elib.suub.uni-bremen.deelib.suub.uni-bremen.de/diss/docs/E-Diss534_Aktani.pdf · 2004-07-21 · Erratum Erratum to: “AKTANI, U. 2003. Fish communities as related to substrate characteristics

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.

iii

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

iv

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.

v

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

vi

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.

vii

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.

viii

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

xi

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

xii

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. ..............


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. ..............


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. .....


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. .....


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

xiv

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


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


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


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


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


Abu

ndan

ce (%

)

October 2000

��

��

��

��

��

��

��

��

��

0

10

20

30

40

50

60

70

80

90


Abu

ndan

ce (%

)

March 2001

��

��

��

��

��

��

��

0

10

20

30

40

50

60

70

80

90


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


Abu

ndan

ce (%

)

June 2001

��

��

��

��

��

��

��

��

��

��

��

��

��

�� 0

10

20

30

40

50

60

70

80

90


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


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


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


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


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


Indi

vidu

al n

umbe

r

Planktivore��

Omnivore Benthic feeder��

Herbivore��

Detritivore��

Corallivore��

Piscivore

August 2001


��

��

��

��

��

��

��

��

��

��

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


(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 '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

)


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 '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

)


P. KA BIRA (C)


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 '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

)


P. PUTRI TIMUR (D)


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 '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

)


P. MELINJO (E)


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 '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

)


P. KA GENTENG (F)


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 '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

)


ALL ISLANDS


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.

72

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

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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.

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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

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96

APP

END

IX 3

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97

APP

END

IX 3

. C

ontin

ued

P.

Pan

dan

(A)

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pak

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ar (B

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KA

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‘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

Documents

elib.suub.uni-bremen.deelib.suub.uni-bremen.de/diss/docs/E-Diss534_Aktani.pdf · 2004-07-21 · Erratum Erratum to: “AKTANI, U. 2003. Fish communities as related to substrate characteristics