analis C. fasciatus C. fasciatus A. curacao C. analis 8 L. dimidiatus A. leucogaster A. curacao T. lunare C. fasciatus 9 P. alexanderae C. octofasciatus C. analis C. fasciatus T. lunare
10 M. ornatus T. lunare P. amboinensis C. analis P. lacrymatus FIGURE 3.17. Rank abundance plot (a) and linear regression analysis of fish abundance (b) in Putri (the linear relationship is highly significant, P<0.01). Sampling time was in October 2000, March 2001, April 2001, June 2001 and August 2001. The table shows the ten most abundant species. R2 = coefficient of determination.
(a)
(b)
54
0
1
2
3
4
5
6
0 5 10 15 20 25 30 35 40 45 50 55 60
Rank of abundance
Abun
danc
e (ln
)Oct '00Mar '01Apr '01Jun '01Aug '01
Oct '00y = -0.12x + 4.49R2 = 0.94
Mar '01y = -0.10x + 4.41R2 = 0.94
Apr '01y = -0.09x + 4.58R2 = 0.95
Jun '01y = -0.11x + 4.39R2 = 0.9422
Aug '01y = -0.11x + 4.46R2 = 0.95
0
1
2
3
4
5
6
0 5 10 15 20 25 30 35 40 45 50 55 60
Rank of abundance
Abun
danc
e (ln
)
Oct '00Mar '01Apr '01Jun '01Aug '01
P. MELINJO (E)
Time Rank October 2000 March 2001 April 2001 June 2001 August 2001
1 P. alexanderae C. cyanopleura P. alexanderae P. alexanderae C. cyanopleura 2 H. melanurus P. alexanderae A. vaigiensis C. cyanopleura P. alexanderae 3 H. argus A. vaigiensis C. cyanopleura C. analis T. lunare 4 C. analis N. nigroris A. sexfasciatus T. lunare N. nigroris 5 T. lunare C. analis Scarus sp. 1 N. nigroris A. curacao 6 A. curacao T. lunare C. analis A. curacao H. melanurus 7 A. sexfasciatus A. curacao N. nigroris C. octofasciatus Pomacentrus sp. 1 8 C. octofasciatus H. melanurus Pomacentrus sp. 1 Pomacentrus sp. 1 C. analis 9 N. nigroris C. octofasciatus C. atripectoralis A. leucogaster A. leucogaster
10 H. hortulanus P. trivittatus T. lunare P. amboinensis A. compressus FIGURE 3.18. Rank abundance plot (a) and linear regression analysis of fish abundance (b) in Melinjo (the linear relationship is highly significant, P<0.01). Sampling time was in October 2000, March 2001, April 2001, June 2001 and August 2001. The table shows the ten most abundant species. R2 = coefficient of determination.
(a)
(b)
55
0
1
2
3
4
5
6
0 5 10 15 20 25 30 35 40 45 50 55 60
Rank of abundance
Abun
danc
e (ln
)Oct '00Mar '01Apr '01Jun '01Aug '01
Oct '00y = -0.15x + 5.27R2 = 0.9739
Mar '01y = -0.11x + 4.76R2 = 0.98
Apr '01y = -0.11x + 4.97R2 = 0.99
Jun '01y = -0.12x + 4.90R2 = 0.95
Aug '01y = -0.13x + 4.80R2 = 0.98
0
1
2
3
4
5
6
0 5 10 15 20 25 30 35 40 45 50 55 60
Rank of abundance
Abun
danc
e (ln
)
Oct '00Mar '01Apr '01Jun '01Aug '01
P. KA GENTENG (F)
Time Rank October 2000 March 2001 April 2001 June 2001 August 2001
1 P. lepidogenys P. lepidogenys A. curacao P. lepidogenys P. lepidogenys 2 A. sexfasciatus A. curacao P. lepidogenys A. curacao C. cyanopleura 3 A. vaigiensis N. anabatoides A. sexfasciatus P. alexanderae N. anabatoides 4 C. atripectoralis T. lunare C. analis C. cyanopleura C. analis 5 C. viridis P. alexanderae T. lunare A. sexfasciatus P. alexanderae 6 N. anabatoides A. leucogaster C. octofasciatus N. anabatoides A. compressus 7 H. argus C. analis C. atripectoralis C. analis C. octofasciatus 8 M. ornatus A. sexfasciatus N. nigroris T. lunare N. nigroris 9 A. curacao N. nigroris P. grammorhyncus C. octofasciatus A. curacao
10 C. analis C. octofasciatus C. cyanopleura N. nigroris T. lunare FIGURE 3.19. Rank abundance plot (a) and linear regression analysis of fish abundance (b) in Genteng (the linear relationship is highly significant, P<0.01). Sampling time was in October 2000, March 2001, April 2001, June 2001 and August 2001. The table shows the ten most abundant species. R2 = coefficient of determination.
(a)
(b)
56
0
1
2
3
4
5
6
7
8
0 5 10 15 20 25 30 35 40 45 50 55 60 65 70 75 80 85 90
Rank of abundance
Abun
danc
e (ln
)Oct '00Mar '01Apr '01Jun '01Aug '01
Oct '00y = -0.08x + 5.89R2 = 0.97
Mar '01y = -0.07x + 5.97R2 = 0.98
Apr '01y = -0.08x + 6.35R2 = 0.99
Jun '01y = -0.08x + 6.05R2 = 0.9784
Aug '01y = -0.08x + 5.92R2 = 0.98
0
1
2
3
4
5
6
7
8
0 5 10 15 20 25 30 35 40 45 50 55 60 65 70 75 80 85 90
Rank of abundance
Abun
danc
e (ln
)
Oct '00Mar '01Apr '01Jun '01Aug '01
ALL ISLANDS
Time Rank October 2000 March 2001 April 2001 June 2001 August 2001
1 H. argus P. alexanderae P. alexanderae C. cyanopleura C. cyanopleura 2 C. atripectoralis C. cyanopleura P. lepidogenys P. alexanderae P. lepidogenys 3 P. lepidogenys P. lepidogenys C. cyanopleura P. lepidogenys P. alexanderae 4 A. curacao A. curacao A. curacao A. curacao Scarus sp. 1 5 C. analis C. analis Scarus sp. 1 C. analis C. analis 6 T. lunare T. lunare C. analis N. nigroris A. curacao 7 P. alexanderae N. nigroris N. nigroris T. lunare N. nigroris 8 N. anabatoides A. vaigiensis C. octofasciatus C. octofasciatus T. lunare 9 A. sexfasciatus C. octofasciatus T. lunare Scarus sp. 1 Pomacentrus sp. 1
10 C. octofasciatus N. anabatoides A. sexfasciatus A. leucogaster P. grammorhyncus FIGURE 3.20. Rank abundance plot (a) and linear regression analysis of fish abundance (b) in all islands (the linear relationship is highly significant, P<0.01). Sampling time was in October 2000, March 2001, April 2001, June 2001 and August 2001. The table shows the ten most abundant species. R2 = coefficient of determination.
(a)
(b)
57
3.6. FISH COMMUNITY STRUCTURE
In general the fish community structure in all surveyed islands could be
separated into two groups: one from west monsoon and another from the east
monsoon. The cluster analysis based on the Bray-Curtis similarity of all fish species
from all islands displayed three different groups at the 56 % similarity level (Fig.
3.21). The first group considered solely the fish community in Pandan in October
2000. The second group was the fish community in October 2000 from all islands,
except Pandan. The third group was the fish community from the following
observations.
The result of NMDS indicated two different groups of fish species composition
and community structure between west and east monsoon (Fig. 3.22). In this analysis,
the stress value was 0.16, which indicates in fair condition to interpreted.
A PCA-plot of fish community with 30.6 % variation in PC-1 and 23.9 % in
PC-2 (Fig. 3.23) gave a different pattern compared to the dendrogram (Fig. 3.21) and
the NMDS-plot (Fig. 3.22). In the first quadrant can be found a fish community from
all islands during west monsoon, except Opak. In the three other quadrants were the
fish communities from east monsoon. They were split into three different groups thus
showing another compositions and pattern of fish communities in each quadrant. In
the first quadrant, the fish community was mainly characterized Chromis
atripectoralis, Halichoeres argus (see Appendix 3). The second quadrant was
characterized by Abudefduf vaigiensis, Amblyglyphidodon leucogaster and
Neoglyphidodon nigroris. The third quadrant was more dominated by Cirrhilabrus
cyanopleura, Pomacentrus alexanderae, Scarus and Plectroglyphidodon lacrymatus
and the forth quadrant by Pomacentrus lepidogenys, Pomacentrus grammorhyncus,
Chromis analis and Chaetodon octofasciatus.
58
A-O
ct '0
0
E-O
ct '0
0
F-O
ct '0
0
B-O
ct '0
0
C-O
ct '0
0
D-O
ct '0
0
B-Ap
r '01
F-Au
g '0
1
F-M
ar '0
1
F-Ap
r '01
F-Ju
n '0
1
B-M
ar '0
1
B-Ju
n '0
1
B-Au
g '0
1
A-M
ar '0
1
A-Au
g '0
1
A-Ap
r '01
A-Ju
n '0
1
C-A
pr '0
1
C-A
ug '0
1
C-M
ar '0
1
C-J
un '0
1
E-M
ar '0
1
E-Ap
r '01
E-Ju
n '0
1
E-Au
g '0
1
D-M
ar '0
1
D-A
pr '0
1
D-J
un '0
1
D-A
ug '0
1
100
80
60
40
20
Sim
ilarit
y
FIGURE 3.21. Dendrogram of hierarchical clustering with group linkage methods of the fish community, based on species abundance. Three replicate samples were made from each island at each sampling. (A=Pandan, B=Opak, C=Bira, D=Putri, E=Melinjo, F=Genteng).
A-Oct '00
A-Mar '01
A-Apr '01 A-Jun
A-Aug '01
B-Oct '00 B-Mar '01
B-Apr '01 B-Jun '01
B-Aug '01
C-Oct '00
C-Mar '01 C-Apr '01 C-Jun '01
C-Aug '01
D-Oct '00
D-Mar '01
D-Apr '01 D-Jun '01
D-Aug '01
E-Oct '00
E-Mar '01 E-Apr '01
E-Jun '01 E-Aug '01
F-Oct '00
F-Mar '01 F-Apr '01
F-Jun '01 F-Aug '01
Stress: 0.16
East Monsoon West Monsoon
FIGURE 3.22. Non-metric multidimensional scaling ordination of the fish community based on species abundance. Three replicate samples were made for each island at each sampling. (A=Pandan, B=Opak, C=Bira, D=Putri, E=Melinjo, F=Genteng).
West Monsoon East Monsoon
59
A-Oct '00
A-Mar '01
A-Apr '01
A-Jun '01
A-Aug '01
B-Oct '00B-Mar '01
B-Apr '01
B-Jun '01B-Aug '01
C-Oct '00C-Mar '01
C-Apr '01
C-Jun '01
C-Aug '01D-Oct '00
D-Mar '01D-Apr '01
D-Jun '01D-Aug '01
E-Oct '00
E-Mar '01
E-Apr '01
E-Jun '01
E-Aug '01 F-Oct '00
F-Mar '01F-Apr '01F-Jun '01
F-Aug '01
-12
-10
-8
-6
-4
-2
0
2
4
6
8
10
-10 -8 -6 -4 -2 0 2 4 6 8 10
PC-2: 23.9 %
PC-1: 30.6 %
FIGURE 3.23. PCA-plot of fish communities based on species abundance. Three replicate samples were made for each island at each sampling. (A=Pandan, B=Opak, C=Bira, D=Putri, E=Melinjo, F=Genteng).
The PCA-biplot of the trophic fish group displayed the change in fish
composition throughout the survey periods (Fig. 3.24). The starting point of the fish
community was in October 2000 for each island, then continued to the following
months, March – August 2001. In general the fish community returned again to a
similar composition at the beginning (see the arrow direction of each fish community
from each island in Fig. 3.24.).
60
PCA-biplot of fish trophic group
F-Aug
F-JunF-Apr
F-Mar
F-Oct
E-Aug
E-Jun
E-Apr
E-Mar
E-Oct
D-Aug
D-Jun
D-Apr D-Mar
D-Oct
C-Aug
C-Jun
C-Apr
C-Mar
C-OctB-Aug
B-Jun
B-Apr
B-MarB-Oct
A-AugA-Jun
A-Apr
A-Mar
A-OctPiscivore
Benthic feeder
Corallivore
Detritivore
Planktivore
Omnivore
Herbivore
-20
-15
-10
-5
0
5
10
-20 -15 -10 -5 0 5 10 15 20 25
PC-2: 31.0 %
PC-1: 52.7
FIGURE 3.24. PCA-biplot of trophic group of fish produced by SVD method. The sampling times were October 2000 and March, April, June, and August 2001. (A=Pandan, B=Opak, C=Bira, D=Putri, E=Melinjo, F=Genteng).
61
3.7. RELATING BENTHIC HABITAT AND FISH COMMUNITY STRUCTURE
CCA ordination plots were created several times and then four species of fish
were selected that had relatively strong relationships with certain life form categories
(environmental variables) (Fig. 3.25). The selected fish species were Chaetodon
octofasciatus (Ctoc), Chromis analis (Cran), Pomacentrus alexanderae (Pmal), and
Pomacentrus lepidogenys (Pmle). The CCA eigenvalue of the first axis was 0.28
(explaining 68.5 % of the variance), the second axis had an eigenvalue 0.08
(explaining 18.3 % of the variance), and the sum of all CCA eigenvalues was 0.41.
The CCA ordination plot are interpreted by means of the centroid principle, the
distance rule, the biplot rule, and the biplot rule for compositional data (ter Braak &
Verdonschot 1995). Using the centroid principle, the sites close to the species point
then described to have the higher relationship than the sites far from the species point
(ter Braak & Verdonschot 1995). In Genteng and Opak, Chaetodon octofasciatus was
more abundant in April, June and August 2001; Chromis analis in August 2001, and
Pomacentrus lepidogenys in October 2000, compared to the other islands. In Pandan,
P. lepidogenys was most abundant in March, April, June and August 2001. In Bira C.
analis occurred in high abundance in October 2000, March, April, and June 2001,
while C. octofasciatus was most abundant in October 2000. In Melinjo, Pomacentrus
alexanderae was more abundant in April, June and August 2001. However, the
centroid rule creates good results when the eigenvalues had at least a value of 0.4 (ter
Braak & Verdonschot 1995).
62
CCA-
trip
lot o
f sel
ecte
d fis
h sp
ecie
s an
d lif
e fo
rm c
ateg
orie
s
Cto
c-Oc
Cra
n-Oc
Pmal-Oc
Pmle-Oc
Cto
c-Ma
Cra
n-Ma
Pmal-Ma
Pmle-Ma
Cto
c-Ap
Cra
n-Ap
Pmal-Ap
Pmle-Ap
Cto
c-Ju
Cra
n-Ju
Pmal-Ju
Pmle-Ju
Cto
c-Au
Cra
n-Au
Pmal-Au
Pmle-Au
ACB
ACD
ACT
CB
CECFCM
CMR
CS
CME
CHL
OT
AL
DC
P. P
anda
n (A
)
P. O
pak
Besa
r (B)
P. K
A Bi
ra (C
) P.
Put
ri Ti
mur
(D)
P. M
elin
jo (E
)
P. K
A G
ente
ng (F
)
-2.0
-1.5
-1.0
-0.50.0
0.5
1.0
1.5
-1.5
-1.0
-0.5
0.0
0.5
1.0
1.5
2.0
FI
GU
RE
3.2
5.
CC
A-tr
iplo
t of t
he d
istri
butio
n of
sel
ecte
d fis
h-sp
ecie
s fo
und
durin
g O
ctob
er 2
000-
Aug
ust 2
001
in s
ix is
land
s: fi
sh s
peci
es (s
olid
circ
le),
life
form
and
ben
thic
var
iabl
es (h
ollo
w c
ircle
), an
d th
e is
land
s (s
olid
squ
are)
. The
ben
thic
var
iabl
es w
ere:
Acr
opor
a B
ranc
hing
(AC
B),
Acro
pora
Dig
itate
(AC
D),
Acro
pora
Tab
ulat
e (A
CT)
, Cor
al B
ranc
hing
(C
B),
Cor
al E
ncru
stin
g (C
E), C
oral
Fol
iose
(C
F), C
oral
Mas
sive
(C
M),
Cor
al S
ub-m
assi
ve (
CS)
, Mus
hroo
m
Cor
al (
CM
R),
Mill
epor
a (C
ME)
, Hel
iopo
ra (
CH
L), O
ther
Fau
na (
OT)
, Alg
ae (
AL)
, and
Dea
d C
oral
(D
C).
The
fish
spec
ies
wer
e C
haet
odon
oct
ofas
ciat
us
(Cto
c), C
hrom
is a
nalis
(C
ran)
, Pom
acen
trus
ale
xand
erae
(Pm
al),
and
Pom
acen
trus
lepi
doge
nys
(Pm
le).
The
sam
plin
g tim
es w
ere
Oct
ober
200
0 (O
c), a
nd
Mar
ch (M
a), A
pril
(Ap)
, Jun
e (J
u) a
nd A
ugus
t 200
1 (A
u).
63
CC
A-tr
iplo
t of s
elec
ted
fish
spec
ies
and
life
form
cat
egor
ies
Cto
c-Oc
Cra
n-Oc
Pmal-Oc
Pmle-Oc
Cto
c-Ma
Cra
n-Ma
Pmal-Ma
Pmle-Ma
Cto
c-Ap
Cra
n-Ap
Pmal-Ap
Pmle-Ap
Cto
c-Ju
Cra
n-Ju
Pmal-Ju
Pmle-Ju
Cto
c-Au
Cra
n-Au
Pmal-Au
Pmle-Au
ACB
ACD
ACT
CB
CE
CF
CM
CM
R
CS
CM
EC
HL
OT
AL
DC
P. P
anda
n (A
)
P. O
pak
Besa
r (B)
P. K
A Bi
ra (C
) P.
Put
ri Ti
mur
(D)
P. M
elin
jo (E
)
P. K
A G
ente
ng (F
)
-2.0
-1.5
-1.0
-0.50.0
0.5
1.0
1.5
-1.5
-1.0
-0.5
0.0
0.5
1.0
1.5
2.0
FI
GU
RE
3.2
6. C
CA
-trip
lot o
f the
dis
tribu
tion
of s
elec
ted
fish-
spec
ies
foun
d du
ring
Oct
ober
200
0-A
ugus
t 200
1 in
six
isla
nds:
fish
spe
cies
(sol
id c
ircle
), lif
e fo
rm a
nd b
enth
ic v
aria
bles
(ho
llow
circ
le),
and
the
isla
nds
(sol
id s
quar
e).
An
arro
w (
dash
line
) w
as p
roje
cted
alo
ng th
e Ac
ropo
ra B
ranc
hing
var
iabl
e th
at
indi
catin
g a
grad
ient
; the
per
pend
icul
ar d
ash
line
in th
e ar
row
indi
cate
d th
e po
sitio
n of
the
isla
nds a
long
this
gra
dien
t. (R
efer
to F
igur
e 3.
25 fo
r abb
revi
atio
ns).
64
Since the first two eigenvalues of CCA (Fig. 3.25) were low, the biplot rule was
also used, as it would be more informative (Gabriel 1971, 1982; ter Braak &
Verdonschot 1995). The environmental variable “ Acropora Branching” was chosen
and then an arrow was projected along the line (Fig. 3.26). The projected arrow
showed that Opak had the highest cover of “Acropora Branching” being followed by
Genteng. The lowest coverage was in Melinjo, as the variable does not change in
value in the perpendicular direction (ter Braak & Verdonschot 1995). However, the
ordination plot did not display the data table (Appendix 1) exactly, because the plot
uses only two dimensions whereas the data table is multidimensional (ter Braak &
Verdonschot 1995). The projected arrow of the ACB gradient showed that P.
lepidogenys and C. octofasciatus were more abundant when the value of ACB
coverage was higher. Vice versa, C. analis and P. alexanderae were more abundant
in the area with low coverage of ACB.
The length of the line of life form categories (environmental variables) can be
used to indicate the importance of the variable (ter Braak & Verdonschot 1995).
Therefore the Acropora Branching variable was the most important Acropora life
form, because this variable had the longest line (Fig. 3.26).
The CCA of the five most abundant fish families (Fig. 3.27) showed that the
first axis had a CCA eigenvalue of 0.04 (37.2 % of the variance) and the second axis
had 0.03 (26.1 % of the variance) summing-up to an eigenvalue was 0.10. The
projected arrow along the Dead Coral (DC) variable showed that Putri had the highest
weighted value of dead coral coverage, followed by Melinjo, Opak, Pandan, Bira and
Genteng. The most abundant fish families at the DC high coverage were Labridae,
Nemipteridae and Scaridae. In contrast, Chaetodontidae and Pomacentridae
65
CC
A-tr
iplo
t of m
ost a
bund
ant f
ish
fam
ily a
nd li
fe fo
rm c
ateg
orie
s
Cha-Oc
Poc-Oc
Lab-Oc
Sca-Oc
Nem-Oc
Cha-Ma
Poc-Ma
Lab-Ma
Sca-Ma
Nem-Ma
Cha-Ap
Poc-Ap
Lab-Ap
Sca-Ap
Nem-Ap
Cha-Ju
Poc-Ju
Lab-JuSca-Ju
Nem-Ju
Cha-Au
Poc-Au
Lab-Au
Sca-Au
Nem-Au
ACB
ACD AC
T
CB
CE
CF
CM
CM
R
CS
CM
E
CH
L
OT
AL
DC
P. P
utri
Tim
ur (D
)
P. O
pak
Besa
r (B)
P. K
A Bi
ra (C
)
P. M
elin
jo (E
)
P. P
anda
n (A
)
P. K
A G
ente
ng (F
)
-3-2-10123
-4-3
-2-1
01
23
4
FI
GU
RE
3.2
7.
CC
A-tr
iplo
t of
mos
t abu
ndan
t of
fish-
fam
ilies
fro
m O
ctob
er 2
000-
Aug
ust 2
001
in s
ix is
land
s: f
ish
fam
ilies
(so
lid c
ircle
), lif
e fo
rm a
nd
bent
hic
varia
bles
(ho
llow
circ
le),
and
the
isla
nds
(sol
id tr
iang
le).
The
bent
hic
varia
bles
wer
e: A
crop
ora
Bra
nchi
ng (
AC
B),
Acro
pora
Dig
itate
(A
CD
), Ac
ropo
ra T
abul
ate
(AC
T), C
oral
Bra
nchi
ng (C
B),
Cor
al E
ncru
stin
g (C
E), C
oral
Fol
iose
(CF)
, Cor
al M
assi
ve (C
M),
Cor
al S
ub-m
assi
ve (C
S), M
ushr
oom
C
oral
(CM
R),
Mill
epor
a (C
ME)
, Hel
iopo
ra (C
HL)
, Oth
er F
auna
(OT)
, Alg
ae (A
L), a
nd D
ead
Cor
al (D
C).
The
fish
fam
ilies
wer
e: P
omac
entri
dae
(Poc
), La
brid
ae (
Pmal
), Sc
arid
ae (
Sca)
, Cha
etod
ontid
ae (C
ha) a
nd N
emip
terid
ae (N
em).
The
sam
plin
g tim
es w
ere
Oct
ober
200
0 (O
c), a
nd M
arch
(M
a), A
pril
(Ap)
, Jun
e (J
u) a
nd A
ugus
t 200
1 (A
u).
66
dominated islands with low DC coverage. However, these families preferred islands
with high coverage of ACB and CF (Coral Foliose) (Fig. 3.27).
The CCA ordination plot of trophic group of fish (Fig. 3.28 showed that the first
axis had a CCA eigenvalue of 0.06 (explaining 43.6 % of the variance), the second
axis had 0.03 (explaining 19.6 % of the variance), and the sum of eigenvalues was
0.14. The projected arrow along the DC gradient displayed the highest DC coverage
in Bira followed by Putri, Pandan, Opak, Melinjo and Genteng. Herbivores were
most abundant in those islands that had a high cover of DC, since the Algae category
(AL) was positively correlated with DC. The planktivores occurred in high numbers
in islands where DC coverage was low, but where the cover of Coral Branching (CB),
Acropora Tabulate (ACT) and Coral Encrusting (CE) were high (Opak). The
piscivores preferred the same environmental parameter as the planktivores. The
carnivores preferred islands like Genteng and Melinjo, as these were mostly covered
by CF, ACB and CHL (Heliopora). Benthic feeders and detritivores had no specific
preference for any life form variable. However, the group of omnivores preferred
islands that were mostly covered by other fauna (OT), coral sub-massive (CS),
mushroom coral (CMR) and Millepora (CME).
67
CC
A-tr
iplo
t of t
roph
ic g
roup
of f
ish
and
life
form
cat
egor
ies
DC
ALO
T
CHLCM
ECS
CMR
CM
CFCE
CBACT
ACD
ACB
Pi-Au
B-Au
C-Au
D-Au
P-Au
O-Au
H-Au
Pi-Ju
B-Ju
C-Ju
D-Ju
P-Ju
O-Ju
H-Ju
Pi-Ap
B-Ap
C-Ap
D-Ap
P-Ap
O-Ap
H-Ap
Pi-MaB-Ma
C-Ma
D-Ma
P-Ma
O-Ma
H-Ma
Pi-Oc
B-Oc
C-Oc
P-Oc
O-Oc
H-Oc
P. K
A G
ente
ng (F
)
P. P
anda
n (A
)
P. M
elin
jo (E
)
P. K
A Bi
ra (C
)
P. O
pak
Besa
r (B)
P. P
utri
Tim
ur (D
)
-2.0
-1.5
-1.0
-0.50.0
0.5
1.0
1.5
2.0
-3.0
-2.5
-2.0
-1.5
-1.0
-0.5
0.0
0.5
1.0
1.5
2.0
FI
GU
RE
3.2
8. C
CA
-trip
lot o
f tro
phic
gro
ups
of fi
sh fo
und
from
Oct
ober
200
0-A
ugus
t 200
1 in
six
isla
nds:
fish
fam
ilies
(sol
id c
ircle
), lif
e fo
rm a
nd b
enth
ic
varia
bles
(ho
llow
circ
le),
and
the
isla
nds
(sol
id t
riang
le).
The
bent
hic
varia
bles
wer
e: A
crop
ora
Bra
nchi
ng (
AC
B),
Acro
pora
Dig
itate
(A
CD
), Ac
ropo
ra
Tabu
late
(AC
T), C
oral
Bra
nchi
ng (C
B),
Cor
al E
ncru
stin
g (C
E), C
oral
Fol
iose
(CF)
, Cor
al M
assi
ve (C
M),
Cor
al S
ub-m
assi
ve (C
S), M
ushr
oom
Cor
al (C
MR
), M
illep
ora
(CM
E), H
elio
pora
(CH
L), O
ther
Fau
na (O
T), A
lgae
(AL)
, and
Dea
d C
oral
(DC
). Tr
ophi
c gr
oups
of f
ish:
her
bivo
re (H
), om
nivo
re (O
), pl
ankt
ivor
e (P
), de
tritiv
ore
(D),
bent
hic
feed
er (B
), co
raliv
ore
(C) a
nd p
isci
vore
(Pi).
The
sam
plin
g tim
es w
ere
Oct
ober
200
0 (O
c), a
nd M
arch
(Ma)
, Apr
il (A
p), J
une
(Ju)
an
d A
ugus
t 200
1 (A
u).
68
4. DISCUSSION
The coral reef fish communities in six islands were studied in three different
management zones of the Kepulauan Seribu Marine National Park between October
2000 and August 2001. The study focused on the assessment of the distribution of
fish communities in a coral reef area and the response to blast fishing activities that
had ceased five years ago. Univariate and multivariate analysis tools were used to
analyse the fish communities, the reef structure and relationships between them.
4.1. VARIATION IN CORAL REEF COVERAGE ALONG THE GRADIENT OF BLAST
FISHING IMPACT
The impact of blast fishing on coral reefs was reflected by the presence of many
fields of dead coral, particularly dead coral rubble, throughout the Kepulauan Seribu.
The expected smallest blast fishing impact on coral reef coverage was in the
Sanctuary Zone in which any activities are prohibited, followed by the Intensive
Utilization Zone and the Traditional Utilization Zone. Surprisingly, the percent coral
cover in the Sanctuary Zone, Bira and Putri islands, in fact was the lowest and this
zone could be classified as in ‘bad’ condition in coral cover (according to Gomez &
Alcala 1984), with 19.6 % and 7.6 % for Bira and Putri, respectively (Fig. 3.1). In
contrast, coral coverage in the Intensive Utilization Zone, Melinjo (25.0 %) and
Genteng (42.75 %), can be classified as in ‘fair’ condition. Even in Pandan (29.1 %),
located in the Traditional Utilization Zone and expected to have the lowest coral
coverage, the coral coverage was significantly higher when compared to Sanctuary
Zone islands. Coral coverage in Opak (Traditional Utilization Zone) was in ‘bad’
condition (18.2 %), as expected. Thus in term of hard coral coverage the islands can
69
be ranked such as: (1) Genteng (42.8 %), (2) Pandan (29.1 %), (3) Melinjo (25.1 %),
(4) Bira (19.6 %), (5) Opak (18.2 %) and (6) Putri (7.6 %).
In this study, dead coral rubble was found to cover the largest part of the study
area (Fig. 3.4). Most of the live hard corals of all islands grew on substrate with coral
rubble underlying them. This fact is a strong indication that blast fishing happened
many times throughout all islands before 1995.
All multivariate exploratory techniques showed the same tendency in
separation of the islands into their geographic position: all islands were grouped into
‘west side’ and ‘east side’ (see map on Fig. 2.1). Cluster analysis was not successful
to group islands, according to the expected impact gradient of blast fishing or the
zoning management (Fig. 3.6). Cluster analysis displayed Bira and Putri in a group as
expected (the Sanctuary Zone), but for other islands the dendrogram did not show a
clear pattern in the zoning management as expected. NMDS-plot (Fig. 3.7) displayed
a clearer tendency to separate the islands based on their benthic categories
composition. NMDS technique displayed the island groups according to the zoning
management, but for the Traditional Utilization Zone was only Opak.
But, although in this case the NMDS ordination might be interpreted ‘perfectly’,
the PCA-ordination depicted better in the grouping of the islands. The grouping of
the islands with PCA technique gave a clear separation based on the composition of
life form categories (Fig. 3.8, Appendix 1). Bira and Putri (Sanctuary Zone), which
had a high cover of dead corals rubble (Fig. 3.4) grouped as different,. Genteng and
Pandan had a high cover of foliose corals as another group. The PCA-plot (Fig. 3.8)
also gave another possibility to be interpreted: the PC-2 (principal component)
separated the islands into two geographic groups, the ‘northern part’ (Bira, Putri and
70
Melinjo) and the ‘southern part’(Pandan, Opak and Genteng). This possibility will be
discussed further together with the fish community in the next sub-chapter.
These results confirm previous studies; De Vantier et al. (1998) found the
highest coral cover in Kepulauan Seribu in 1985 to be around 30 %. In 1995 these
authors found a decrease of coral cover due to the blast fishing practices, temperature
stress associated with ENSO events and from pollution. The coral cover in P.
Belanda (the Sanctuary Zone, close to Bira) in 1985 was 39.7 %, but in the pre-survey
of this study only less than 10 % was recorded. Russ & Alcala (1989) noted that blast
fishing and drive-net fishing reduced live coral cover in reserve areas in Sumilon
Island (the Philippines) from 50 to 25 %. And McManus et al. (1997) found 10-30 %
hard coral cover and 60 % dead coral cover in a former blast fishing area in the
Philippines.
Hutomo (1987) and Edinger et al. (1998) noted that coral coverage in
Kepulauan Seribu was positively correlated with distance from the mainland. This,
however, was not the case in the present study; since the hard coral cover sequence
(from on- to offshore) was 29.1 % (Pandan), 18.1 % (Opak), 42.7 % (Genteng), 19.6
% (Bira), 7.6 % (Putri) and 25.1 % (Melinjo).
Since the coral reef as a substrate is biologically generated and coral growth,
form and distribution are influenced by many factors (Luckhurst & Luckhurst 1978b),
we expect that corals would recover after several years of no blast fishing. In general,
there was a high recruitment at all islands; with at least 80 % of the colonies were the
new recruits (Fig. 3.5, Appendix 2). However, the new recruits lacked a pattern along
the impact gradient of blast fishing or the zoning management. All islands had almost
the same percentage of coral colonies in each size category. This fact relates with the
previous finding that the coral rubble is an unstable substrate that can move several
71
centimeters per day, depending on the current speed and rubble fields may even
inhibit coral recovery (Fox et al. 2001).
The islands separation according to geographic location was probably related to
the influence of the monsoon cycle that caused different coverage of foliose and sub-
massive coral in the west and east side of the archipelago (Fig. 3.3). Foliose coral
mostly dominated the west side and sub-massive coral dominated the east side.
Ongkosongo & Sukarno (1986) cited that the wind was strongest from east to south
(i.e. the east monsoon exerts a stronger influence to the island formation), then from
the north and the other wind directions playing a minor role in island formation.
Furthermore the monsoon influence on the currents is clearly marked; the westward
current runs approximately eight months per year and the eastward current flows
around four months per year that has almost twice the strength of westward current.
These physical factors both govern the morphology of the islands and the structure of
the benthic communities that caused mostly of the lateral growth of reef in Kepulauan
Seribu along an east-west axis (Soekarno 1989, Tomascik et al. 1997).
The assumption that the hard coral coverage positively relates to the zoning
management or the expected gradient of blast fishing was wrong. There is also no
correlation between the distance of the island from the mainland Java and the hard
coral cover. However, a new result is that foliose coral characterize the west side
islands (Pandan, Genteng and Melinjo) and sub-massive coral dominate the east side
islands (Opak, Bira and Putri). Bira, Putri and Melinjo (the “northern part”) were
dominated by higher cover of dead coral rubble, while Pandan, Opak and Genteng
(the “southern part”) were characterized by higher coverage of foliose coral,
branching coral, Acropora branching and Acropora digitate.
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4.2. VARIATION IN FISH COMMUNITY ALONG THE GRADIENT OF BLAST FISHING
IMPACT
According to univariate analysis, the structure of the coral reef fish community
of all islands did not reveal a clear pattern according to the expected gradient of blast
fishing impact or the zoning management. Fish diversity did also not correlate with
the expected gradient of blast fishing impact. Furthermore, the percent cover of hard
coral and number of fish species did not correlate with each other. But, multivariate
analysis showed a clear pattern for the relationship between the fish community
structure and the life form categories. The fish community seemed to be separated
according to the composition of benthic groups and life form categories. The PCA
ordination for the islands position, based on benthic groups and life form categories
(Fig. 3.8) and based on the abundance of each fish species (Fig. 3.23) displayed Bira,
Putri and Melinjo in one group of geographic position, the “northern part” of the
studied islands (Fig. 2.1). Pandan, Opak and Genteng displayed in another group as
the “southern part” of the studied islands. Apparently the fish community distribution
is influenced by the composition of benthic and life form categories, because the
“northern part” has mostly a higher cover of dead coral rubble and the “southern part”
mostly foliose, branching and encrusting coral (Fig. 3.8). The most abundant fish in
the “northern part” were Pomacentrus alexanderae and Cirrhilabrus cyanopleura
(Fig. 3.14 - 3.20). Pandan, Opak and Genteng (the “southern part”), seem to be
characterized by the high fish abundance of Pomacentrus lepidogenys. This fish
species was always most abundant in Pandan, Opak and Genteng, but in Bira, Putri
and Melinjo (the “northern part”) it was never among the ten most abundant fish
species (except in Bira in October 2000) (Fig. 3.14 - 3.20).
In all islands a domination of certain fish species was never found throughout
the study period. The evenness index had moderate level for all sites (Table 3.2).
73
Pomacentridae was the most abundant fish family throughout the sampling period and
throughout the islands (Fig. 3.9 & 3.10), followed by Labridae (except in August
2001, when Pomacentridae ranked second most abundant in Melinjo).
The species-abundance relationship of the fish community was best described
and fitted by the log normal model for the pooled data of all islands (Table 3.2, Fig.
3.20). However, the χ2 distribution values were below the conventional 95 %
significance level. When fish data were pooled at each island, the log series model
displayed a better fit in most sampling times (Table 3.2, Fig. 3.14 – 3.19). Geometric
series displayed only one time in Opak in March 2001 (Table 3.2).
Thus, since the fish community throughout all islands performed a better fit
with the log normal, the fish community after around five years of no blast fishing
activities tended to be already in mature level. But some islands, that were still in a
succession process to a mature fish community.
This could be used to explain why the pooled fish data of all islands performed
a log normal distribution model, while mostly the pooled data from each island
performed log series distribution model. For the sum of all islands it is a “complete”
fish community, while for each island it is only part of a “complete” fish community.
Only few fish species had a consistently strong association with certain life form
category, most of the fish species in this study did not show a consistent association.
The CCA-triplot (Fig. 3.25) confirmed that P. lepidogenys is strongly associated with
encrusting coral, except in October 2000, and mostly abundant in Pandan, Genteng
and Opak (the “southern part”). Pomacentrus alexanderae is strongly associated
with mushroom coral and dead coral, mostly found in Bira, Putri and Melinjo (the
“northern part”) (Fig. 3.25). Chaetodon octofasciatus and Chromis analis are mostly
74
found in the islands covered by Acropora-Branching, Acropora-Tabulate, Acropora-
Digitate, sub-massive coral, Heliopora and Millepora.
However, at the family level, Chaetodontidae was consistently associated with
Acropora-Branching, foliose coral, encrusting coral and Heliopora (Fig. 3.27). The
other families had only a weak tendency to associate with a certain life form category,
but not as strong as Chaetodontidae.
Planktivores and omnivores were the two most abundant trophic groups in all
islands (Fig. 3.11 & 3.12). No particular pattern was found along the fishing impact
gradient. The PCA-biplot (Fig. 3.24) also separated the fish trophic groups into
groups of islands. But, the grouping was not as clear as the Fig. 3.8 and Fig. 3.23.
Bira and Melinjo (the “northern part”) were dominated by the high abundance of
benthic feeders and omnivores in all sampling times. Pandan and Genteng (the
“southern part”) were mostly dominated by planktivores (Fig. 3.24). Opak and Putri
were mostly characterized by detritivore, herbivore, piscivore and coralivore fishes.
Furthermore, from CCA-triplot (Fig. 3.28), the herbivores were consistently
associated with areas covered by algae, dead corals and massive coral in Pandan and
Putri. The planktivores associated with branching coral, Acropora-Tabulate and
encrusting coral, mostly located in Opak and Pandan.
The relationship between number of fish species and living coral cover have
been studied for several times by many researchers, but some of these studies resulted
in positive correlation (e.g. Hutomo & Adrim 1986, Hutomo 1987, Gomez et al.
1988) and the others resulted in no correlation (e.g. Luckhurst & Luckhurst 1978a,
McManus et al. 1981). This study did not find a relationship between number of fish
species and percent coverage of hard coral. This confirms previous studies: with no
significant correlations (Luckhurst & Luckhurst 1978a). But these authors found
75
substrate complexity to be the decisive factor for fish species richness and diversity.
In addition, according to Smith (1977), space is the limiting factor for structuring the
fish community, instead of food availability. Many species are considered to have
evolved behavior patterns to ensure an adequate amount of living space (Luckhurst &
Luckhurst 1978b) to be used for feeding, nursery and spawning (Smith 1977). Hence,
the lack of reef complexity in the study area might be the reason for the lack of
relationships between fish diversity and live coral coverage, since most of the hard
corals are new recruits of little complexity (Fig. 3.5). As many researchers have
noted that nowadays coral reef ecosystems have to face natural and anthropogenic
disturbances, which cause the reduction in their topographic complexity and the loss
of habitats (e.g. Carpenter et al. 1981, Sorokin 1995, McManus et al. 1997,
Kunzmann 1997, Edinger et al. 1998, Hodgson 1999, Fox et al. 2001). Sano et al.
(1984) found that the destruction of hermatypic corals leads to changes in fish
community structure because of the change of food resources and the decrease in
structural complexity of coral colonies. The high diversity of reef fish communities
may on the other hand be maintained by unpredictable environmental changes that
prevent development of an equilibrium community (Sale 1977). Therefore most
fishes living in coral reefs have a special form, color and behavior, suitable for a coral
reef biotope (Smith 1977). Their specialization allowed many species to live together
without direct competition for the coral reef’s limited resources (Smith 1977). Bell &
Galzin (1984) noted that the presence and amount of live coral cover may be more
important in structuring fish communities than previously thought.
The impact of blast fishing was still remaining throughout the studied islands,
indicated by the presence of many fields of dead coral (particularly dead coral rubble).
Thus, this case became another reason why there was no clear pattern of the fish
76
community throughout all islands. Riegl & Luke (1998) found significant changes in
coral and fish community composition within dynamited sites. Russ & Alcala (1989)
found that the intense fishing pressure had both direct and indirect effects on the fish
assemblage, and lead to significant changes in the community structure. The study of
Gaudian et al. (1995) also confirmed that coral reef fisheries have a significant impact
on the structure of fish assemblages. In a recent study, Russ & Alcala (1998a)
contradict their previous findings, as they did not find that species richness and the
relative abundance of the families/trophic groups of reef fish in the community were
affected by fishing, and that there was no evidence of phase shifts of the community
in response to fishing. Russ & Alcala (1998b) found that perturbation of the
community by fishing did not alter the relative abundance of major families or trophic
groups of reef fish significantly, except during a period of use of explosives and drive
nets. However, the new fish community would not be the same when certain reef
fishes have been removed from their habitat and community was allowed to-re-
establish naturally thereafter (Smith 1977).
The fish community in the studied islands seemed to be separated according to
the composition of benthic groups and life form categories. This fact confirmed that
the distribution and abundance of species of coral reef fish appears to be strongly
influenced by physical factors (wave exposure, sediment loads, water depth and
topographical complexity) as well as by biological factors (Williams 1982). Galzin
et al. (1994) stated that species diversity of reef fishes within a given family appears
to be affected more by ecological parameters, such as living coral cover, food
diversity, and reproductive behavior, than morphological features. According to
Jennings & Polunin (1997) a single dominant process rarely governs the structure of
reef fish communities.
77
Pomacentridae and Labridae were the most abundant fish families throughout
the islands. This finding coincides with results of Hutomo (1987) who also conducted
a survey in Kepulauan Seribu. Russ & Alcala (1989) found an increasing abundance
of Labridae (Cirrhilabrus and Thalassoma), decreasing abundances of planktivore
Pomacentridae and Caesionidae, and a significant decrease of Chaetodontidae with an
increase in the coverage of coral rubble in fishing grounds that used explosives and
drive-net fishing. According to Russ (1985) and Russ & Alcala (1989), the
abundance (or density) of fishes was a more useful indicator than species richness.
Bouchon-Navaro et al. (1985) found also that the abundance of Chaetodontidae had a
significant positive correlation with coral coverage.
Based on the log normal species-abundance model that fitted for fish
community at all islands together; this indicates that the reef fish communities are
already in a mature stage, consisting of a large heterogeneous assembly of fish species
(May 1975, Ludwig & Reynolds 1988, Magurran 1988). The relative abundance of
fish is most likely a product of many independent factors, which were related to the
function of fish species in diverse ecological roles (May 1975, Ludwig & Reynolds
1988, Magurran 1988). However, mostly the pooled data of fish community from
each island performed log series distribution model, this case indicates a situation
where one or few environment factors determine/regulate the ecology of the
community (Magurran 1988). In this study, the composition of the benthic groups
and life form categories seem to be the determining factors (Fig. 3.3, 3.8 and 3.23).
Furthermore Magurran (1988) noted that the log series model describes a community,
which consists of a small number of abundant species and many species with low
abundance, and this model predicted that species arrive at an unsaturated habitat, at
78
random intervals of time and then occupy the remaining niche (Fig. 3.14 – 3.20,
Appendix 3).
Geometric series displayed only one time in Opak in March 2001 (Table 3.2).
According to May (1975) and Magurran (1988), this model described that the species
arrived at an unsaturated habitat at a regular intervals of time and occupy remaining
fraction of niche. Magurran (1988) noted from field data that geometric series
distribution was found primarily in species poor environment or in the early stages of
a succession - then while a succession proceeds or a condition improve, species
abundance pattern changes into the log series distribution. However, during a
succession of a fish community it is difficult to differentiate between natural or human
disturbances (van Woesik & Done (1997).
Throughout the study period, planktivores and omnivores were the two most
abundant trophic groups in all islands (Fig. 3.11 & 3.12). This finding was not the
same with Sano et al. (1984), who found herbivorous fishes, zooplankton feeders and
omnivores fishes were significantly more abundant and of higher species richness on
the living coral colonies than on damaged coral colonies, and vice versa: when
structural complexity of the coral reef was decreased by bio- and physical-erosion,
diversity and abundance of resident reef fishes decreased. Smith (1977) noted that
food supplies appeared to be quite stable in a coral reef, but did not mean the food
was readily available, and yet the fishes that live there, exhibit a wide variety of
feeding adaptations and specializations of behavior as well as the community
structure. In contrast, Sale (1980) stated that food and space have been considered
most likely to limit the abundance of reef fish. Reef fish were specialized upon
different resources, exhibiting low overlap in the use of food or habitat space (Sale
1977). Munro & Williams (1985) found that the enormous abundance of planktivores
79
fish in Indo-Pacific reefs was related to a higher productivity potential of Indo-
Pacific-reef fisheries.
Fish community is more dependent on benthic groups and life form categories
than the coverage of hard coral. P. lepidogenys is abundant at a substrate mostly
covered by encrusting coral and P. alexanderae is abundant at a substrate with
mushroom and dead coral. C. octofasciatus and C. analis are more abundant in areas
dominated by Acropora corals. In relation with tropic groups, benthic feeders and
omnivores preferred substrates with high cover of dead coral and planktivores
preferred foliose corals. Not a surprise that herbivore is associated with algae and
dead coral with algae locations.
4.3. SEASONAL CHANGES IN FISH COMMUNITY STRUCTURE
In relation with seasonal changes, the multivariate analysis also gave a clear
pattern for the fish community. The differences in fish community structure among
the islands suggest that monsoon cycle and benthic substrate composition were the
major affecting factors (Fig. 3.8 and Fig 3.23). Unfortunately the seasonal changes of
fish species and abundance were not clear enough since data of December 2000 and
January 2001 were missing (Fig. 3.13).
The fish communities tend to be separated clearly into two groups along the
monsoonal season (west monsoon in October 2000 and east monsoon from March to
August 2001) as performed by cluster analysis and NMDS-ordination (Fig. 3.21 &
3.22). The PCA ordination also showed a clear tendency for the grouping of fish
communities by the monsoon cycle (Fig. 3.23), the entire fish community from each
island was displayed in the first quadrant of the PCA-plot as the fish group from the
west monsoon and the others three quadrants displayed the fish communities from the
east monsoon.
80
Fish species richness and the total number of species in the Sanctuary Zone
were also fluctuating seasonally. In support, the comparison of Shannon diversity
index of all islands indicated that the diversity index in October 2001 (west monsoon)
was always significantly lower than in March and April 2001 (the beginning of east
monsoon) (Tables 3.4). Thus, both methods, univariate and multivariate analysis,
revealed a strong tendency that the fish community was different between the two
monsoons.
Weather and currents were two important major factors in determining reef fish
community (Wals 1983). The difficulties to measure the actual impact of destructive
fishing practice is due to the fact that the effect of human activities and natural
processes (wave action, storm, temperature fluctuation, tectonic events, climatic
disruptions, terrestrial runoff, diseases, predator outbreaks) were difficult to separate
(Cesar et al. 1997; Pet-Soede et al. 1999).
The monsoon influences the community structure of fish in the surveyed
islands. There are two different fish communities along the monsoonal cycle.
4.4. VARIATION IN FISH DIVERSITY WITHIN THE ZONING MANAGEMENT
Bira is located in the Sanctuary Zone and was expected to have the highest fish
species richness, but had in fact lower fish species richness compared to the other
islands. Only in March and April 2001, species richness was high in Bira (Table 3.2).
However, when the Shannon diversity index of fish is considered, Bira was generally
similar to other islands, sometimes even lower in diversity index (Table 3.3). Only
once (in April 2001) the fish community in Bira had a significantly higher diversity
index compared to Melinjo (Intensive Utilization Zone).
Also Putri, located within the border of the Sanctuary Zone was also expected to
have higher fish diversity. In fact, it had only a higher diversity during certain periods
81
(Table 3.3). This higher diversity in Putri might be the result of the intensive
surveillance of the surrounding area by the private coast guard of Putri tourist resort,
which reduced the fishing pressure here.
The marine park (especially the Sanctuary Zone) aims at protecting and
maintaining high species richness, are shown e.g. by Samoilys (1988). In addition,
Russ (1985) found that in a protected area the densities of large predatory fishes and
overall abundance and species richness of the reef fish assemblage was significantly
higher compared to non-protected areas. Unfortunately, this expectation is not met in
Kepulauan Seribu Marine Park. Not only the fish abundance, but also the fish
diversity was lower in the Sanctuary Zone compared to other management zones. The
other potential factor lowering the fish diversity might be ongoing illegal fishing in
the Sanctuary Zone with destructive methods. Also Robert (2000) found that most
existing marine reserves are based on social criteria and opportunism rather than
scientific studies. Thus, the zoning management of the national park in Seribu Islands
did not perform succeed in maintaining high diversity of fish in the Sanctuary Zone.
4.5. METHODOLOGICAL ASPECTS 4.5.1. ASSESSMENT OF LIFE FORM CATEGORIES AND BENTHIC GROUPS
The photographic method is usually used for monitoring the biological
condition, growth, mortality and recruitment of corals in a permanent quadrate
(English et al. 1994). However, in this study the photographic method was used for
mapping and assessment of the cover of coral life form categories and benthic groups,
instead of line intercept transect (LIT). The photographic method was used at the
beginning and at the end of the study. It has advantages, but also disadvantages.
Photographic methods need little time in the field for assessment of the substrate
coverage compared to LIT (Line Intercept Transect). It also provides details and
82
allows for a careful observation, a permanent record and a non-destructive sampling
(English et al. 1994). However, it needs a relatively flat area (English et al. 1994),
which sometimes is difficult to find. The permanent transect along 50-m was also
difficult to be maintained during the entire study, therefore only the average percent
cover was used for the further analysis. The photographic method is costly compared
to LIT, because this method needs camera set and negative film, and then requires the
negative film to be scanned into digital picture. Finally, too much time was consumed
to determine the life form categories and measure the cover in the computer. To
analyse one photograph needed around 30-80 minutes, depending on the complexity
of the picture. Thus, all 1,296 photographs were analyzed in 972 hours (with the
average of 45 minute per photograph). Another limitation was that the photograph
resolution was not enough to determine all corals to the genus level, so that just the
life form categories could be determined. The reef rugosity could not be measured
with this method, since it only gives a two-dimensional picture. The photographic
method, however, fulfilled most of the important requirements for substrate mapping
better than LIT.
4.5.2. FISH VISUAL CENSUS
According to Russell et al. (1978), the fundamental problem in quantitative
assessments of fish on coral reef is caused by the sampling. Whereas many fishes are
highly mobile, others are sedentary (Russell et al. 1978). Underwater visual census
(UVC) has errors and biases, caused by the observer, the proper fish behavior, and the
sampling method, most of which result in an underestimation of the population
densities (e.g. Chapman et al. 1974, Brock 1982, Buckley & Hueckel 1989, Greene &
Alevizon 1989, English et al. 1994, Harvey et al. 2002 and Labrosse et al. 2002).
Using UVC Brock (1982) counted only 65 % of the fish species that were collected by
83
rotenone (poison) at the same area, and he only saw 26 % of the cryptic species.
Accordingly, Sale & Sharp (1983) underestimated the density of fish between 11.1 –
26.7 % in a 1-m wide transect.
The ability to spot all fishes present was also depending on the fish behavior and
the divers activity: there are neutral, shy, curious and secretive fishes (Chapman et al.
1974, Kulbicki 1998). Activities and the swimming speed of the observer also
contributed to the bias (Chapman et al. 1974). If the observer moves too slowly, an
overestimation will be the result, and vice versa (Sale & Sharp 1983, Smith 1988).
The air bubbles originating from an open circuit SCUBA also influence the behavior
of the fish (Chapman et al. 1974). While writing data on a slate, the observer might
have overlooked fish when starting again to count (Sale & Sharp 1983). The other
sources of bias were the distance of the diver from the substratum, the diver
experience, and the diver’s physiology in the aquatic environment (Sale & Sharp
1983, Smith 1988, Harvey et al. 2001, Labrosse et al. 2002). The surrounding
environment also gave some limitation for UVC, the visibility of the water, the state
of the ocean and the weather conditions (Labrosse et al. 2002). However, according
to Bell et al. (1985), a trained observer provides consistency in estimating abundance
and length frequency estimations of the same population.
During this study only one observer counted all the fish, in order to minimize
errors and to keep the bias constant (also done by Samoilys & Carlos 2000). The
UVC was done between 10.00 a.m. and 03.00 p.m. to avoid the diurnal-nocturnal
change of fish behavior. During the preliminary study a list of fish species was
developed from all surveyed islands to minimize miss-identification, and to include
also those fish that were caught by net during the study.
84
5. CONCLUSIONS AND OUTLOOK
5.1. CONCLUSIONS
This study showed that after five years of no blast fishing activities, the impact
of these activities on coral reefs are still reflected by the presence of many fields of
dead coral, particularly dead coral rubble. Therefore, the coral reef fish community in
each island is also not yet a mature community stage, but is still in a succession
process.
The fish community is more dependent on benthic groups and life form
categories than on the coverage of hard coral. P. lepidogenys is abundant at a
substrate mostly covered by encrusting coral and P. alexanderae is abundant at a
substrate with mushroom and dead coral. C. octofasciatus and C. analis are more
abundant in area dominated by Acropora corals. Benthic feeders and omnivores
preferred substrate with high cover of dead coral and planktivores preferred foliose
corals. Herbivores are associated with algae and dead coral with algae locations.
The monsoon influences the fish community structure in the surveyed islands.
There are two different fish communities along the monsoonal cycle.
The assumption that the hard coral coverage positively relates to the zoning
management or the expected gradient of blast fishing impact is wrong. There is also
no correlation between the distance of the island from the mainland Java and the hard
coral cover. Thus, the zoning management of the national park is not successful.
85
5.2. OUTLOOK
This study is a step forward to understand the coral reef ecosystem in
Kepulauan Seribu after blast fishing activities. The study has revealed some
unexpected and surprising results. Contrasting results to previous studies on coral
coverage and fish diversity along a distance gradient from the mainland were found.
This study finds a different composition of benthic groups and life form categories
between the east and west side of the archipelago. The fish community is also
different between the monsoon periods.
Regarding the expectations of this study, the succession stage of the fish
community is found; it is still in succession process. The fish biodiversity can be
maintained by intensive surveillance, which performed in Putri. Thus, this
information hopefully can be used to manage the national park.
Considering the current status of the coral reef ecosystem in Kepulauan Seribu,
some questions emerge from the weakness of this work:
1. Is it true that monsoon separates the fish community into two groups? How
does the monsoon influence the fish community? (Since this study had only
one sample of the fish community during west monsoon, this question could
not be answered).
2. Is it necessary to place the sampling sites surrounding each island, in order to
have a better understanding of the changes of the fish community? (This study
only placed the sampling sites at the northeast parts of each island).
3. What should the marine park management do to improve the performance of
the zoning management? Is it necessary to relocate the Sanctuary Zone? Or is
it enough to improve the surveillance and law enforcement?
86
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94
APPENDIX 1. Complete list of the percent cover of the major benthic groups and life form categories (%) at the different study sites.
Study sites Categories P. Pandan
(A) P. Opak Besar
(B) P. KA Bira
(C) P. Putri Timur
(D) P. Melinjo
(E) P. KA Genteng
(F) Hard coral 29.11 18.17 19.62 7.60 25.09 42.75
Acropora Branching 0.21 2.41 2.39 0.32 1.14 9.16 Acropora Digitate 0.04 0.21 0.09 0.09 0.05 0.05 Acropora Tabulate 0.34 3.35 0.46 0.67 1.07 0.66 Coral Branching 6.49 1.71 2.46 0.62 2.05 4.31 Coral Encrusting 2.73 0.78 0.78 0.37 0.58 3.09 Coral Foliose 14.32 0.53 2.66 1.19 13.86 19.91 Coral Massive 2.78 1.77 2.11 0.81 0.45 0.28 Coral Mushroom 0.64 0.42 1.22 0.40 0.68 0.41 Coral Sub-massive 1.39 6.97 5.57 2.11 4.77 2.96 Millepora 0.17 0.03 1.86 1.01 0.24 1.34 Heliopora 0.00 0.00 0.02 0.00 0.20 0.58
Dead Coral 64.27 70.76 70.25 83.43 65.66 51.64
DC (Branching) 6.47 0.00 0.25 0.00 0.00 0.00 DC (Massive) 17.08 19.89 11.02 24.07 18.05 11.67 DC (Rubble) 40.52 30.60 57.87 58.56 47.62 39.15 DC (Tabulate) 0.20 0.00 0.00 0.00 0.00 0.00 DC Algae 0.00 20.27 1.12 0.80 0.00 0.81
Other Fauna 1.52 4.13 6.29 3.42 5.22 4.37
Acanthaster plancii 0.00 0.02 0.00 0.03 0.00 0.00 Sea Anemone 0.03 0.05 0.02 0.03 0.54 0.05 Ascidian 0.09 0.31 0.01 0.44 0.05 0.02 Bryozoan 0.00 0.05 0.00 0.00 0.00 0.00 Lily 0.10 0.15 0.18 0.06 0.21 0.17 Sea Star 0.00 0.03 0.00 0.08 0.01 0.00 Sea Urchin 0.31 0.76 2.58 1.28 1.24 0.12 Soft Coral 0.34 0.50 1.22 0.75 0.95 0.65 Sponge 0.63 1.77 2.27 0.76 2.23 2.97 Tridacna 0.00 0.02 0.00 0.00 0.00 0.00 Zooanthid 0.00 0.41 0.01 0.00 0.00 0.40 Tubipora 0.00 0.07 0.00 0.00 0.00 0.00
Algae 5.10 6.95 3.84 5.55 4.03 1.24
Caulerpa 4.80 6.01 2.78 4.58 2.99 1.13 Halimeda 0.00 0.91 1.01 0.97 0.99 0.01 Macro Algae 0.30 0.03 0.05 0.00 0.05 0.10
95
APP
END
IX 2
. N
umbe
r of c
oral
col
onie
s di
ffer
entia
ted
by th
eir g
row
th fo
rm a
t the
stu
dy s
ites
assu
min
g th
at c
oral
gro
wth
is 2
.4 m
m p
er m
onth
an
d in
circ
ular
dire
ctio
n, S
=sm
all (
< 65
1 cm
2 ; gro
wth
dur
ing
five
year
s), M
=med
ium
(651
- 94
0 cm
2 ; gro
wth
dur
ing
six
year
s) a
nd L
=lar
ge (>
94
0 cm
2 ; gro
wth
dur
ing
seve
n or
mor
e ye
ars)
(van
Moo
rsel
198
8).
Life
form
cat
egor
ies P
. Pan
dan
(A) P
. Opa
k B
esar
(B) P
. KA
Bira
(C) P
. Put
ri Ti
mur
(D) P
. Mel
injo
(E) P
. KA
Gen
teng
(F)
S
M
L S
M
L S
M
L S
M
L S
M
L S
M
L Ac
ropo
ra B
ranc
hing
7
0 1
9 2
4 22
6
6 5
1 1
9 1
4 24
2
20
Acro
pora
Dig
itate
3
0 0
5 1
0 5
1 0
3 1
2 3
0 0
3 0
0 Ac
ropo
ra T
abul
ate
3 1
1 18
2
13
9 0
2 5
1 4
6 2
3 8
2 3
Cor
al B
ranc
hing
72
9
16
76
3 2
137
2 3
21
1 0
83
3 3
118
17
7 C
oral
Enc
rust
ing
250
5 3
21
1 3
53
3 0
42
1 3
39
1 2
28
1 5
Cor
al F
olio
se
201
23
48
11
0 1
81
5 8
35
2 4
348
20
37
180
18
52
Cor
al M
assi
ve
63
6 8
76
3 3
50
2 6
50
0 2
15
1 1
10
0 1
Cor
al M
ushr
oom
95
0
0 51
0
6 97
0
0 40
0
3 81
0
0 43
0
0 C
oral
Sub
-mas
sive
50
3
3 24
2 12
20
19
8 11
13
10
0 3
7 20
9 7
8 97
4
9 M
illep
ora
7 2
0 1
0 0
45
4 5
52
2 3
8 1
1 18
1
2 H
elio
pora
0
0 0
0 0
0 0
0 0
1 0
10
7 0
1 17
1
1
96
APP
END
IX 3
. C
ompl
ete
list o
f fis
h sp
ecie
s acc
ordi
ng to
thei
r sys
tem
atic
ord
er a
nd th
eir a
bund
ance
s at e
ach
site
thro
ugho
ut th
e st
udy
perio
d.
P.
Pan
dan
(A)
P. O
pak
Bes
ar (B
) P.
KA
Bira
(C)
P. P
utri
Tim
ur (D
) P.
Mel
injo
(E)
P. K
A G
ente
ng (F
) N
o. S
peci
es
Oct
‘00
Mar
‘00
Apr
‘00
Jun
‘00
Aug
‘00
Oct
‘00
Mar
‘00
Apr
‘00
Jun
‘00
Aug
‘00
Oct
‘00
Mar
‘00
Apr
‘00
Jun
‘00
Aug
‘00
Oct
‘00
Mar
‘00
Apr
‘00
Jun
‘00
Aug
‘00
Oct
‘00
Mar
‘00
Apr
‘00
Jun
‘00
Aug
‘00
Oct
‘00
Mar
‘00
Apr
‘00
Jun
‘00
Aug
‘00
Mur
aeni
dae
1 G
ymno
thor
ax s
p.
1
Hol
ocen
trid
ae
2 M
yrip
ristis
adu
sta
1
7
7
1
3 M
yrip
ristis
vio
lace
a
2
4
Myr
ipris
tis s
p.
1
4
5 Sa
rgoc
entro
n pr
aslin
1
2
3
15
Syn
odon
tidae
6
Syno
dus
sp.
1
3 2
1
1
3 2
2
2
1
1
1
Aul
osto
mid
ae
7 Au
lost
omus
chi
nens
is
2
1
Fis
tula
riida
e
8
Fist
ular
ia c
omm
erso
nii
2
1
4
Tet
raro
gida
e
9
Abla
bys
taen
iano
tus
1 3
4 2
4
1
3
1 2
1
2
2 3
2
1
2
3
1
2
Sco
rpae
nida
e
10
Pte
rois
vol
itans
1
1
S
erra
nida
e
11
Cep
halo
phol
is a
rgus
6
1 1
2
3
4 2
1
7
12
Cep
halo
phol
is b
oena
k
1 3
1
2 1
1
1
1
1 6
2 2
13
Cep
halo
phol
is s
p. 1
4 1
6 2
1
1 2
3
1 1
1 1
1
2 2
1 14
Cep
halo
phol
is s
p. 2
2
2
1 2
15
Epi
neph
elus
sp.
1
1
2
2
2
1
16 E
pine
phel
us s
p. 2
1
1
1
17
Epi
neph
elus
sp.
3
1
Apo
goni
dae
18 A
pogo
n co
mpr
essu
s 2
39
5 3
1 7
7 25
23
13
7
5 2
29
2 5
1 2
5 6
29
1 22
8
2 57
19
Che
ilodi
pter
us m
acro
don
1
1
3
20 C
heilo
dipt
erus
qui
nque
linea
tus
1
20
2
32
5
15
1
2 21
Sph
aera
mia
nem
atop
tera
3
15
16
L
utja
nida
e
22
Lut
janu
s bi
gutta
tus
1
1
2 2
23 L
utja
nus
decu
ssat
us
1
1 10
1 3
1
1
1
1
2 1
3 1
24
Lut
janu
s fu
lvifl
amm
us
1
2
1
Hae
mul
idae
25
Ple
ctor
hinc
hus
chae
todo
noid
es
1
1
1
Nem
ipte
ridae
26
Pen
tapo
dus
trivi
ttatu
s
1 25
29
14
12
37
43
20
25
6
17
9 13
7
3 3
8 17
7
7 22
4
20
19
6 4
1 10
1
27 S
colo
psis
bilin
eata
4
1 2
13
4 4
4 12
2
9 4
5 7
10
1 7
4 11
13
16
9
6 5
3 1
2 2
5 1
5 28
Sco
lops
is li
neat
us
5
2
3
11
32
2
1
7
13
7 22
4
1
29 S
colo
psis
mar
garit
ifer
2
7 14
1
2
3
1 7
7 4
7
1 4
1
2 3
1 2
1 2
15
10
97
APP
END
IX 3
. C
ontin
ued
P.
Pan
dan
(A)
P. O
pak
Bes
ar (B
) P.
KA
Bira
(C)
P. P
utri
Tim
ur (D
) P.
Mel
injo
(E)
P. K
A G
ente
ng (F
) N
o. S
peci
es
Oct
‘00
Mar
‘00
Apr
‘00
Jun
‘00
Aug
‘00
Oct
‘00
Mar
‘00
Apr
‘00
Jun
‘00
Aug
‘00
Oct
‘00
Mar
‘00
Apr
‘00
Jun
‘00
Aug
‘00
Oct
‘00
Mar
‘00
Apr
‘00
Jun
‘00
Aug
‘00
Oct
‘00
Mar
‘00
Apr
‘00
Jun
‘00
Aug
‘00
Oct
‘00
Mar
‘00
Apr
‘00
Jun
‘00
Aug
‘00
Mul
lidae
30
Par
upen
eus
barb
erin
us
1
3
6 1
5
1 2
7 1
6
2 2
Eph
ippi
dae
31 P
lata
x sp
.
1
1
1
C
haet
odon
tidae
32
Cha
etod
on a
urig
a 1
33
Cha
etod
on o
ctof
asci
atus
26
25
49
56
34
37
60
79
38
49
35
36
31
29
20
15
22
15
9
6 28
25
31
33
28
29
36
81
75
57
34
Cha
etod
on v
agab
undu
s 1
35
Che
lmon
rost
ratu
s
2
36
Hen
ioch
us s
p.
1
1
1
1
Pom
acan
thid
ae
37 C
entro
pyge
bic
olor
2
38
Cha
etod
onto
plus
mes
oleu
cus
3 14
4
16
5 2
1
1 3
2 2
6 9
4 2
6 8
2 2
2
3 4
4 3
2 13
5
6 39
Pom
acan
thus
sp.
2
P
omac
entr
idae
40
Abu
defd
uf v
aigi
ensi
s
1 21
3 21
11
2
1 7
4
4
11
15
7 15
14
11
9 15
5 6
10
0 5
29
8
41 A
bude
fduf
ben
gale
nsis
1
42
Abu
defd
uf s
exfa
scia
tus
4
4 5
4
12
10
1
12
1
14
4 16
7 36
18
10
1 3
5 12
7 55
10
9 10
0
43 A
mbl
ygly
phid
odon
cur
acao
33
47
39
52
51
69
80
63
80
66
13
3 14
6 14
6 20
6 75
45
51
37
41
44
39
38
36
41
50
67
12
5 20
7 11
7 44
44
Am
blyg
lyph
idod
on le
ucog
aste
r
11
14
8 6
15
17
34
13
79
45
81
36
3 28
55
20
7
3
7 25
33
58
13
16
14
45 A
mbl
ygly
phid
odon
tern
aten
sis
5
31
19
1 5
2
25
1 10
1
3 1
3
1
2 21
30
9
46 A
mph
iprio
n fre
natu
s
2
2
47
Am
phip
rion
ocel
laris
3 3
6
2
2
6
48
Am
phip
rion
perc
ula
1
1
49 A
mph
iprio
n sa
ndar
acin
os
1 11
6
6 7
2 50
Am
phip
rion
sp.
1
36
16
51 C
heilo
prio
n la
biat
us
6
6
1
16
4 12
3
2 14
16
22
1 4
10
15
8
1
52
Chr
omis
ana
lis
28
27
47
68
62
57
89
57
81
84
46
39
35
58
41
32
46
36
32
33
97
56
80
73
36
46
57
93
92
87
53 C
hrom
is a
tripe
ctor
alis
20
0 25
5
3 14
20
0 27
7
7 6
104
3 23
2 10
0 5
13
11
11
1
39
100
5 77
8 54
Chr
omis
flav
ipec
tora
lis
1
55 C
hrom
is v
iridi
s
2
100
56 C
hrom
is w
eber
i 10
0
3 8
34
1 13
1
57 C
hrom
is x
anth
ura
2
1
1
58 C
hrom
is s
p.
23
13
3
2
9
6
59 C
hrys
ipte
ra ro
lland
i
1
1
4 1
1
17
1
60 C
hrys
ipte
ra s
p.
4
61 D
ascy
llus
arua
nus
1
2
62 D
ascy
llus
trim
acul
atus
2 9
1
4
1
2 7
4 9
4 6
63
Dis
chis
todu
s m
elan
otus
4
64
Dis
chis
todu
s pr
osop
otae
nia
5
3
1
1
9 6
7 4
65 N
eogl
yphi
dodo
n bo
nang
1
3 3
1
66
Neo
glyp
hido
don
mel
as
2 1
2
1
1 5
24
1 2
6 3
13
1
7 3
1 1
10
3
3 2
6 67
Neo
glyp
hido
don
nigr
oris
2
22
24
45
30
25
59
32
33
10
50
46
50
68
13
41
53
49
64
26
59
59
43
65
21
43
67
59
51
68
Neo
pgly
phid
odon
oxy
odon
18
12
69
Neo
pom
acen
trus
anab
atoi
des
100
3
10
0 10
0
1
100
100
20
100
100
98
APP
END
IX 3
. C
ontin
ued
P.
Pan
dan
(A)
P. O
pak
Bes
ar (B
) P.
KA
Bira
(C)
P. P
utri
Tim
ur (D
) P.
Mel
injo
(E)
P. K
A G
ente
ng (F
) N
o. S
peci
es
Oct
‘00
Mar
‘00
Apr
‘00
Jun
‘00
Aug
‘00
Oct
‘00
Mar
‘00
Apr
‘00
Jun
‘00
Aug
‘00
Oct
‘00
Mar
‘00
Apr
‘00
Jun
‘00
Aug
‘00
Oct
‘00
Mar
‘00
Apr
‘00
Jun
‘00
Aug
‘00
Oct
‘00
Mar
‘00
Apr
‘00
Jun
‘00
Aug
‘00
Oct
‘00
Mar
‘00
Apr
‘00
Jun
‘00
Aug
‘00
70 N
eopo
mac
entru
s az
ysro
n
31
7
32
20
3
71 P
lect
rogl
yphi
dodo
n la
crym
atus
33
44
43
47
45
6
5
2 4
1 1
1 1
1 14
16
22
42
28
1
2
1
72
Pom
acen
trus
alex
ande
rae
16
1 17
0 18
2 20
7
123
7 57
19
44
24
3 16
1 30
0 17
7 23
64
12
0 47
70
13
1 22
4 30
0 22
8 10
4 25
82
22
11
0 61
73
Pom
acen
trus
ambo
inen
sis
9
25
14
11
7
99
18
10
2
1 4
2
6 31
18
24
17
23
23
9
14
12
11
8 74
Pom
acen
trus
gram
mor
hync
us
8 32
32
37
41
1
56
71
48
84
44
27
63
43
42
1 1
15
13
20
19
21
8
22
47
37
7 75
Pom
acen
trus
lepi
doge
nys
30
0 30
0 30
0 30
0 20
1 15
0 20
0 14
5 18
5 12
0 38
46
56
8
18
6 20
0 20
1 20
0 20
0 20
9 76
Pom
acen
trus
philip
pinu
s
11
2
1
9
2
77 P
omac
entru
s sp
. 1
15
33
41
53
5 31
10
18
2 13
2
11
8 21
29
44
64
11
51
30
39
3 10
8
2 17
78
Pom
acen
trus
sp. 2
11
5
79 P
omac
entru
s sp
. 3
1
1
1
80 P
omac
entru
s ta
enio
met
opon
1 6
1 4
1
20
17
2
81
Ste
gast
es fa
scio
latu
s
1
1
L
abrid
ae
82 A
nam
pses
sp.
2
2
1
1
2
3 2
1
3 5
1
1
1 83
Che
ilinus
chl
orou
rus
2
2
2
29
10
9 12
2 4
4 3
6
6 13
9
4
7 7
7
3 6
1 10
84
Che
ilinus
fasc
iatu
s 4
10
13
11
11
17
41
46
15
16
9 19
15
11
8
37
29
49
38
31
18
17
12
21
4 17
20
37
17
13
85
Che
ilinus
und
ulat
us
1 4
16
11
11
4
17
1 2
5
6 6
2
15
1
1
2 4
3
2 12
17
8
4 86
Cho
erod
on a
ncho
rago
2
8
2
2
1
1
1
2
2
1
1 8
1
87
Cirr
hila
brus
cya
nopl
eura
2
49
200
300
300
100
116
2 30
0 15
3
30
150
216
33
100
209
200
200
200
30
0 11
1 21
6 30
0
9 40
11
0 10
1 88
Dip
roct
acan
thus
xan
thur
us
1 4
5 12
10
8 2
4 12
7
8 3
16
11
2 1
10
5 3
2 2
2 9
11
13
5 14
27
23
89
Epi
bulu
s in
sidi
ator
2 6
7 5
5 2
4 3
2 1
2 2
2 2
3 2
6
7 1
2 90
Gom
phos
us v
ariu
s 3
1 2
3
2
3 2
1
3
91
Hal
icho
eres
arg
us
5
1 10
2 59
202
8 2
2
100
6 1
100
2
100
92 H
alic
hoer
es c
hlor
opte
rus
3
5
3 1
26
14
3
12
6 7
5 8
2 9
9 4
5 1
2
3
9 1
93
Hal
icho
eres
hor
tula
nus
2 5
7 7
4
21
10
13
14
12
13
21
17
18
14
16
17
9 10
23
13
2
3 5
3 15
16
94
Hal
icho
eres
mel
anur
us
1 27
23
38
29
1
20
37
12
26
19
31
27
16
12
16
11
10
117
36
29
22
43
24
39
20
21
95
Hal
icho
eres
pur
pure
scen
s 2
6 10
20
5
15
24
12
8
23
21
36
18
4
4 8
6 6
11
9 8
15
20
2 15
29
16
12
96
Hal
icho
eres
vro
likii
7 12
2
1
1
2 1
3 1
1
1
2 5
1
1 5
4 97
Hem
igym
nus
mel
apte
rus
1 7
8 4
4
2 12
1
4
6 4
3 1
2
13
10
3 10
2
4 1
1 4
4
7 1
98 L
abro
ides
dim
idia
tus
5 8
14
16
2 6
5 9
7 3
13
11
15
11
11
30
13
13
17
8 11
11
6
20
7 5
9 6
5 8
99 M
acro
phar
yngo
don
orna
tus
5
1
5
21
1
100
100
Pter
agog
us s
p.
4
101
Stet
hoju
lis s
trigi
vent
er
3
18
2
1
7 4
1
11
1
102
Thal
asso
ma
hard
wic
ke
1 1
1 2
7
2
2
1
1
4
5 2
1
103
Thal
asso
ma
luna
re
40
43
34
55
28
50
49
49
19
26
33
14
45
30
34
51
22
27
40
31
73
39
38
48
70
34
83
88
76
32
104
Thal
asso
ma
lute
scen
s
4
10
5 Th
alas
som
a pu
rpur
eum
1
S
carid
ae
106
Scar
us g
hobb
an
7
2
2
107
Scar
us n
iger
1
1
4
3 1
1
4
2 3
10
8 C
hlor
urus
sor
didu
s
4 16
2
2 15
30
10
1
2
1
12
1
109
Scar
us v
iridi
fuca
tus
8
1 31
5
1
3
4 7
4
1
1
11
0 Sc
arus
sp.
1
16
10
5 11
15
0 14
10
0 46
100
1 30
1
100
153
7
100
101
59
18
16
100
3
1
2
11
1 Sc
arus
sp.
2
1
2
6
99
APP
END
IX 3
. C
ontin
ued
P.
Pan
dan
(A)
P. O
pak
Bes
ar (B
) P.
KA
Bira
(C)
P. P
utri
Tim
ur (D
) P.
Mel
injo
(E)
P. K
A G
ente
ng (F
) N
o. S
peci
es
Oct
‘00
Mar
‘00
Apr
‘00
Jun
‘00
Aug
‘00
Oct
‘00
Mar
‘00
Apr
‘00
Jun
‘00
Aug
‘00
Oct
‘00
Mar
‘00
Apr
‘00
Jun
‘00
Aug
‘00
Oct
‘00
Mar
‘00
Apr
‘00
Jun
‘00
Aug
‘00
Oct
‘00
Mar
‘00
Apr
‘00
Jun
‘00
Aug
‘00
Oct
‘00
Mar
‘00
Apr
‘00
Jun
‘00
Aug
‘00
Ble
nniid
ae
112
Mei
acan
thus
sm
ithi
1
1
1
4 4
1
1
1
Mic
rode
smid
ae
113
Pter
eleo
tris
evid
es
2
12
3 3
2
2 A
cant
hurid
ae
114
Acan
thur
us li
neat
us
1
Sig
anid
ae
115
Siga
nus
cana
licul
atus
4 2
4
2
11
6 Si
ganu
s co
rallin
us
1
1
1
1
1
1
2
117
Siga
nus
vulp
inus
2
3
3
1 1
1
4
O
stra
ciid
ae
118
Ost
raci
on c
ubic
us
2
1
1
1
Tet
raod
ontid
ae
119
Arot
hron
sp.
1
1
100
APPENDIX 4. Trophic group of all fish species observed (Sources: Lieske & Myers 1997; Fish Base www.fishbase.org).
Species Trophic group Species Trophic group Muraenidae Chaetodontidae
1 Gymnothorax sp. Piscivore 32 Chaetodon auriga Benthic feeder Holocentridae 33 Chaetodon octofasciatus Omnivore
2 Myripristis adusta Planktivore 34 Chaetodon vagabundus Omnivore 3 Myripristis violacea Benthic feeder 35 Chelmon rostratus Benthic feeder 4 Myripristis sp. Planktivore 36 Heniochus sp. Benthic feeder 5 Sargocentron praslin Benthic feeder Pomacanthidae
Synodontidae 37 Centropyge bicolor Omnivore 6 Synodus sp. Piscivore 38 Chaetodontoplus mesoleucus Omnivore
Aulostomidae 39 Pomacanthus sp. Omnivore 7 Aulostomus chinensis Piscivore Pomacentridae Fistulariidae 40 Abudefduf vaigiensis Omnivore 8 Fistularia commersonii Piscivore 41 Abudefduf bengalensis Omnivore Tetrarogidae 42 Abudefduf sexfasciatus Omnivore 9 Ablabys taenianotus Piscivore 43 Amblyglyphidodon curacao Omnivore Scorpaenidae 44 Amblyglyphidodon leucogaster Benthic feeder
10 Pterois volitans Piscivore 45 Amblyglyphidodon ternatensis Omnivore Serranidae 46 Amphiprion frenatus Omnivore
11 Cephalopholis argus Piscivore 47 Amphiprion ocellaris Omnivore 12 Cephalopholis boenak Piscivore 48 Amphiprion percula Planktivore 13 Cephalopholis sp. 1 Piscivore 49 Amphiprion sandaracinos Omnivore 14 Cephalopholis sp. 2 Piscivore 50 Amphiprion sp. Planktivore 15 Epinephelus sp. 1 Piscivore 51 Cheiloprion labiatus Omnivore 16 Epinephelus sp. 2 Piscivore 52 Chromis analis Planktivore 17 Epinephelus sp. 3 Piscivore 53 Chromis atripectoralis Planktivore
Apogonidae 54 Chromis flavipectoralis Planktivore 18 Apogon compressus Benthic feeder 55 Chromis viridis Omnivore 19 Cheilodipterus macrodon Piscivore 56 Chromis weberi Planktivore 20 Cheilodipterus quinquelineatus Benthic feeder 57 Chromis xanthura Planktivore 21 Sphaeramia nematoptera Benthic feeder 58 Chromis sp. Planktivore
Lutjanidae 59 Chrysiptera rollandi Omnivore 22 Lutjanus biguttatus Piscivore 60 Chrysiptera sp. Planktivore 23 Lutjanus decussatus Piscivore 61 Dascyllus aruanus Omnivore 24 Lutjanus fulviflammus Benthic feeder 62 Dascyllus trimaculatus Omnivore
Haemulidae 63 Dischistodus melanotus Herbivore
25 Plectorhinchus chaetodonoides Benthic feeder 64 Dischistodus prosopotaenia Herbivore Nemipteridae 65 Neoglyphidodon bonang Omnivore
26 Pentapodus trivittatus Benthic feeder 66 Neoglyphidodon melas Omnivore 27 Scolopsis bilineata Benthic feeder 67 Neoglyphidodon nigroris Omnivore 28 Scolopsis lineatus Benthic feeder 68 Neopglyphidodon oxyodon Omnivore 29 Scolopsis margaritifer Benthic feeder 69 Neopomacentrus anabatoides Planktivore
Mullidae 70 Neopomacentrus azysron Planktivore 30 Parupeneus barberinus Benthic feeder 71 Plectroglyphidodon lacrymatus Herbivore
Ephippidae 72 Pomacentrus alexanderae Omnivore
31 Platax sp. Omnivore 73 Pomacentrus amboinensis Herbivore
101
APPENDIX 4. Continued.
Species Trophic group Species Trophic group 74 Pomacentrus grammorhyncus Herbivore 100 Pteragogus sp. Benthic feeder 75 Pomacentrus lepidogenys Planktivore 101 Stethojulis strigiventer Benthic feeder 76 Pomacentrus philippinus Herbivore 102 Thalassoma hardwicke Benthic feeder 77 Pomacentrus sp. 1 Omnivore 103 Thalassoma lunare Benthic feeder 78 Pomacentrus sp. 2 Omnivore 104 Thalassoma lutescens Benthic feeder 79 Pomacentrus sp. 3 Omnivore 105 Thalassoma purpureum Benthic feeder 80 Pomacentrus taeniometopon Herbivore Scaridae
81 Stegastes fasciolatus Herbivore 106 Scarus ghobban Herbivore
Labridae 107 Scarus niger Herbivore
82 Anampses sp. Benthic feeder 108 Chlorurus sordidus Detritivore 83 Cheilinus chlorourus Benthic feeder 109 Scarus viridifucatus Herbivore 84 Cheilinus fasciatus Benthic feeder 110 Scarus sp. 1 Herbivore 85 Cheilinus undulatus Benthic feeder 111 Scarus sp. 2 Herbivore 86 Choerodon anchorago Benthic feeder Blenniidae
87 Cirrhilabrus cyanopleura Planktivore 112 Meiacanthus smithi Omnivore 88 Diproctacanthus xanthurus Coralivore Microdesmidae
89 Epibulus insidiator Benthic feeder 113 Ptereleotris evides Planktivore 90 Gomphosus varius Benthic feeder Acanthuridae 91 Halichoeres argus Benthic feeder 114 Acanthurus lineatus Herbivore 92 Halichoeres chloropterus Benthic feeder Siganidae 93 Halichoeres hortulanus Benthic feeder 115 Siganus canaliculatus Herbivore 94 Halichoeres melanurus Benthic feeder 116 Siganus corallinus Herbivore 95 Halichoeres purpurascens Benthic feeder 117 Siganus vulpinus Herbivore 96 Halichoeres vrolikii Benthic feeder Ostraciidae 97 Hemigymnus melapterus Benthic feeder 118 Ostracion cubicus Omnivore 98 Labroides dimidiatus Benthic feeder Tetraodontidae
99 Macropharyngodon ornatus Benthic feeder 119 Arothron sp. Omnivore