3phil.reef fish-go.pmd JA Anticamara and others 1 SCIENCE DILIMAN (JANUARY-JUNE) 27:1, 1-47 National Patterns of Phil ippine Reef Fish Diversity and Its Impl ications on the Current Municipal-Level Management Jonathan A. Anticamara University of the Philippines Diliman Kevin Thomas B. Go* University of the Philippines Diliman Stevenson S. Ongsyping University of the Philippines Diliman Francesco Antonio T. Valdecañas University of the Philippines Diliman Ryan Gabriel S. Madrid University of the Philippines Diliman _______________ *Corresponding Author ISSN 0115-7809 Print / ISSN 2012-0818 Online ABSTRACT Recent national-level assessments of Philippine reef f ish diversity have b e e n m a i n l y b a s e d o n s p e c i e s r i c h n e s s s u r v e y s , b u t g e n e r a l l y d o n o t a c c o u n t f o r r e e f f i s h a b u n d a n ce a n d b i o m a s s — m e t r i c s t h a t b e t t e r d e s c r i b e f i s h co m m u n i t y a s s e m b l a g e s . G i v e n t h a t t h e P h i l i p p i n e s i s considered a major biodiversity hotspot and is heavily reliant on coastal r e s o u r c e s , t h e r e i s a g r e a t n e e d t o q u a n t i f y t h e c u r r e n t s t a t u s o f i t s r e e f f i s h d i v e r s i t y u s i n g s t a n d a r d i ze d m e t h o d s . H e r e , s t a n d a r d i z e d Underwater Visual Census (UVC) belt transect sampling methods were u s ed to q u a n t i f y c u r r e n t l eve l s of r ee f f i s h s p ec i e s r i c h n e s s , r e l a t i ve a b u n d a n ce, a n d r e l a t i ve b i o m a s s t h r o u g h o u t t h e P h i l i p p i n e s . Re s u l t s showed that most surveyed municipalities were still species-rich (22.2 ± 0.8 reef f ish species per 100 m 2), but appeared depleted in terms of r e e f f i s h a b u n d a n c e a n d b i o m a s s . Pa r t i t i o n i n g a n a l y s i s r e v e a l e d s i g n i f i c a n t d i f f e r e n c e s i n r ee f f i s h s p e c i e s r i c h n e s s p a t t e r n s a c r o s s municipalities, suggesting the presence of a few restricted-range and r a r e s p e c i e s p e r s i te . H o w ev e r, p a r t i t i o n i n g a n a l y s i s a cco u n t i n g f o r National Patterns of Philippine Reef Fish Diversity 2 r e l a t i v e a b u n d a n c e s h o w e d t h a t r e e f f i s h d i v e r s i t y w a s g e n e r a l l y h o m o g e n o u s a c r o s s s t u d y s i t e s , s u g g e s t i n g t h e d o m i n a n c e o f a f e w highly-abundant species. SIMPER analysis revealed that Philippine reefs were generally dominated by small and medium-bodied species, rather than large-bodied species—the latter of which are especially vulnerable to f i s h i n g d u e to ce r t a i n l i fe h i s to r y t r a i t s (e.g. , l a te a g e a t m a t u r i t y a n d s l o w g r o w t h r a t e ) a n d c o m m e r c i a l e x p l o i t a t i o n . W h i l e c u r r e n t municipal-level management may be suff icient for restricted-range f ish species, large-scale conservation efforts (i.e. , in the form of collaborative marine reserve networks) are needed for wide-range and l a r g e - b o d i e d s p e c i e s t h a t a r e n o t c o n f i n e d t o p o l i t i c a l l y - d e f i n e d m u n i c i p a l b o u n d a r i e s . I n a d d i t i o n , l o n g - t e r m a n d n a t i o n w i d e e f f o r t s t o systematically monitor Philippine reef diversity are needed to provide up-to-date knowledge of the status of Philippine reef diversity that will h e l p s u p p o r t s c i e n c e - b a s e d r e e f m a n a g e m e n t a n d r e c o v e r y e f f o r t s throughout the country. Keywords: Conservation, coastal management, marine reserves, Philippine r e e f s , r ee f f i s h e r i e s LAYMAN’S ABSTRACT Recent national-level assessments of Philippine reef f ish diversity are mainly based on the number of species present, but generally do not account for the abundance and biomass of these species—metrics that b e t t e r d e s c r i b e t h e f i s h co m m u n i t y co m p o s i t i o n . U n d e r s t a n d i n g t h e current status of reef f ish in the Philippines is impor tant , considering that the country is a marine biodiversity hotspot, and is greatly reliant on marine resources for food and livelihood. To address this, we conducted underwater reef f ish surveys throughout t h e c o u n t r y, r e c o r d i n g f i s h a b u n d a n ce a n d s i z e u s i n g s t a n d a r d i ze d methods. We found that most surveyed municipalities still held a high n u m b e r o f s p e c i e s , b u t a p p e a r e d d e p l e t e d i n t e r m s o f r e e f f i s h a b u n d a n c e a n d b i o m a s s . F u r t h e r a n a l y s i s s u g g e s t e d t h a t m o s t m u n i c i p a l i t i e s w e r e h o m e t o s o m e r e s t r i c t e d - r a n g e a n d r a r e s p e c i e s , b u t w e r e d o m i n a t e d b y a f e w h i g h l y - a b u n d a n t s p e c i e s . F u r t h e r m o r e , Philippine reefs were generally dominated by small- and medium-bodied s p e c i e s , r a t h e r t h a n l a r g e - b o d i e d s p e c i e s . L a r g e - b o d i e d f i s h a r e e s p e c i a l l y v u l n e r a b l e to f i s h i n g d u e to t h e i r h i g h co m m e r c i a l v a l u e , w h i c h m a k e s t h e m d e s i r a b l e f i s h e r i e s t a r g e t s . I n a d d i t i o n , ce r t a i n JA Anticamara and others 3 characteristics, such as slow growth and low reproductive rate, also add to the vulnerability of large-bodied species. A l t h o u g h c u r r e n t m u n i c i p a l - l eve l m a n a g e m e n t m a y b e s u f f i c i e n t f o r r e s t r i c t e d - r a n g e f i s h s p e c i e s , l a r g e - s c a l e c o n s e r v a t i o n e f f o r t s a r e n e e d e d f o r w i d e - r a n g e a n d l a r g e - b o d i e d s p e c i e s t h a t t r a v e r s e l a r g e areas, often across politically-def ined municipal boundaries. An example o f l a r g e - s c a l e c o n s e r v a t i o n e f f o r t s w o u l d b e c r e a t i n g n e t w o r k s o f “marine reserves,” which are areas of protected sea where f ishing and other exploitative activities are not allowed. In addition, long-term and nationwide efforts to systematically monitor Philippine reef diversity are needed to provide up-to-date knowledge on the status of Philippine reef diversity. This knowledge is key in helping suppor t science-based reef management and recovery effor ts throughout the country. INTRODUCTION The Philippines, an archipelagic nation located in the Coral Triangle of the Indo- Pacif ic region, is considered to be one of the global centres of marine f ish diversity (Carpenter and Springer 2005). The country is also a known biodiversity hotspot— a place that is rapidly losing biodiversity in a short amount of time due to extensive habitat destruction and exploitation (Licuanan and Gomez 2000; Roberts and others 2002; Possingham and Wilson 2005; Allen 2008). Over the past two decades, pressure from overexploitation and destructive human activities have contributed to the degradation of Philippine reefs and the deterioration of coastal resources (Gomez and others 1994; Gomez 1997; White and Vogt 2000; Nanola and others 2006; Briones 2007). Declines in coastal resource production, particularly in the f isheries sector, is a matter of concern for the Filipino people, since many Filipinos rely heavily on f isheries products for both food and livelihood (Gjertsen 2005; BFAR 2012). Therefore, it is important to ask what is left of reef f ish diversity in the Philippines, considering the country’s growing population and the potential increase in demands for f isheries-related products that may follow. Most recent national or regional analyses of reef f ish diversity in the Philippines have been based on species presence or absence data obtained through a variety of sources, including Underwater Visual Census (UVC) assessments, museum collections, published literature, and expert opinion (Carpenter and Springer 2005; Allen 2008; Nañola Jr. and others 2011). These studies used presence-absence data from various sources to create species distribution maps that allow assessments of biodiversity patterns over large areas, and pinpoint hotspots of conservation National Patterns of Philippine Reef Fish Diversity 4 importance. Other assessments have used vulnerability scores (often put together by experts based on information on species population trends, distribution, life history, ecology, threats, and existing conservation measures) (Comeros-Raynal and others 2011) or local ecological knowledge (Lavides and others 2009) to map areas at risk of potential species loss. While useful, these assessments do not account for f ish abundance and biomass—metrics that are needed in estimating potential f isheries production or yield, the effectiveness of management schemes (e.g. , Marine Reserve [MR] enforcement), and describing f ish community assemblages in greater detail than species presence-absence data. To date, only a few studies have presented estimates of reef f ish diversity in the Philippines that account for not only species richness, but also species relative abundance and biomass (Go and others in press; Nanola and others 2006). However, many of these publications, generally included only a few sites in the country (Allen 2002; Stockwell and others 2009; Anticamara and others 2010), were limited to a few commercial species (Alcala 1988; Russ and Alcala 2004; Russ and others 2005), or were mainly focused on studying the effectiveness of select no-take Marine Reserves (MRs). Thus, there is still a great need to quantify the current status of reef f ish diversity in representative sites throughout the Philippines by gathering reef f ish diversity data that not only reflects estimates of species richness, but also show the relative abundance and biomass of reef f ish species. Collecting such data is vital in providing up-to-date knowledge for science-based decision making and marine resource management in the country (Walton and others 2014). In the Philippines, marine resource management began with a centralized, top- down, and use-oriented structure (Alcala and Russ 2006). Such early Philippine policies encouraged greater use of natural resources, which lead to depletion and habitat degradation. With the top-down approach, management responsibility often fell upon the central government, or government bureaucracies centred around large cities such as Manila and Cebu (Pomeroy and Carlos 1997; Alcala and Russ 2006). Unfortunately, in the case of the Philippines, this top-down approach to management was mostly ineffective, as the governing bodies were unable to properly manage resource exploitation and the expansion of f isheries (which included destructive and illegal f ishing methods) in the country (Alcala and Russ 2006). However, in recent times, the responsibilities and power to establish marine resource policy in the Philippines has since shifted towards community-based co- management, which involves the municipal LGUs and, more importantly, the primary resource users themselves—the local f ishers and coastal communities. A number of well-enforced MRs built on community co-management and collaboration have been documented in the Philippines, although these have only covered specif ic localities throughout the country (White and Courtney 2002; Alcala and Russ 2006; JA Anticamara and others 5 Arceo and others 2008; Cabral and others 2014). Resource co-management tends to have better continuity over human generations, particularly if the local communities enforcing these policies are convinced of its effectiveness and have a strong desire to participate (Alcala and Russ 2006). Conversely, a lack of belief and participation in the management system could easily lead to non-compliance and resistance (Oracion and others 2005). There is generally a lack of standards in managing MRs among Philippine municipal governments, and the quality of management can vary with the skills and interests of local off icials (White and Courtney 2002). Inconsistencies in enforcement may be limiting the effectiveness of marine resource policies across the country, and need to be addressed, especially with the turnover of jurisdictions with every change in local government administration after elections. Current national policy devolves biodiversity conservation and management effort in the Philippines to the municipal level. For example, the Local Government Code (LGC) of 1991 provided municipal Local Government Units (LGUs) with authority to carry-out specif ic functions, including the establishment of policies regarding the conservation and management of natural resources, such as the establishment of reserves and protected areas (Philippine Government 1991). In addition, the Republic of the Philippines Fisheries Code Republic Act (RA) 8550 states that all Philippine municipalities must allocate about 15% of its municipal waters (i.e. , coastal waters from foreshore to 15 km away from the coasts) as MRs (Department of Agriculture 1998). However, while the number of well-enforced MRs has increased over the years (Maypa and others 2012), recent estimates suggest that only about 1% of Philippine coral reef areas are well-protected (White and others 2014), and 90% of the 1,000+ MRs currently existing in the Philippines are small or < 1km2 (Weeks and others 2010). Despite the pressing need to improve coastal resource management in the country (Weeks and others 2014), the current status of Philippine reef f ish biodiversity remains largely unmeasured, except for surrogate data from a few sites, or select (usually commercial) families and species (Russ and Alcala 2004; Russ and others 2005; Stockwell and others 2009). The main goal of this paper is to present the results of a recent and systematic assessment of Philippine reef f ish diversity, accounting for reef f ish species richness, relative abundance, and relative biomass across representative sites throughout the country. In addition, this paper will explore patterns of reef f ish diversity and dominance across the Philippines. It is not within the scope of this paper to quantify the effects of municipal-level management on Philippine reef f ish diversity, but rather to discuss how diversity patterns revealed in the study are potentially related to the existing municipal-level “devolution of power” of marine resource National Patterns of Philippine Reef Fish Diversity 6 management in the country to date. Therefore, much of the analysis in this paper on Philippine reef f ish biodiversity patterns will be conducted at the municipal level (e.g. , comparing biodiversity between and across municipalities). Specif ically, the paper seeks to answer the following questions: (1) What is the general picture of reef f ish biodiversity throughout the Philippines to date, based on different biodiversity metrics (e.g. , species richness, evenness, abundance, and biomass)?; (2) How does reef f ish biodiversity vary across Philippine municipalities based on these metrics?; (3) What patterns can be observed in reef f ish assemblages throughout the country?; and (4) What types of f ish species dominate Philippine coral reefs to date? METHODS Study site selection and survey methods Using Google Earth satellite images, we selected sites that most likely had coral reefs close to shore. In addition, study sites were selected to represent the Philippines’ three major island groups (e.g. , Luzon, Visayas, and Mindanao) and the six marine biogeographic regions proposed by Aliño and Gomez (1994), while accounting for budget and logistical constraints such as travel time, costs, issues of site accessibility, traveling with lots of equipment, and safety. Surveyed reef sites within each municipality included areas inside and outside MRs (where MR boundary demarcation was clearly established), and were often referred by local f ishers, boatmen, or Local Government Unit (LGU) off icers, whom we asked to direct us to reef areas where we could record as much of the local f ish diversity as possible. Due to budget and logistical limitations, the number of surveyed transects varied per municipality and biogeographic region (Appendix 1). To quantify Philippine reef f ish diversity, standardized Underwater Visual Census (UVC) belt transects surveys were conducted throughout the Philippines. A total of 420 belt transects, belonging to 119 reef sites, forty-nine municipalities, and six Philippine marine biogeographic regions were surveyed from March 2012 to June 2014 (approximately two-year period), spanning north to south of the Philippines (Figure 1). The UVC belt transect method used is an established non-destructive reef f ish survey method, for quantifying reef f ish species diversity (Brock 1982; Samoilys 1997; Samoilys and Carlos 2000). To conduct UVC surveys, a diver (J. Anticamara) swam along a 20 x 5 m transect and recorded all size (cm) and abundance estimates of non-cryptic reef f ish species with a minimum length of 1 cm encountered within the transect boundaries. JA Anticamara and others 7 Typically, a minimum length of 10-11 cm is recommended to avoid errors in length and abundance estimates (Bellwood and Alcala 1988). However, we initially observed that many of our survey sites were dominated by small-bodied species and individuals, so setting the minimum length of our methods to 10 cm would exclude a signif icant portion of the reef f ish community. Therefore, we decided to include all reef f ish species down to a minimum length of 1 cm, to appropriately represent the true status of reef f ish diversity throughout our survey sites. The estimated length of recorded individual f ish species was later converted into weight using Length-Weight (LW) relationships available in FishBase (Froese and Pauly 2014). In cases where the LW relationship of a particular f ish species was not Figure 1. Map of surveyed reef sites throughout the Philippines. The broken lines represent demarcations of the Philippine marine biogeographic regions as proposed by Aliño and Gomez (1994). The number of transects and municipalities surveyed per biogeographic region can be found in Appendix 1. National Patterns of Philippine Reef Fish Diversity 8 available in FishBase, the LW of the congener or family member of similar shape and maximum total length was used (Anticamara and others 2010). All surveyed transects were conducted at depths ranging from 3–6 m to capture as much f ish diversity as possible at a manageable depth, since reef f ish diversity and abundance are often high at this depth range relative to other depth ranges (Friedlander and Parrish 1998; Friedlander and others 2003). To avoid variation due to surveyor’s error, we only analyzed UVC survey data obtained by one of the authors (J. Anticamara), who has had nearly twenty years of experience conducting underwater surveys in Philippine coral reefs (Samoilys and others 2007; Anticamara 2009; Anticamara and others 2010). All surveys were conducted during daylight hours, and each transect was surveyed for approximately 20 minutes. Our choice of transect dimensions (20 x 5 m), number of transect replicates (3-4 transect replicates per reef site, or 8–10 transects per municipality) and total surveyed reef area per site (300–400 m2total reef area per site, or 800–1,000 m2 per municipality) is comparable to UVC methods used in other studies quantifying reef f ish diversity (Brock 1982; Friedlander and Parrish 1998; T issot and others 2004; Nakamura and Tsuchiya 2008; Shibuno and others 2008; Honda and others 2 0 1 3 ) . The UVC belt transect method underestimates the abundance of cryptic reef f ish species (e.g. , Blennies, Gobies, Dottybacks, and Eels) and nocturnal species (e.g. , Sweepers, Soldierf ishes, and Priacanthids), since such species may remain hidden from census divers (Willis 2001). On the other hand, highly-mobile species may be overestimated, due to their conspicuous movements in the diver’s f ield of vision (Smith 1988). To address these limitations, we conducted UVC surveys at slow swim speeds of about 5 m2 min-1 (or roughly 20 min per 100 m 2 transect), which improves counting accuracy, search eff iciency (Samoilys and Carlos 2000), and avoids scaring away skittish f ish, while taking care not to double-count individuals that re-enter the transect area. In addition, photographs of all encountered reef f ish species were taken for identif ication verif ication using a number of references (Allen and others 2003; Kuiter and Debelius 2006; Froese and Pauly 2014). Data analysis First, to present the adequacy of our current sampling effort in capturing Philippine reef f ish diversity, we constructed Species Accumulation Curves (SACs) for each of the six sampled biogeographic regions. A SAC is a graph of recorded number of JA Anticamara and others 9 species as a function of sampling effort and allows for the estimation of the total number of species in a given area per increased sampling unit (Colwell and others 2004). Initially, the SAC rises steeply as common and abundant species are recorded, then more slowly as rare species are recorded (Ugland and others 2003). To examine general patterns of reef f ish diversity across the Philippines, histograms of reef f ish species richness, abundance, and biomass per municipality were constructed. Then, to examine potential differences in reef f ish diversity between municipalities across the country, bar plots showing mean (and standard errors SE) species richness, abundance, and biomass per municipality were also produced. Examining spatial trends at the municipal level coincides with the paper’s objectives to discuss our research f indings in relation to the current municipal-level policy of Philippine coastal management. To explore patterns of reef f ish assemblages throughout the country, non-metric Multidimensional Scaling (MDS) analysis was performed to help visualize potential grouping-by-similarity of reef f ish assemblages at various spatial scales, namely: transects, reef sites, municipalities, or marine biogeographic regions. MDS analysis uses a constructed Bray-Curtis similarity matrix to visually map the similarities of the 420 sampled transects—i.e. , transects that are more similar are plotted closer- together, while transects that are dissimilar are plotted further apart. For example, if transects from a given biogeographic region grouped more closely-together than with transects from other biogeographic regions, this would suggest that reef f ish assemblages in that biogeographic region are distinct from the other biogeographic regions. Similarly, if transects from a given municipality grouped more closely together than with transects from other municipalities, this would suggest that reef f ish assemblages in sampled transects within that municipality are similar and are distinct from transects sampled from the other municipalities. MDS analysis would therefore allow us to determine if transects grouped at certain spatial scales or did not show any clear grouping at all. MDS analysis was performed separately for reef f ish assemblage similarity based on reef f ish abundance and biomass data. To further examine reef-f ish assemblages at multiple spatial scales, additive diversity partitioning was performed. In additive diversity partitioning, total diversity (�) is the sum of the mean local diversity or the mean diversity within samples or transects ( ), and the diversity between samples (�) at various def ined scales (e.g. , between transects, reef sites, municipalities, or biogeographic regions). In an unbalanced, hierarchal sampling design, such as in our case, each sample level can be represented as hierarchal spatial scales (Veech and others 2002). Specif ically, in this study, represents mean within-transect reef f ish diversity, �1 represents � _ � _ National Patterns of Philippine Reef Fish Diversity 10 1 —∑ pijklnpijk 2 between-transec t d i versity, � 2 represents between-reef site d i versity, � 3 represents between-municipality diversity, �4 represents between-biogeographic region diversity, and � represents the estimated total Philippine reef f ish diversity based on all our samples, as summarized in the following equations: where is the mean diversity for each hierarchal spatial scale. Estimated total Philippine diversity (�) based on all our samples can be expressed as: (5) Therefore, diversity partitioning can be used to determine the proportional contributions (percentage) of each level of and �-diversity to �-diversity, wherein total diversity � = 100% (Lande 1996). However, the interpretation of and � varies depending on the particular diversity index used. The general equation for is presented below: (6) Based on the above equation, the sample weight q ij is the proportion of the total number of individuals found in each sample j, and sampling level i. In addition, can be calculated using different diversity indices D ij . When partitioning is based on species richness index, D ij is the number of species in transect j, at sampling level i. When par titioning is based on Shannon’s Diversity index, D ij = , where p ijk is the proportional abundance of species k in transect j. Similarly, when partitioning is based on Simpson’s Diversity index, D ij = (Crist and others 2 0 0 3 ) . PARTITION v2 freeware (Veech and Crist 2007) was used to run additive diversity partitioning analysis, and to test for signif icant differences between our data (e.g. , “observed diversity”) and randomly-generated null models (e.g. , “expected diversity”) (Veech and Crist 2007). Null models were generated based on 999 � _ � _ �1transects = Dsites — transects� __ �3municipalities = Dbiogeographic—Dmunicipalities _ _ �4biogeographic = � Dbiogeographic _ � = � + �1 + �2 + �3 + �4 • � = ∑ _ n i t = 1 D ij q ij 1 —∑ pijklnpijk �2sites = Dmunicipalities—Dsites _ _ _ D (1) (2) (3) (4) 0 � _ � _ JA Anticamara and others 11 individual-based randomizations. A signif icant difference between the observed data portioning and randomized data partitioning means that the observed hierarchical patterns of diversity are real and unlikely to be produced by a randomized assignment of species to samples at various def ined scales. Finally, to examine dominance patterns in reef f ish assemblages, a Similarity Percentage (SIMPER) analysis was run separately for reef f ish species abundance and biomass. SIMPER determines the contributions of each reef f ish species to the pair-wise Bray-Curtis similarities of all surveyed transects within each municipality. The species with the highest contributions are usually the most abundant or have the highest total biomass and therefore are generally the most dominant species. Both MDS and SIMPER analyses were run using Primer v6 (Clarke and Gorley 2006). The body sizes (maximum total length (max TL)) of the top f ive dominant reef f ish species for all municipalities was obtained from FishBase and categorized as small-bodied (max TL < 10 cm), medium-bodied (max TL = 10.1 - 30 cm), and large-bodied (max TL > 30.1 cm). RESULTS Overall, we identif ied a total of 375 non-cryptic reef f ish species, belonging to forty-eight families, across the 420 transects that we surveyed throughout the entire Philippines. Species accumulation curves per biogeographic region showed increasing number of species with every additional transect (Figure 2). However, the rate of increase in the number of species per additional transect started to plateau or visibly slow-down at around 20-30 transects per biogeographic region, at which point over 100 reef f ish species had been recorded. This suggests that most common or dominant species in each biogeographic region have been recorded after surveying about 20–30 100 m2 transects, and additional species detected by surveying more transects may be rare or cryptic species. Patterns in the frequency distribution of transects with respect to mean reef f ish species richness (22.2 ± 0.8 species), abundance (387.7 ± 66.1 f ish), and biomass (2.5 ± 0.3 kg) differed. Transects were normally distributed in terms of species richness (Figure 3a). On the other hand, transects were skewed to the left in terms of both abundance and biomass, indicating that most transects had abundance and biomass values way below the mean for both metrics (Figure 3d-e). Bar plots on mean species richness, abundance, and biomass per municipality showed generally lower variation in species richness within and among municipalities, but 0 National Patterns of Philippine Reef Fish Diversity 12 higher variation in terms of both abundance and biomass (Figure 4). Most municipalities (61% of forty-nine municipalities) had high species richness (i.e. , “high” def ined as having values within or above the range values of the mean ± SE across all municipalities, and “low” def ined as having values below it). On the other hand, most municipalities had low abundance (63%) and biomass (59%). MDS analysis of the Bray-Curtis Similarity in reef f ish species assemblages with respect to species relative abundance or relative biomass did not show clear grouping of surveyed transects according to either biogeographic regions or municipalities (Appendix 2). However, some transects from the same municipality tended to group together, indicating within-municipal similarity of reef f ish assemblages, at least for those municipalities. Additive par tition of reef f ish diversity in terms of species richness and species evenness (e.g. , Shannon’s diversity and Simpson’s diversity) showed different spatial patterns of Philippine reef f ish biodiversity. In terms of species richness, observed �1 and �2-diversity did not signif icantly differ from the expected null model, while observed �3 and �4-diversity were signif icantly higher than the expected Figure 2. Species Accumulation Curves (SACs) per biogeographic region, for the Visayas Sea (a), Northern Philippine Sea (b), Sulu Sea (c), Southern Philippine Sea (d), South China Sea (e), and Celebes Sea (f ). SAC curves show the cumulative number of f ish species found in every additional transect based on 1,000 permutations of transect ordering. JA Anticamara and others 13 null model. Furthermore, most of species richness � -diversity was accounted for by �3 (37.7%) and �4 (41.8%). In contrast, accounted for much of Shannon’s (50.0%) and Simpson’s (81.13%) �-diversity. Dominance analysis using SIMPER indicated that each municipality generally had different sets of top f ive dominant species in terms of species’ relative abundance (Appendix 3a) and biomass (Appendix 3b). The top f ive dominant species within each municipality generally accounted for about 73.7 ± 1.6% of the total abundance Figure 3. Histograms showing per-transect frequency distributions for reef f ish species richness (a), Shannon’s diversity (b), Pielou’s evenness (c), abundance (d), and biomass (e). The broken line on each graph represents the mean value per transect for that respective metric. � _ National Patterns of Philippine Reef Fish Diversity 14 and 65.6 ± 2.0% of the total biomass. Of the 375 reef f ish species recorded in our study, only 66 and 68 species comprised the top f ive dominant species in terms of abundance and biomass for all municipalities, respectively. Certain reef f ish families tended to dominate the top f ive species per municipality. For example, of the 66 species included in the dominant species listed for all municipalities based on Figure 4. Bar plots showing per-municipality mean ± SE bars for reef f ish species richness (a), Shannon’s diversity (b), Pielou’s evenness (c), abundance (d), and biomass (e), arranged by biogeographic region, and from north to south of the Philippines. Municipality codes can be found in Appendix 1. JA Anticamara and others 15 abundance, 58% were Damself ishes and 13% were Wrasses. In terms of biomass, 18% of the 68 top f ive dominant species were Wrasses, 16% were Damself ishes, and 13% were Butterflyf ishes. Fur thermore, many of these dominant species in terms of both abundance and biomass were small to medium-bodied species. In terms of abundance, small-bodied, medium-bodied, and large-bodied species made up 36%, 58%, and 6% of the top f ive dominant species, respectively. In terms of biomass, small-bodied, medium-bodied, and large-bodied species made up 9%, 57%, and 34% of the top f ive dominant species, respectively. However, most of the medium to large-bodied species that appeared as top f ive dominant in terms of biomass were mainly omnivores, herbivores, and corallivores, suggesting the decline of most large-bodied carnivorous reef f ish species throughout the Philippines. DISCUSSION Overall, results from this research show that many coral reef areas throughout the Philippines are still highly diverse, only if reef f ish species richness is used as the sole measure of biodiversity. However, while still considerably species-rich (i.e. , most municipalities having at least 21-23 species per 100 m2), many areas throughout the Philippines appear to be exhibiting signs of depletion in terms of f ish abundance and biomass, and in fact, previous studies have described the depletion of species richness as well (Lavides and others 2009; Nañola Jr. and others 2011). Diversity partitioning analysis revealed that Philippine reef f ish assemblages can be characterized as having high variations in species richness across municipalities, but generally low species evenness (e.g. , Shannon’s and Simpson’s diversity indices). Differences in species richness rather than evenness best explained differences in reef f ish assemblages between municipalities, suggesting the presence of restricted-range species (i.e. , species found in only a few of the surveyed municipalities) in each municipality (Go and others in press). On the other hand, Shannon’s and Simpson’s diversity were best explained by within- transect diversity, suggesting that most surveyed reefs were dominated by a few, highly-abundant reef f ish species. Further investigation of dominance patterns via SIMPER analysis revealed that most surveyed reef sites were dominated by abundant small and medium-bodied reef f ish species. However, there was also a general rarity of large-bodied species throughout most Philippine reefs—a f inding which suggests overexploitation due to f ishing, considering that large-bodied, high trophic-level species are particularly targeted by f isheries (Pauly and others 1998), and are especially vulnerable to f ishing due to particular life history traits (Abesamis and others 2014). These f indings suggest that the current “municipality-by- National Patterns of Philippine Reef Fish Diversity 16 municipality” policy to biodiversity conservation in the Philippines may be affecting reef f ish assemblages throughout the country (but see qualif ied discussions and elaborations of this point below). Although previous work accounting for Philippine reef f ish abundance, biomass, and functional diversity has been done (Carpenter and others 1981; Russ and Alcala 1998a; Russ and Alcala 1998b; Nanola and others 2006), to date, the most common measure of reef f ish diversity in the Philippines is species richness (Allen 2002; Carpenter and Springer 2005; Allen 2008; Nañola Jr. and others 2011). Species richness or species presence-absence data is useful in estimating species range, restriction or expansion of range, and potential local extirpation (Lavides and others 2009; Nañola Jr. and others 2011). However, results of the current study show that species richness alone is not always a good indicator of reef f ish diversity status in the country. For example, while reef f ish species richness remains high in most places throughout the Philippines, examination of the other metrics reveals that most places in the country actually have low reef f ish abundance and biomass. Indeed, other studies have also found that reef f ish species richness may exhibit less-obvious changes than reef f ish species abundance, in response to disturbances such as exploitation and habitat degradation (Alcala 1988; Harmelin and others 1995; Jones and others 2004)—human-induced disturbances that are common in many coastal areas of the Philippines. Therefore, the effects of such disturbances on reef f ish assemblages may be underestimated, if only species richness is taken into account. Many species still exist, but most in very low population size or abundance throughout the sampled municipalities. Patterns of Phil ippine Reef Fish D iversity: Restricted-range Species and the Dominance of a Few Highly-abundant Species Results of additive partitioning analysis suggest two main findings regarding spatial patterns of reef f ish assemblages throughout the country: (1) the presence of restricted-range species influences the differences in species richness between municipalities; and (2) only a few, abundant species tend to dominate reef f ish assemblages throughout the country, and greatly influence species evenness. With regards to our f irst f inding—additive diversity partitioning analysis showed that between-municipality (�3) and between-biogeographic region (�4) diversity species richness was signif icantly greater than that predicted by the null models, and also accounted for a relatively large portion of � -diversity. This means that differences in reef f ish species richness between municipalities and between biogeographic regions may reflect real variations in species richness at these spatial scales JA Anticamara and others 17 (Belmaker and others 2008). The high contribution of �3-diversity to �-diversity in terms of species richness indicates that the presence of restricted-range and rare species may account for the difference in species richness between municipalities (Rodríguez-Zaragoza and others 2011). Indeed, many of the reef f ish species included in our study exhibited restricted ranges (Go and others in press). However, we suspect that the restricted ranges of many of these species is not due to evolutionary or ecological factors, considering the nationwide distributions of most of these species based on previous records (Carpenter and Springer 2005; Nañola Jr. and others 2011; Froese and Pauly 2014), but rather due to the high rates of exploitation and reef degradation in the Philippines to date. With regards to our second f inding of diversity partitioning analysis—it is possible to infer that most surveyed areas were dominated by a few, highly-abundant species, because -diversity accounted for much of Shannon’s and Simpsons’ � -diversity, but not for species richness’ � -diversity (Rodríguez-Zaragoza and others 2011). High -diversity when accounting for species relative abundance (e.g. , evenness metrics like Shannon’s and Simpson’s indices) can be interpreted as homogeneity of species assemblages across surveyed transects, because the very high abundance of a few species common to all sites overwhelms the small amounts of between- site (�) variation contributed by the non-abundant species (Rodríguez-Zaragoza and others 2011). These f indings suggest that surveyed reef f ish assemblages per municipality are generally characterized by high species richness, but low evenness (Rodríguez-Zaragoza and others 2011). The differences in observed patterns from diversity partitioning analysis between species richness and evenness again highlights the importance of measuring biodiversity using different metrics (Gering and others 2003). Dominance Patterns in Phil ippine Reef Fish Assemblages: The Abundance of Small and Med ium-bod ied Species Analysis of reef f ish species assemblages based on Bray-Curtis SIMPER showed that most of the dominant species in surveyed reefs were small and medium- bodied species such as Wrasses, Damself ishes, and Butterflyf ishes, in terms of abundance. Although some large-bodied species were among the top f ive dominant species in terms of biomass, this does not necessarily mean that these species are abundant in Philippine reefs—indeed, large-bodied reef f ish species such as Emperors, Groupers, Jacks, Snappers, and Sweetlips were rarely dominant in terms of abundance across all surveyed reefs. This may be due to the fact that: (1) larger maximum body size—along with other life history traits such as slower growth � _ � _ National Patterns of Philippine Reef Fish Diversity 18 rate, longer lifespan, later age at maturity, and lower rates of natural mortality—has been associated with increased vulnerability to f ishing (Abesamis and others 2014); and (2) large-bodied f ish species are especially targeted by f ishers for their higher commercial value than small-bodied species (Russ and Alcala 1996; Pauly and others 1998). Shifts in f ish assemblages from dominance of larger-bodied species and individuals towards dominance of smaller-bodied species and individuals have been documented in the past, following high levels of exploitation (Greenstreet and Hall 1996; Bianchi and others 2000; Rogers and Ellis 2000; Levin and others 2006). Thus, high exploitation rates in the Philippines, accompanying increasing demands for f ish production and a growing human and f ishing population, may be threatening most commercially-important, large-bodied reef f ish species in the country. The exploitation-induced depletion of large-bodied reef f ish species may have negative implications on Philippine f isheries production and the food security of many Filipinos, who are largely-dependent on f ish products as a dietary protein source, and actually prefer to consume large-bodied f ish species (BFAR 2012). However, formal assessments on the threatened status of many reef f ish species in the Philippines are limited by the lack of available species abundance and distribution data in the past and recent years. This makes assessment criteria commonly used by internationally-recognized conservation organizations like the IUCN (such as population decline and range contraction) diff icult to apply for many reef f ish species in the Philippines. As a result, many Philippine reef f ish species remain under-assessed or totally unassessed (Go and others in press; IUCN 2014)—an issue that should be addressed by conservation and management efforts in the country. Caveats and Limitations The main caveat of the current study is that the number of surveyed transects varied across municipalities and biogeographic regions. This could lead to under- representation of reef f ish diversity for biogeographic regions that had a disproportionally fewer number of surveyed transects than the other surveyed regions (e.g. , Celebes Sea, in our study). Nañola Jr. and others (2011), who presented reef f ish species richness patterns across Philippine marine biogeographic regions, showed with SACs that the number of species recorded per additional transect surveyed increased rapidly until about 40 to 50 transects per biogeographic region, after which the addition of new species recorded per additional transect slowed down. This suggests that around 40 to 50 surveyed transects are required to account JA Anticamara and others 19 for most of the common or abundant species in each biogeographic region. Based on this estimate, reef f ish diversity in the Celebes Sea biogeographic region is under- represented in our study. However, based on our own SACs and data, our sampling effort adequately captured reef f ish diversity for all biogeographic regions, as all of our SACs per biogeographic region approached asymptotic patterns. In addition, we surveyed reef sites and municipalities that were geographically far apart, and selected sites referred by local f ishers, boatmen, or LGU off icers to capture as much representative reef f ish biodiversity per municipality and per biogeographic region as possible. However, despite these efforts to survey as much reef f ish diversity as possible, none of our SACs approached the 350–500 reef f ish species recorded per biogeographic region at 40-50 transects reported by Nañola Jr and others (2011), even after we re-ran the SAC construction using Jackknife 2 estimators —e.g. , the estimator used by Nañola Jr. and others (2011), which is based on species presence-absence data (Smith and Pontius 2006). This may suggest a general depletion of reef f ish diversity throughout the Philippines (Lavides and others 2009; Nañola Jr. and others 2011), considering that Nañola Jr. and others (2011) included reef f ish survey data from 1991 to 2008. To account for the differences in the number of transects per municipality and biogeographic region when conducting diversity partitioning analysis, we used an unbalanced sampling design in PARTITION v2 (Veech and Crist 2007). Unbalanced sampling in diversity partitioning has been used in previous studies on reef f ish diversity patterns as well, where sampling effort was not uniform across study areas (Rodríguez-Zaragoza and Arias-Gonzalez 2008; Francisco-Ramos and Arias- González 2013). Impl ications for Management The observed patterns in reef f ish assemblages throughout the country may be affected by the municipal-level organization of coastal management in the Philippines today. For example, signif icant differences in diversity metrics (particularly abundance and biomass) between municipalities, as well as the presence of restricted-range species in each municipality, may be potentially due to the variations in management effectiveness (e.g. , MR enforcement) between these municipalities (although this is not tested in the current study). The positive effect of well-enforced MRs on local f ish diversity has been documented in previous studies (Russ and Alcala 1999; Walmsley and White 2003; Samoilys and others National Patterns of Philippine Reef Fish Diversity 20 2007; Maypa and others 2012; Bergseth and others 2013), and it is highly possible that surveyed municipalities that exhibited high diversity metrics were also those municipalities that had well-enforced MRs. However, quantifying the effects of varied management on reef f ish diversity across municipalities is diff icult given available datasets, since only 14 of the 49 municipalities included in our study had MRs with available enforcement ratings on the recently established Philippine Marine Protected Area Network (Cabral and others 2014). In addition, management effectiveness may be linked to the interest and support of stakeholders. For example, the distribution of MRs in the Philippines is concentrated in the Visayas region (Weeks and others 2010)—a region where academic institutions and non-government organizations (NGOs) continue to support MR establishment (Pollnac and others 2001), and where the f irst efforts of Philippine MR establishment began (Alcala and Russ 2006). While much has been done to quantify the extent of MR establishment and enforcement throughout the country (Weeks and others 2010; Maypa and others 2012), the effectiveness in terms of biological indicators (e.g. , reef f ish species abundance, biomass, and f ish yield) of most Philippine MR’s is still largely unknown. Maypa and others (2012) presented the most recent analysis of Philippine MR effectiveness on coral reef health, but only included a few (n = 56) MRs from the Visayas region that had available biophysical data. Thus, there is still a great need to monitor biological indicators of MR effectiveness throughout the Philippines. To date, the Coral Triangle Marine Protected Area System (CTMPAS), created by the Coral Triangle Initiative (CTI) in 2009, hopes to achieve well-managed MPAs throughout the six coral triangle countries by integrating the aforementioned ecological, social, and governance factors through a consistent and science-based system of MR establishment (Walton and others 2014). However, there is still a great need to improve the enforcement, monitoring, socioeconomic accountability, governance, and f inancial support of many MRs in the Philippines (White and others 2014). In addition, facilitating collaboration and communication between multiple stakeholders, increasing local capacity to manage MRs, and developing learning networks across MR managers are invaluable in achieving successful MR enforcement (Weeks and others 2014). Finally, it is important to account for ecological factors in MR design, such as adequate habitat representation, protection of critical areas used in a species’ different life history stages (e.g. , spawning grounds, nurseries), ensuring connectivity between protected habitats, accounting for resilience or vulnerability to climate change, and minimizing local anthropogenic threats (e.g. , land-based runoff and siltation) (Green and others 2014). For example, while current small-scale municipal-level management may be suff icient for restricted-ranged species, implementing large-scale (e.g. , across networks of MRs JA Anticamara and others 21 rather than at select, individual MRs) and long-term management and monitoring of reef f ish diversity would help ref ine and adjust marine biodiversity conservation strategies for the country, effectively manage species that traverse large spatial units beyond municipal boundaries (e.g. , large-bodied species such as Groupers, Snappers, Sharks, and Whales, etc.), and ensure proper connectivity of reef f ish diversity throughout the country (Kramer and Chapman 1999; Beets and others 2003; Lowry and others 2009; Matias and others 2013; Green and others 2014). Results from this research provide the most recent analysis on the current status of reef f ish diversity throughout the Philippines using a standardized or systematic survey strategy. The results and conclusions from this research suggest that there is a great need to fully enforce the current marine biodiversity conservation policies of the Philippines, to conduct national coral reef assessments that are systematic, scientif ically-sound, well-organized (Licuanan and Aliño 2014), and to mitigate reef degradation and the depletion of valuable marine biodiversity resources. By ensuring that 15% of Philippine municipal waters receive effective protection from further overexploitation and destructive f ishing (e.g. , dynamite f ishing and the use of poison), the remaining reef areas of the Philippines will have some chance of recovery, which will allow them to continue to provide benef its to Philippine f isheries and food security, and maintain the high levels of diversity in the country (Russ and others 2004; Russ and others 2005; Anticamara and others 2 0 1 0 ) . ACKNOWLEDGMENTS We would like to thank the mayors and local government units and agencies of all the municipalities included in this research for their support. We also would like to thank the Local People’s Organizations in Bolinao (Kisahan ng mga Samahan Alay sa Kalikasan [KAISAKA]), Masinloc (Samahang Pangkaunlarang San Salvador [SPSS]), and Mabini (Samahang Pangkaunlarang San Teodoro [SPSTI]). Special thanks to the state universities who helped in this research: Ateneo de Naga (Joanaviva Plopeni and Shane Bimeda), Bicol State University (Karina Luth Discaya, Antonino Mendoza, Meek Salvador), Mariano Marcos University (Wilnorie Rasay), Palawan State University (Arselene Bitara and Dr. Michael Pido), and University of Eastern Philippines (Saula Gabona). We also thank two Research Assistants who helped in the f ield work: Ambrosio Melvin Matias and Justin Albert de Ramos. Funding for this research comes from the following: Foundation for the Philippine Environment (FPE), Off ice of the Vice Chancellor for Research and Development UP OVCRD (111104 PhDIA), Natural Sciences Research Institute UP NSRI (Project Code: Bio-11-1-08), and the Center for Integrative and Development Studies UP CIDS. National Patterns of Philippine Reef Fish Diversity 22 REFERENCES Abesamis RA, Green AL, Russ GR, Jadloc CRL. 2014. The intrinsic vulnerability to f ishing of coral reef f ishes and their differential recovery in f ishery closures. Rev Fish Biol Fish. 24:1033–1063. Alcala AC. 1988. Effects of marine reserves on coral f ish abundance and yields of Philippine coral reefs. Ambio. 17:194–199. Alcala AC and Russ GR. 2006. No-take marine reserves and reef f isheries management in the Philippines: a new people power revolution. Ambio. 35:245–254. Aliño PM, Gomez ED. 1994. Philippine coral reef conservation: its signif icance to the S o u t h C h i n a S e a . 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Establishing a functional region-wide Coral Triangle marine protected area system. Coast Manag. 42:107–127. Weeks R, Aliño PM, Atkinson S, and others. 2014. Developing marine protected area networks in the Coral Triangle: good practices for expanding the Coral Triangle marine protected area system. Coast Manag. 42:183–205. Weeks R, Russ GR, Alcala AC, White AT. 2010. Effectiveness of marine protected areas in the Philippines for biodiversity conservation. Conserv Biol. 24:531–540. National Patterns of Philippine Reef Fish Diversity 28 _____________ Jonathan A. Anticamara is Associate Professor and Head of the JA Lab Group at the Institute of Biology, University of the Philippines, Diliman. He is lead author in the Intergovernmental Platform on Biodiversity and Ecosystem Services (Deliverable 3c). He currently holds the Energy Development Corporation Chair on Biodiversity Research. Kevin Thomas B. Go is a Research Associate at the JA Lab Group, where he has been working since his graduation from the Institute of Biology, UP Diliman in 2012. Steven S. Ongsyping is a medical student at the University of Santo Tomas Faculty of Medicine and Surgery. Before his graduation from the Institute of Biology, UP Diliman in 2013, he worked at the JA Lab Group for his undergraduate thesis on reef f ish diversity. Francesco Antonio T. Valdecañas is a medical student at the University of the Philippines College of Medicine. Before his graduation from the Institute of Biology, UP Diliman in 2013, he worked at the JA Lab Group for his undergraduate thesis on coral and reef f ish relationships. Ryan Gabriel S. Madrid is an environmental consultant at Berkman International Inc. Before his graduation from the Institute of Biology, UP Diliman in 2013, he worked at the JA Lab Group for his undergraduate thesis on coral diversity and distribution. White AT, A l i ñ o P M , Cros A , and others. 2014. Marine protected areas in the co r a l triangle: progress, issues, and options. Coast Manag. 42:87–106. White AT and Cour tney CA . 2002. Experience with marine protected area planning and management in the Philippines. Coast Manag. 30:1–26. White AT, Vogt HP. 2000. Philippine coral reefs under threat: lesson learned after 25 years of community-based reef conservation. Mar Pollut Bull. 40:537–550. Willis TJ. 2001. Visual census methods underestimate density and diversity of cryptic reef f ishes. J Fish Biol. 59:1408–1411. JA Anticamara and others 29 1.01 Pagudpod 2 South China Sea 1.02 Burgos 2 South China Sea 1.03 Curimao 6 South China Sea 1.04 Sinait 4 South China Sea 1.05 Bolinao 19 South China Sea 1.06 Alaminos 25 South China Sea 1.07 Masinloc 11 South China Sea 1.08 El Nido 5 South China Sea 1.09 Quezon 6 South China Sea Total 80 South China Sea 2.01 Sta. Ana 18 North East Philippine Sea 2.02 Baler 8 North East Philippine Sea 2.03 Caramoan 8 North East Philippine Sea 2.04 Tabaco 7 North East Philippine Sea 2.05 Lavesarez 4 North East Philippine Sea 2.06 Catarman 6 North East Philippine Sea 2.07 Laoang 4 North East Philippine Sea Total 61 North East Philippine Sea 3.01 Mabini 17 Visayas Sea 3.02 Puerto Galera 9 Visayas Sea 3.03 Bongabong 5 Visayas Sea 3.04 Romblon 6 Visayas Sea 3.05 San Fernando 5 Visayas Sea 3.06 Mandaon 6 Visayas Sea 3.07 Cataingan 6 Visayas Sea 3.08 Malay 3 Visayas Sea 3.09 Buruanga 6 Visayas Sea 3.10 Inopacan 11 Visayas Sea 3.11 Getafe 5 Visayas Sea 3.12 Tubigon 6 Visayas Sea 3.13 Calape 2 Visayas Sea 3.14 Panglao 6 Visayas Sea 3.15 Mambajao 5 Visayas Sea 3.16 Mahinog 5 Visayas Sea Total 97 Visayas Sea Municipality Code Municipality Number of Transects Biogeographic Region Appendix 1 All surveyed municipalities, with corresponding codes and biogeographic regions National Patterns of Philippine Reef Fish Diversity 30 4.01 Anini-y 10 Sulu Sea 4.02 Nueva Valencia 8 Sulu Sea 4.03 Puerto Princesa 8 Sulu Sea 4.04 Bataraza 10 Sulu Sea 4.05 Bongao 14 Sulu Sea 4.06 Simunul 5 Sulu Sea Total 55 Sulu Sea 5.01 Lawaan 16 Southern Philippine Sea 5.02 Balangiga 6 Southern Philippine Sea 5.03 Giporlos 10 Southern Philippine Sea 5.04 Quinapondan 8 Southern Philippine Sea 5.05 Salcedo 6 Southern Philippine Sea 5.06 Guiuan 31 Southern Philippine Sea 5.07 Surigao 11 Southern Philippine Sea 5.08 Mati 23 Southern Philippine Sea Total 111 Southern Philippine Sea 6.01 Parang 4 Celebes Sea 6.02 Glan 6 Celebes Sea 6.03 Sarangani 6 Celebes Sea Total 16 Celebes Sea Municipality Code Municipality Number of Transects Biogeographic Region Appendix 1 All surveyed municipalities, with corresponding codes and biogeographic regions (cont’n.) JA Anticamara and others 31 Appendix 2 MDS plots of Bray-Curtis similarity among municipalities in terms of species abundance (a) and species biomass (b). Transects with the same shape denote transects from withinthe same biogeographic region. Municipality codes can be found in Appendix 1 National Patterns of Philippine Reef Fish Diversity 32 1.01 Thalassoma amblycephalum 34.48 Labridae 16.0 medium Similarity: 39.7 Chromis margaritifer 34.48 Pomacentridae 9.0 small Top 5: 93.1 Pomacentrus bankanensis 17.24 Pomacentridae 9.0 small Others: 6.9 Chaetodon kleinii 3.45 Chaetodontidae 15.0 medium Centropyge vroliki 3.45 Pomacanthidae 12.0 medium 1.02 Ctenochaetus striatus 20.33 Acanthuridae 26.0 medium Similarity: 15.0 Chromis margaritifer 16.26 Pomacentridae 9.0 small Top 5: 70.7 Thalassoma amblycephalum 16.26 Labridae 16.0 medium Others: 29.3 Plectroglyphidodon dickii 9.76 Pomacentridae 11.0 medium Chromis vanderbilti 8.13 Pomacentridae 4.5 small 1.03 Pomacentrus philippinus 25.09 Pomacentridae 10.0 small Similarity: 16.9 Ctenochaetus striatus 17.89 Acanthuridae 26.0 medium Top 5: 66.02 Thalassoma hardwicke 9.16 Labridae 20.0 medium Others: 33.98 Neoglyphidodon nigroris 8.19 Pomacentridae 13.0 medium Ctenochaetus cyanocheilus 5.69 Acanthuridae 13.7 medium 1.04 Plectroglyphidodon lacrymatus 28.91 Pomacentridae 10.0 small Similarity: 40.1 Pomacentrus philippinus 17.56 Pomacentridae 10.0 small Top 5: 78.9 Chromis margaritifer 13.19 Pomacentridae 9.0 small Others: 21.1 Pomacentrus lepidogenys 10.59 Pomacentridae 9.0 small Neoglyphidodon nigroris 8.62 Pomacentridae 13.0 medium 1.05 Chromis margaritifer 25.38 Pomacentridae 9.0 small Similarity: 12.1 Thalassoma hardwicke 12.72 Labridae 20.0 medium Top 5: 64.3 Plectroglyphidodon lacrymatus 10.20 Pomacentridae 10.0 small Others: 35.7 Ctenochaetus striatus 9.35 Acanthuridae 26.0 medium Coris batuensis 6.61 Labridae 17.0 medium 1.06 Pomacentrus chrysurus 22.66 Pomacentridae 9.0 small Similarity: 12.6 Neoglyphidodon melas 18.94 Pomacentridae 18.0 medium Top 5: 60.2 Plectroglyphidodon lacrymatus 8.01 Pomacentridae 10.0 small Others: 39.8 Macropharyngodon meleagris 5.98 Labridae 15.0 medium Stethojulis trilineata 4.64 Labridae 15.0 medium Appendix 3A Table showing the top 5 dominant species in terms of abundance, based on Bray-Curtis SIMPER analysis within municipalities, for each municipality, arranged from north to south of the Philippines. Names of each municipality and biogeographic region can be found in Appendix 1. Also shown are the average within-municipality similarity % (“Similarity”), the total contributory % of the top 5 dominant species for each municipality (“Top 5”), and the total contributory % of all other species not included in the top 5 (“Others”) Municipality Code Species Contributory % Family Max TL (cm) Body Size JA Anticamara and others 33 1.07 Ctenochaetus striatus 37.47 Acanthuridae 26.0 medium Similarity: 27.8 Chromis margaritifer 15.63 Pomacentridae 9.0 small Top 5: 80.4 Thalassoma hardwicke 14.48 Labridae 20.0 medium Others: 19.6 Stegastes fasciolatus 6.63 Pomacentridae 15.0 medium Plectroglyphidodon dickii 6.15 Pomacentridae 11.0 medium 1.08 Plectroglyphidodon lacrymatus 49.18 Pomacentridae 10.0 small Similarity: 19.4 Abudefduf sexfasciatus 7.94 Pomacentridae 16.0 medium Top 5: 78.1 Hemiglyphidodon plagiometopon 7.38 Pomacentridae 18.0 medium Others: 21.9 Thalassoma lunare 6.85 Labridae 25.0 medium Labroides dimidiatus 6.78 Labridae 11.5 medium 1.09 Plectroglyphidodon lacrymatus 35.52 Pomacentridae 10.0 small Similarity: 12.0 Neoglyphidodon nigroris 30.56 Pomacentridae 13.0 medium Top 5: 81.5 Amblyglyphidodon curacao 7.14 Pomacentridae 11.0 medium Others: 18.5 Apogon griffini 4.21 Apogonidae 13.5 medium Thalassoma lunare 4.08 Pomacentridae 25.0 medium 2.01 Ctenochaetus striatus 43.81 Acanthuridae 26.0 medium Similarity: 22.0 Chrysiptera rex 6.12 Pomacentridae 7.0 small Top 5: 66.1 Plectroglyphidodon lacrymatus 5.98 Pomacentridae 10.0 small Others: 33.9 Zanclus cornutus 5.96 Zanclidae 23.0 medium Pomacentrus coelestis 4.23 Pomacentridae 9.0 small 2.02 Ctenochaetus striatus 23.65 Acanthuridae 26.0 medium Similarity: 39.3 Chrysiptera rex 21.73 Pomacentridae 7.0 small Top 5: 83.8 Plectroglyphidodon lacrymatus 19.81 Pomacentridae 10.0 small Others: 16.2 Pomacentrus lepidogenys 14.23 Pomacentridae 9.0 small Pomacentrus bankanensis 4.35 Pomacentridae 9.0 small 2.03 Abudefduf sexfasciatus 14.18 Pomacentridae 16.0 medium Similarity: 21.1 Chaetodon octofasciatus 13.87 Chaetodontidae 12.0 medium Top 5: 56.0 Pomacentrus bankanensis 13.27 Pomacentridae 9.0 small Others: 44.0 Chrysiptera rex 7.93 Pomacentridae 7.0 small Amblyglyphidodon curacao 6.74 Pomacentridae 11.0 medium Appendix 3A Table showing the top 5 dominant species in terms of abundance, based on Bray-Curtis SIMPER analysis within municipalities, for each municipality, arranged from north to south of the Philippines. Names of each municipality and biogeographic region can be found in Appendix 1. Also shown are the average within-municipality similarity % (“Similarity”), the total contributory % of the top 5 dominant species for each municipality (“Top 5”), and the total contributory % of all other species not included in the top 5 (“Others”) (cont’n.) Municipality Code Species Contributory % Family Max TL (cm) Body Size National Patterns of Philippine Reef Fish Diversity 34 2.04 Pomacentrus moluccensis 21.31 Pomacentridae 9.0 small Similarity: 23.1 Pomacentrus lepidogenys 18.72 Pomacentridae 9.0 small Top 5: 68.8 Amblyglyphidodon curacao 13.84 Pomacentridae 11.0 medium Others: 31.2 Pomacentrus bankanensis 9.04 Pomacentridae 9.0 small Plectroglyphidodon lacrymatus 5.85 Pomacentridae 10.0 small 2.05 Chromis atripectoralis 66.22 Pomacentridae 12.0 medium Similarity: 55.5 Pomacentrus lepidogenys 6.48 Pomacentridae 9.0 small Top 5: 87.9 Pomacentrus moluccensis 5.68 Pomacentridae 9.0 small Others: 12.1 Amblyglyphidodon curacao 5.40 Pomacentridae 11.0 medium Neoglyphidodon nigroris 4.08 Pomacentridae 13.0 medium 2.06 Chrysiptera rex 25.25 Pomacentridae 7.0 small Similarity: 36.9 Pomacentrus lepidogenys 24.04 Pomacentridae 9.0 small Top 5: 78.3 Pomacentrus bankanensis 12.58 Pomacentridae 9.0 small Others: 21.7 Thalassoma hardwicke 8.28 Labridae 20.0 medium Ctenochaetus striatus 8.15 Acanthuridae 26.0 medium 2.07 Pomacentrus simsiang 25.64 Pomacentridae 7.0 small Similarity: 11.3 Thalassoma hardwicke 13.85 Labridae 20.0 medium Top 5: 68.1 Pomacentrus opisthostigma 10.85 Pomacentridae 6.5 small Others: 31.9 Labrichthys unilineatus 10.04 Labridae 17.5 medium Neoglyphidodon nigroris 7.72 Pomacentridae 13.0 medium 2.08 Scarus rivulatus 23.48 Scaridae 40.0 large Similarity: 19.5 Pomacentrus alexanderae 14.93 Pomacentridae 9.0 small Top 5: 67.0 Pomacentrus moluccensis 12.52 Pomacentridae 9.0 small Others: 33.0 Amblyglyphidodon curacao 11.23 Pomacentridae 11.0 medium Pomacentrus simsiang 4.87 Pomacentridae 7.0 small 2.09 Pomacentrus chrysurus 71.09 Pomacentridae 9.0 small Similarity: 42.0 Chrysiptera rex 9.73 Pomacentridae 7.0 small Top 5: 89.2 Thalassoma hardwicke 3.11 Pomacentridae 20.0 medium Others: 10.8 Scolopsis lineatus 2.72 Nemipteridae 23.0 medium Naso unicornis 2.59 Acanthuridae 70.0 large Appendix 3A Table showing the top 5 dominant species in terms of abundance, based on Bray-Curtis SIMPER analysis within municipalities, for each municipality, arranged from north to south of the Philippines. Names of each municipality and biogeographic region can be found in Appendix 1. Also shown are the average within-municipality similarity % (“Similarity”), the total contributory % of the top 5 dominant species for each municipality (“Top 5”), and the total contributory % of all other species not included in the top 5 (“Others”) (cont’n.) Municipality Code Species Contributory % Family Max TL (cm) Body Size JA Anticamara and others 35 2.10 Pomacentrus chrysurus 40.35 Pomacentridae 9.0 small Similarity: 16.9 Scarus rivulatus 11.85 Scaridae 40.0 large Top 5: 77.1 Chrysiptera rex 9.11 Pomacentridae 7.0 small Others: 22.9 Pomacentrus moluccensis 8.20 Pomacentridae 9.0 small Pomacentrus alexanderae 7.58 Pomacentridae 9.0 small 2.11 Chromis ternatensis 34.07 Pomacentridae 10.0 small Similarity: 24.7 Chrysiptera cyanea 17.41 Pomacentridae 8.5 small Top 5: 77.7 Pomacentrus burroughi 9.89 Pomacentridae 8.5 small Others: 22.3 Pomacentrus alexanderae 8.93 Pomacentridae 9.0 small Hemiglyphidodon plagiometopon 7.39 Pomacentridae 18.0 medium 2.12 Pomacentrus moluccensis 16.84 Pomacentridae 9.0 small Similarity: 28.8 Scarus rivulatus 16.61 Scaridae 40.0 large Top 5: 62.5 Amblyglyphidodon curacao 14.49 Pomacentridae 11.0 medium Others: 37.5 Chromis ternatensis 7.75 Pomacentridae 10.0 small Dischistodus prosopotaenia 6.80 Pomacentridae 17.0 medium 2.13 Pomacentrus coelestis 15.08 Pomacentridae 9.0 small Similarity: 17.4 Scarus rivulatus 11.61 Scaridae 40.0 large Top 5: 53.1 Pomacentrus chrysurus 11.03 Pomacentridae 9.0 small Others: 46.9 Pomacentrus moluccensis 8.29 Pomacentridae 9.0 small Pomacentrus burroughi 7.07 Pomacentridae 8.5 small 3.01 Pseudanthias huchti 22.52 Serranidae 12.0 medium Similarity: 19.1 Pomacentrus moluccensis 21.86 Pomacentridae 9.0 small Top 5: 59.6 Pomacentrus brachialis 6.05 Pomacentridae 8.0 small Others: 40.4 Chromis viridis 4.94 Pomacentridae 8.0 small Centropyge vroliki 4.26 Pomacentridae 12.0 medium 3.02 Acanthochromis polyacanthus 39.84 Acanthuridae 14.0 medium Similarity: 23.0 Pomacentrus moluccensis 14.6 Pomacentridae 9.0 small Top 5: 67.5 Chaetodon kleinii 4.71 Chaetodontidae 15.0 medium Others: 32.5 Amblyglyphidodon curacao 4.63 Pomacentridae 11.0 medium Chaetodon lunulatus 3.71 Chaetodontidae 14.0 medium Appendix 3A Table showing the top 5 dominant species in terms of abundance, based on Bray-Curtis SIMPER analysis within municipalities, for each municipality, arranged from north to south of the Philippines. Names of each municipality and biogeographic region can be found in Appendix 1. Also shown are the average within-municipality similarity % (“Similarity”), the total contributory % of the top 5 dominant species for each municipality (“Top 5”), and the total contributory % of all other species not included in the top 5 (“Others”) (cont’n.) Municipality Code Species Contributory % Family Max TL (cm) Body Size National Patterns of Philippine Reef Fish Diversity 36 3.03 Centropyge vroliki 30.32 Pomacanthidae 12.0 medium Similarity: 20.3 Pomacentrus bankanensis 26.1 Pomacentridae 9.0 small Top 5: 91.4 Thalassoma hardwicke 25.7 Labridae 20.0 medium Others: 8.6 Plectroglyphidodon lacrymatus 5.24 Pomacentridae 10.0 small Pomacentrus coelestis 4.06 Pomacentridae 9.0 small 3.04 Ctenochaetus striatus 18.01 Acanthuridae 26.0 medium Similarity: 21.0 Pomacentrus moluccensis 13.27 Pomacentridae 9.0 small Top 5: 53.3 Dascyllus trimaculatus 9.85 Pomacentridae 11.0 medium Others: 46.7 Centropyge vroliki 6.36 Pomacanthidae 12.0 medium Pomacentrus lepidogenys 5.76 Pomacentridae 9.0 small 3.05 Pomacentrus moluccensis 55.51 Pomacentridae 9.0 small Similarity: 18.1 Pomacentrus bankanensis 9.45 Pomacentridae 9.0 small Top 5: 80.6 Pomacentrus brachialis 5.30 Pomacentridae 8.0 small Others: 19.4 Abudefduf vaigiensis 5.19 Pomacentridae 20.0 medium Pomacentrus chrysurus 5.12 Pomacentridae 9.0 small 3.06 Thalassoma lunare 28.15 Labridae 25.0 medium Similarity: 46.8 Pomacentrus chrysurus 18.45 Pomacentridae 9.0 small Top 5: 78.2 Scarus rivulatus 15.27 Scaridae 40.0 large Others: 21.8 Pomacentrus simsiang 11.61 Pomacentridae 7.0 small Dascyllus trimaculatus 4.77 Pomacentridae 11.0 medium 3.07 Pomacentrus moluccensis 43.24 Pomacentridae 9.0 small Similarity: 32.6 Pomacentrus chrysurus 14.21 Pomacentridae 9.0 small Top 5: 76.1 Amblygliphidodon ternatensis 6.82 Pomacentridae 10.0 small Others: 23.9 Neoglyphidodon melas 6.68 Pomacentridae 18.0 medium Amblyglyphidodon curacao 5.18 Pomacentridae 11.0 medium 3.08 Chaetodon kleinii 54.25 Chaetodontidae 15.0 medium Similarity: 26.2 Plectroglyphidodon lacrymatus 20.13 Pomacentridae 10.0 small Top 5: 94.2 Dascyllus trimaculatus 13.29 Pomacentridae 11.0 medium Others: 5.8 Scarus rivulatus 4.51 Scaridae 40.0 large Dascyllus reticulatus 2.04 Pomacentridae 9.0 small Append ix 3A Table showing the top 5 dominant species in terms of abundance, based on Bray-Curtis SIMPER analysis within municipalities, for each municipality, arranged from north to south of the Philippines. Names of each municipality and biogeographic region can be found in Appendix 1. Also shown are the average within-municipality similarity % (“Similarity”), the total contributory % of the top 5 dominant species for each municipality (“Top 5”), and the total contributory % of all other species not included in the top 5 (“Others”) (cont’n.) Municipality Code Species Contributory % Family Max TL (cm) Body Size JA Anticamara and others 37 3.09 Pomacentrus coelestis 58.18 Pomacentridae 9.0 small Similarity: 19.4 Pomacentrus bankanensis 12.49 Pomacentridae 9.0 small Top 5: 87.0 Chrysiptera cyanea 9.24 Pomacentridae 8.5 small Others: 13.0 Centropyge vroliki 4.20 Pomacanthidae 12.0 medium Ctenochaetus striatus 2.94 Acanthuridae 26.0 medium 3.10 Plectroglyphidodon lacrymatus 27.83 Pomacentridae 10.0 small Similarity: 27.2 Pomacentrus moluccensis 24.62 Pomacentridae 9.0 small Top 5: 75.2 Amblyglyphidodon curacao 13.03 Pomacentridae 11.0 medium Others: 24.8 Zebrasoma scopas 4.99 Zanclidae 20.0 medium Ctenochaetus striatus 4.77 Acanthuridae 26.0 medium 3.11 Thalassoma lunare 30.08 Labridae 25.0 medium Similarity: 26.0 Pomacentrus chrysurus 19.54 Pomacentridae 9.0 small Top 5: 75.5 Pomacentrus simsiang 12.87 Pomacentridae 7.0 small Others: 24.5 Plectroglyphidodon lacrymatus 6.94 Pomacentridae 10.0 small Chromis ternatensis 6.07 Pomacentridae 10.0 small 3.12 Pomacentrus moluccensis 34.66 Pomacentridae 9.0 small Similarity: 35.0 Pomacentrus burroughi 24.03 Pomacentridae 8.5 small Top 5: 74.3 Chromis ternatensis 5.36 Pomacentridae 10.0 small Others: 25.7 Sphaeramia nematoptera 5.23 Apogonidae 8.5 small Chaetodon octofasciatus 5.03 Chaetodontidae 12.0 medium 3.13 Dascyllus aruanus 41.73 Pomacentridae 10.0 small Similarity: 14.5 Pomacentrus moluccensis 28.78 Pomacentridae 9.0 small Top 5: 82.8 Pomacentrus alexanderae 5.76 Pomacentridae 9.0 small Others: 17.2 Amphiprion clarkii 3.60 Pomacentridae 15.0 medium Amblyglyphidodon curacao 2.88 Pomacentridae 11.0 medium 3.14 Pomacentrus moluccensis 34.64 Pomacentridae 9.0 small Similarity: 19.6 Pseudanthias tuka 14.82 Serranidae 12.0 medium Top 5: 83.2 Pseudanthias huchti 13.63 Serranidae 12.0 medium Others: 16.8 Caesio caerulaurea 12.67 Caesionidae 35.0 large Pomacentrus alexanderae 7.47 Pomacentridae 9.0 small Append ix 3A Table showing the top 5 dominant species in terms of abundance, based on Bray-Curtis SIMPER analysis within municipalities, for each municipality, arranged from north to south of the Philippines. Names of each municipality and biogeographic region can be found in Appendix 1. Also shown are the average within-municipality similarity % (“Similarity”), the total contributory % of the top 5 dominant species for each municipality (“Top 5”), and the total contributory % of all other species not included in the top 5 (“Others”) (cont’n.) Municipality Code Species Contributory % Family Max TL (cm) Body Size National Patterns of Philippine Reef Fish Diversity 38 Appendix 3A Table showing the top 5 dominant species in terms of abundance, based on Bray-Curtis SIMPER analysis within municipalities, for each municipality, arranged from north to south of the Philippines. Names of each municipality and biogeographic region can be found in Appendix 1. Also shown are the average within-municipality similarity % (“Similarity”), the total contributory % of the top 5 dominant species for each municipality (“Top 5”), and the total contributory % of all other species not included in the top 5 (“Others”) (cont’n.) Municipality Code Species Contributory % Family Max TL (cm) Body Size 3.15 Scarus rivulatus 29.36 Scaridae 40.0 large Similarity: 33.2 Chromis weberi 17.67 Pomacentridae 13.5 medium Top 5: 68.9 Dascyllus trimaculatus 11.68 Pomacentridae 11.0 medium Others: 31.1 Plectroglyphidodon lacrymatus 5.20 Pomacentridae 10.0 small Centropyge vroliki 5.03 Pomacanthidae 12.0 medium 3.16 Pomacentrus moluccensis 42.01 Pomacentridae 9.0 small Similarity: 33.9 Caesio caerulaurea 23.14 Caesionidae 35.0 large Top 5: 90.7 Amblyglyphidodon curacao 22.6 Pomacentridae 11.0 medium Others: 9.3 Pomacentrus brachialis 1.83 Pomacentridae 8.0 small Neoglyphidodon nigroris 1.14 Pomacentridae 13.0 medium 4.01 Abudefduf vaigiensis 14.50 Pomacentridae 20.0 medium Similarity: 25.7 Plectroglyphidodon lacrymatus 14.04 Pomacentridae 10.0 small Top 5: 61.4 Ctenochaetus striatus 12.55 Acanthuridae 26.0 medium Others: 38.6 Pomacentrus vaiuli 10.93 Pomacentridae 10.0 small Thalassoma hardwicke 9.34 Labridae 20.0 medium 4.02 Plectroglyphidodon lacrymatus 37.35 Pomacentridae 10.0 small Similarity: 14.12 Halichoeres hortulanus 8.30 Labridae 27.0 medium Top 5: 62.7 Pomacentrus moluccensis 6.69 Pomacentridae 9.0 small Others: 37.3 Pomacentrus coelestis 5.62 Pomacentridae 9.0 small Thalassoma lunare 4.71 Labridae 25.0 medium 4.03 Pomacentrus simsiang 18.08 Pomacentridae 7.0 small Similarity: 11.3 Plectroglyphidodon lacrymatus 8.57 Pomacentridae 10.0 small Top 5: 49.5 Dascyllus reticulatus 8.2 Pomacentridae 9.0 small Others: 50.5 Dischistodus prosopotaenia 7.62 Pomacentridae 17.0 medium Apogon griffini 7.0 Apogonidae 13.5 medium 4.04 Pomacentrus moluccensis 26.72 Pomacentridae 9.0 small Similarity: 34.3 Pomacentrus adelus 21.94 Pomacentridae 8.5 small Top 5: 71.5 Plectroglyphidodon lacrymatus 14.4 Pomacentridae 10.0 small Others: 28.5 Thalassoma hardwicke 4.22 Labridae 20.0 medium Amblyglyphidodon curacao 4.2 Pomacentridae 11.0 medium JA Anticamara and others 39 Appendix 3A Table showing the top 5 dominant species in terms of abundance, based on Bray-Curtis SIMPER analysis within municipalities, for each municipality, arranged from north to south of the Philippines. Names of each municipality and biogeographic region can be found in Appendix 1. Also shown are the average within-municipality similarity % (“Similarity”), the total contributory % of the top 5 dominant species for each municipality (“Top 5”), and the total contributory % of all other species not included in the top 5 (“Others”) (cont’n.) Municipality Code Species Contributory % Family Max TL (cm) Body Size 4.05 Pomacentrus moluccensis 26.79 Pomacentridae 9.0 small Similarity: 20.2 Chromis margaritifer 9.98 Pomacentridae 9.0 small Top 5: 62.8 Pomacentrus simsiang 9.74 Pomacentridae 7.0 small Others: 37.2 Dascyllus reticulatus 8.21 Pomacentridae 9.0 small Ctenochaetus striatus 8.06 Acanthuridae 26.0 medium 4.06 Cirrhilabrus cyanopleura 53.02 Labridae 15.0 medium Similarity: 22.7 Pomacentrus lepidogenys 11.76 Pomacentridae 9.0 small Top 5: 83.9 Scolopsis bilineatus 7.79 Nemipteridae 23.0 medium Others: 16.1 Ctenochaetus striatus 6.7 Acanthuridae 26.0 medium Thalassoma lunare 4.65 Labridae 25.0 medium 5.01 Pomacentrus moluccensis 52.49 Pomacentridae 9.0 small Similarity: 27.0 Pomacentrus chrysurus 10.42 Pomacentridae 9.0 small Top 5: 86.4 Amblyglyphidodon curacao 10.34 Pomacentridae 11.0 medium Others: 13.6 Chromis viridis 8.1 Pomacentridae 8.0 small Neoglyphidodon nigroris 5.02 Pomacentridae 13.0 medium 5.02 Acanthochromis polyacanthus 40.12 Pomacentridae 14.0 medium Similarity: 29.9 Pomacentrus moluccensis 18.8 Pomacentridae 9.0 small Top 5: 74.3 Pomacentrus lepidogenys 7.24 Pomacentridae 9.0 small Others: 25.7 Plectroglyphidodon lacrymatus 4.17 Pomacentridae 10.0 small Ctenochaetus striatus 4 Acanthuridae 26.0 medium 6.01 Thalassoma lunare 24.91 Labridae 25.0 medium Similarity: 29.2 Scarus rivulatus 20.96 Scaridae 40.0 large Top 5: 78.5 Chlorurus sordidus 17.74 Pomacentridae 40.0 large Others: 21.5 Halichoeres melanurus 7.55 Labridae 12.0 medium Chaetodon octofasciatus 7.37 Chaetodontidae 12.0 medium 6.02 Dascyllus reticulatus 20.63 Pomacentridae 9.0 small Similarity: 24.7 Plectroglyphidodon lacrymatus 17.27 Pomacentridae 10.0 small Top 5: 67.1 Ctenochaetus striatus 12.58 Acanthuridae 26.0 medium Others: 32.9 Plectroglyphidodon dickii 9.35 Pomacentridae 11.0 medium Pomacentrus moluccensis 7.25 Pomacentridae 9.0 small 6.03 Pomacentrus lepidogenys 19.13 Pomacentridae 9.0 small Similarity: 34.3 Acanthochromis polyacanthus 15.01 Pomacentridae 14.0 medium Top 5: 65.2 Ctenochaetus striatus 12.12 Acanthuridae 26.0 medium Others: 34.8 Plectroglyphidodon lacrymatus 11.63 Pomacentridae 10.0 small Centropyge vroliki 7.28 Pomacanthidae 12.0 medium National Patterns of Philippine Reef Fish Diversity 40 1.01 Zanclus cornutus 38.46 Zanclidae 23 medium Similarity:20.7 Chaetodon ornatissimus 25.62 Chaetodontidae 20 medium Top 5: 94.5 Chaetodon kleinii 25.52 Chaetodontidae 15 medium Others: 5.5 Sufflamen chrysopterus 2.79 Balistidae 30 medium Centropyge vroliki 2.14 Pomacanthidae 12 medium 1.02 Ctenochaetus striatus 28.29 Acanthuridae 26 medium Similarity: 37.4 Chlorurus sordidus 20.20 Scaridae 40 large Top 5: 85.4 Cheilinus chlorourus 13.92 Labridae 45 large Others: 14.6 Zanclus cornutus 11.70 Zanclidae 23 medium Chaetodon kleinii 11.31 Chaetodontidae 15 medium 1.03 Halichoeres hortulanus 28.41 Labridae 27 medium Similarity: 17.9 Ctenochaetus striatus 23.04 Acanthuridae 26 medium Top 5: 75.4 Parupeneus multifasciatus 10.23 Mullidae 35 large Others: 24.6 Thalassoma hardwicke 8.50 Labridae 20 medium Chlorurus sordidus 5.18 Scaridae 40 large 1.04 Epinephelus merra 16.80 Serranidae 31 large Similarity: 28.6 Thalassoma lunare 14.72 Labridae 25 medium Top 5: 59.9 Plectroglyphidodon lacrymatus 10.28 Pomacentridae 10 small Others: 40.1 Halichoeres melanurus 9.33 Labridae 12 medium Labracinus cyclophthalmus 8.72 Pseudochromidae 20 medium 1.05 Ctenochaetus striatus 34.43 Acanthuridae 26 medium Similarity: 12.2 Thalassoma hardwicke 24.79 Labridae 20 medium Top 5: 72.8 Plectroglyphidodon lacrymatus 6.04 Pomacentridae 10 small Others: 27.2 Thalassoma lunare 4.09 Labridae 25 medium Lutjanus decussatus 3.45 Lutjanidae 35 large 1.06 Dischistodus prosopotaenia 23.73 Pomacentridae 17 medium Similarity: 9.6 Neoglyphidodon melas 21.73 Pomacentridae 18 medium Top 5: 69.2 Dascyllus trimaculatus 10.46 Pomacentridae 11 medium Others: 30.8 Choerodon anchorago 7.41 Labridae 38 large Plectroglyphidodon lacrymatus 5.86 Pomacentridae 10 small Appendix 3B Table showing the top 5 dominant species in terms of biomass, based on Bray-Curtis SIMPER analysis within municipalities, for each municipality, arranged from north to south of the Philippines. Names of each municipality and biogeographic region can be found in Appendix 1. Also shown are the average within-municipality similarity % (“Similarity”), the total contributory % of the top 5 dominant species for each municipality (“Top 5”), and the total contributory % of all other species not included in the top 5 (“Others”) Municipality Code Species Contributory % Family Max TL (cm) Body Size JA Anticamara and others 41 1.07 Ctenochaetus striatus 38.67 Acanthuridae 26 medium Similarity: 13.3 Thalassoma hardwicke 11.74 Labridae 20 medium Top 5: 72.3 Balistapus undulatus 10.28 Balistidae 30 medium Others: 27.7 Epinephelus merra 5.94 Serranidae 31 large Stegastes fasciolatus 5.69 Pomacentridae 15 medium 1.08 Thalassoma lunare 19.78 Labridae 25 medium Similarity: 17.5 Ctenochaetus striatus 19.51 Acanthuridae 26 medium Top 5: 70.5 Scolopsis margaritifer 12.77 Nemipteridae 28 medium Others: 29.5 Arothron nigropunctatus 9.73 Tetraodontidae 33 large Thalassoma hardwicke 8.75 Labridae 20 medium 1.09 Lutjanus decussatus 10.22 Lutjanidae 35 large Similarity: 9.2 Dischistodus prosopotaenia 9.35 Pomacentridae 17 medium Top 5: 44.1 Plectroglyphidodon lacrymatus 8.86 Pomacentridae 10 small Others: 55.9 Cheilinus chlorourus 8.18 Labridae 45 large Neoglyphidodon nigroris 7.49 Pomacentridae 13 medium 2.01 Ctenochaetus striatus 42.90 Acanthuridae 26 medium Similarity: 17.5 Zanclus cornutus 13.85 Zanclidae 23 medium Top 5: 71.7 Chaetodon vagabundus 7.50 Chaetodontidae 23 medium Others: 28.3 Halichoeres hortulanus 4.40 Labridae 27 medium Thalassoma hardwicke 3.07 Labridae 20 medium 2.02 Ctenochaetus striatus 46.36 Acanthuridae 26 medium Similarity: 23.0 Chlorurus sordidus 13.45 Scaridae 40 large Top 5: 76.7 Parupeneus multifasciatus 5.99 Mullidae 35 large Others: 23.3 Hemigymnus fasciatus 5.61 Labridae 80 large Plectroglyphidodon lacrymatus 5.31 Pomacentridae 10 small 2.03 Chlorurus sordidus 24.39 Scaridae 40 large Similarity: 16.5 Scarus flavipectoralis 23.29 Scaridae 40 large Top 5: 70.1 Lutjanus decussatus 10.55 Lutjanidae 35 large Others: 29.9 Ctenochaetus striatus 6.29 Chaetodontidae 26 medium Zanclus cornutus 5.58 Zanclidae 23 medium Append ix 3B Table showing the top 5 dominant species in terms of biomass, based on Bray-Curtis SIMPER analysis within municipalities, for each municipality, arranged from north to south of the Philippines. Names of each municipality and biogeographic region can be found in Appendix 1. Also shown are the average within-municipality similarity % (“Similarity”), the total contributory % of the top 5 dominant species for each municipality (“Top 5”), and the total contributory % of all other species not included in the top 5 (“Others”) (cont’n.) Municipality Code Species Contributory % Family Max TL (cm) Body Size National Patterns of Philippine Reef Fish Diversity 42 2.04 Lutjanus decussatus 15.06 Lutjanidae 35 large Similarity: 17.0 Zanclus cornutus 13.37 Zanclidae 23 medium Top 5: 57.5 Ctenochaetus striatus 12.83 Acanthuridae 26 medium Others: 42.5 Halichoeres hortulanus 10.30 Labridae 27 medium Scolopsis bilineatus 5.94 Nemipteridae 23 medium 2.05 Zanclus cornutus 17.01 Zanclidae 23 medium Similarity: 25.8 Chaetodon lunulatus 7.48 Chaetodontidae 14 medium Top 5: 42.6 Hemigymnus fasciatus 6.71 Labridae 80 large Others: 57.4 Chaetodontoplus mesoleucus 6.09 Pomacanthidae 18 medium Labrichthys unilineatus 5.27 Labridae 17.5 medium 2.06 Ctenochaetus striatus 38.67 Acanthuridae 26 medium Similarity: 18.7 Thalassoma hardwicke 13.10 Labridae 20 medium Top 5: 77.6 Chlorurus sordidus 11.99 Scaridae 40 large Others: 22.4 Lutjanus decussatus 8.58 Lutjanidae 35 large Chaetodon citrinellus 5.27 Chaetodontidae 13 medium 2.07 Choerodon anchorago 31.69 Labridae 38 large Similarity: 12.3 Thalassoma hardwicke 24.77 Labridae 20 medium Top 5: 80.0 Pomacentrus simsiang 10.24 Pomacentridae 7 small Others: 20.0 Siganus unimaculatus 7.52 Siganidae 20 medium Amblyglyphidodon curacao 5.33 Pomacentridae 11 medium 2.08 Scarus rivulatus 24.38 Scaridae 40 large Similarity: 17.6 Chlorurus sordidus 20.28 Scaridae 40 large Top 5: 63.3 Lutjanus decussatus 10.05 Lutjanidae 35 large Others: 36.7 Hemigymnus melapterus 4.73 Labridae 90 large Chaetodon octofasciatus 3.89 Chaetodontidae 12 medium 2.09 Thalassoma hardwicke 24.86 Labridae 20 medium Similarity: 12.3 Lutjanus decussatus 12.78 Lutjanidae 35 large Top 5: 68.1 Cheilinus chlorourus 10.75 Lutjanidae 45 large Others: 31.9 Pomacentrus chrysurus 10.59 Pomacentridae 9 small Scolopsis lineatus 9.14 Nemipteridae 23 medium Appendix 3B Table showing the top 5 dominant species in terms of biomass, based on Bray-Curtis SIMPER analysis within municipalities, for each municipality, arranged from north to south of the Philippines. Names of each municipality and biogeographic region can be found in Appendix 1. Also shown are the average within-municipality similarity % (“Similarity”), the total contributory % of the top 5 dominant species for each municipality (“Top 5”), and the total contributory % of all other species not included in the top 5 (“Others”) (cont’n.) Municipality Code Species Contributory % Family Max TL (cm) Body Size JA Anticamara and others 43 2.10 Scarus rivulatus 24.85 Scaridae 40 large Similarity: 15.6 Choerodon anchorago 18.49 Labridae 38 large Top 5: 67.3 Thalassoma hardwicke 13.17 Labridae 20 medium Others: 32.7 Halichoeres melanurus 6.91 Labridae 12 medium Coris batuensis 3.88 Labridae 17 medium 2.11 Hemiglyphidodon plagiometopon27.14 Labridae 18 medium Similarity: 20.4 Chaetodontoplus mesoleucus 15.78 Pomacanthidae 18 medium Top 5: 72.5 Scarus rivulatus 14.19 Scaridae 40 large Others: 27.5 Hemigymnus melapterus 8.20 Labridae 90 large Chaetodon octofasciatus 7.15 Chaetodontidae 12 medium 2.12 Scarus rivulatus 27.12 Scaridae 40 large Similarity: 28.4 Dischistodus prosopotaenia 22.69 Pomacentridae 17 medium Top 5: 70.9 Lutjanus decussatus 8.91 Lutjanidae 35 large Others: 29.1 Halichoeres chloropterus 6.90 Labridae 19 medium Choerodon anchorago 5.24 Labridae 38 large 2.13 Scarus rivulatus 32.13 Scaridae 40 large Similarity: 15.2 Hemigymnus melapterus 10.07 Labridae 90 large Top 5: 61.5 Choerodon anchorago 8.92 Labridae 38 large Others: 38.5 Scolopsis bilineatus 5.19 Nemipteridae 23 medium Hemiglyphidodon plagiometopon 5.17 Labridae 18 medium 3.01 Thalassoma lunare 9.07 Labridae 25 medium Similarity: 13.2 Chaetodon baronessa 7.44 Chaetodontidae 16 medium Top 5: 63.6 Zebrasoma scopas 7.17 Zanclidae 20 medium Others: 36.4 Pomacentrus moluccensis 6.40 Pomacentridae 9 small Chaetodon kleinii 6.31 Chaetodontidae 15 medium 3.02 Chaetodon lunulatus 22.15 Chaetodontidae 14 medium Similarity: 19.8 Chaetodon baronessa 8.29 Chaetodontidae 16 medium Top 5: 50.6 Ctenochaetus striatus 7.24 Acanthuridae 26 medium Others: 49.4 Halichoeres hortulanus 6.84 Labridae 27 medium Thalassoma lunare 6.03 Labridae 25 medium Appendix 3B Table showing the top 5 dominant species in terms of biomass, based on Bray-Curtis SIMPER analysis within municipalities, for each municipality, arranged from north to south of the Philippines. Names of each municipality and biogeographic region can be found in Appendix 1. Also shown are the average within-municipality similarity % (“Similarity”), the total contributory % of the top 5 dominant species for each municipality (“Top 5”), and the total contributory % of all other species not included in the top 5 (“Others”) (cont’n.) Municipality Code Species Contributory % Family Max TL (cm) Body Size National Patterns of Philippine Reef Fish Diversity 44 3.03 Thalassoma hardwicke 54.44 Labridae 20 medium Similarity: 11.7 Centropyge vroliki 17.48 Pomacanthidae 12 medium Top 5: 94.0 Pomacentrus bankanensis 11.98 Pomacentridae 9 small Others: 6.0 Halichoeres hortulanus 8.18 Labridae 27 medium Bodianus mesothorax 1.94 Labridae 25 medium 3.04 Ctenochaetus striatus 17.56 Chaetodontidae 26 medium Similarity: 18.7 Thalassoma lunare 12.39 Lutjanidae 25 medium Top 5: 55.2 Chaetodon vagabundus 10.3 Chaetodontidae 23 medium Others: 45.8 Parupeneus multifasciatus 8.37 Mullidae 35 large Zebrasoma scopas 6.60 Acanthuridae 20 medium 3.05 Thalassoma lunare 14.11 Scaridae 25 medium Similarity: 9.9 Chaetodon baronessa 8.90 Chaetodontidae 16 medium Top 5: 46.9 Plectorhinchus vittatus 8.56 Haemulidae 72 large Others: 53.1 Halichoeres melanurus 8.48 Labridae 12 medium Neoglyphidodon nigroris 6.84 Pomacentridae 13 medium 3.06 Thalassoma lunare 53.96 Labridae 25 medium Similarity: 40.6 Scolopsis bilineatus 18.61 Nemipteridae 23 medium Top 5: 85.9 Cephalopholis boenak 5.70 Serranidae 30 medium Others: 14.1 Halichoeres melanurus 3.91 Labridae 12 medium Scarus rivulatus 3.70 Scaridae 40 large 3.07 Labracinus cyclophthalmus 25.50 Pseudochromidae 20 medium Similarity: 19.9 Thalassoma hardwicke 8.02 Labridae 20 medium Top 5: 53.7 Chaetodontoplus mesoleucus 7.95 Chaetodontidae 18 medium Others: 46.3 Halichoeres chloropterus 6.59 Labridae 19 medium Thalassoma lunare 5.64 Labridae 25 medium 3.08 Chaetodon kleinii 21.83 Chaetodontidae 15 medium Similarity: 11.3 Plectroglyphidodon lacrymatus 13.36 Chaetodontidae 10 small Top 5: 64.3 Dascyllus trimaculatus 11.14 Chaetodontidae 11 medium Others: 35.7 Scarus rivulatus 10.27 Scaridae 40 large Dascyllus reticulatus 7.73 Chaetodontidae 9 small Appendix 3B Table showing the top 5 dominant species in terms of biomass, based on Bray-Curtis SIMPER analysis within municipalities, for each municipality, arranged from north to south of the Philippines. Names of each municipality and biogeographic region can be found in Appendix 1. Also shown are the average within-municipality similarity % (“Similarity”), the total contributory % of the top 5 dominant species for each municipality (“Top 5”), and the total contributory % of all other species not included in the top 5 (“Others”) (cont’n.) Municipality Code Species Contributory % Family Max TL (cm) Body Size JA Anticamara and others 45 3.09 Centropyge vroliki 17.0 Pomacanthidae 12 medium Similarity: 11.2 Thalassoma lunare 13.44 Labridae 25 medium Top 5: 63.7 Chaetodon kleinii 12.92 Chaetodontidae 15 medium Others: 36.3 Dascyllus trimaculatus 10.52 Pomacentridae 11 medium Pomacentrus bankanensis 9.84 Pomacentridae 9 small 3.10 Balistapus undulatus 30.72 Balistidae 30 medium Similarity: 25.5 Ctenochaetus striatus 11.77 Chaetodontidae 26 medium Top 5: 67.5 Chaetodon baronessa 9.09 Chaetodontidae 16 medium Others: 32.5 Chaetodon lunulatus 8.32 Chaetodontidae 14 medium Zebrasoma scopas 7.59 Acanthuridae 20 medium 3.11 Thalassoma lunare 40.76 Labridae 25 medium Similarity: 16.8 Halichoeres chloropterus 13.99 Labridae 19 medium Top 5: 85.3 Halichoeres melanurus 13.64 Labridae 12 medium Others: 14.7 Pomacentrus chrysurus 8.68 Pomacentridae 9 small Pomacentrus simsiang 8.23 Pomacentridae 7 small 3.12 Neoglyphidodon melas 22.17 Pomacentridae 18 medium Similarity: 19.1 Chlorurus sordidus 14.38 Scaridae 40 large Top 5: 57.4 Scarus quoyi 8.25 Scaridae 40 large Others: 42.6 Scarus dimidiatus 6.36 Scaridae 40 large Chaetodontoplus mesoleucus 6.21 Pomacanthidae 18 medium 3.13 Thalassoma lunare 29.49 Labridae 25 medium Similarity: 43.1 Parupeneus multifasciatus 10.86 Mullidae 35 large Top 5: 66.7 Scolopsis bilineatus 9.96 Nemipteridae 23 medium Others: 33.3 Chaetodon baronessa 8.56 Chaetodontidae 16 medium Centropyge vroliki 7.81 Pomacanthidae 12 medium 3.14 Thalassoma hardwicke 17.82 Labridae 20 medium Similarity: 13.4 Scarus niger 9.63 Scaridae 40 large Top 5: 48.7 Acanthurus lineatus 7.72 Acanthuridae 38 large Others: 51.3 Chlorurus sordidus 7.51 Scaridae 40 large Melichthys vidua 6.06 Balistidae 40 large Appendix 3B Table showing the top 5 dominant species in terms of biomass, based on Bray-Curtis SIMPER analysis within municipalities, for each municipality, arranged from north to south of the Philippines. Names of each municipality and biogeographic region can be found in Appendix 1. Also shown are the average within-municipality similarity % (“Similarity”), the total contributory % of the top 5 dominant species for each municipality (“Top 5”), and the total contributory % of all other species not included in the top 5 (“Others”) (cont’n.) Municipality Code Species Contributory % Family Max TL (cm) Body Size National Patterns of Philippine Reef Fish Diversity 46 Append ix 3B Table showing the top 5 dominant species in terms of biomass, based on Bray-Curtis SIMPER analysis within municipalities, for each municipality, arranged from north to south of the Philippines. Names of each municipality and biogeographic region can be found in Appendix 1. Also shown are the average within-municipality similarity % (“Similarity”), the total contributory % of the top 5 dominant species for each municipality (“Top 5”), and the total contributory % of all other species not included in the top 5 (“Others”) (cont’n.) Municipality Code Species Contributory % Family Max TL (cm) Body Size 3.15 Ctenochaetus striatus 27.04 Acanthuridae 26 medium Similarity: 24.2 Chaetodon kleinii 13.11 Chaetodontidae 15 medium Top 5: 63.5 Thalassoma lunare 10.32 Labridae 25 medium Others: 36.5 Dascyllus trimaculatus 6.74 Pomacentridae 11 medium Scarus rivulatus 6.31 Scaridae 40 large 3.16 Pygoplites diacanthus 20.12 Pomacanthidae 25 medium Similarity: 13.1 Chlorurus bleekeri 13.20 Scaridae 49 large Top 5: 63.5 Chlorurus sordidus 12.29 Scaridae 40 large Others: 36.5 Platax boersii 10.82 Ephippidae 40 large Hemigymnus fasciatus 7.05 Labridae 80 large 4.01 Ctenochaetus striatus 35.59 Acanthuridae 26 medium Similarity: 19.9 Acanthurus lineatus 13.87 Acanthuridae 38 large Top 5: 67.3 Thalassoma hardwicke 10.03 Labridae 20 medium Others: 32.7 Parupeneus multifasciatus 4.61 Mullidae 35 large Chaetodon vagabundus 3.25 Chaetodontidae 23 medium 4.02 Acanthurus lineatus 22.18 Acanthuridae 38 large Similarity:14.6 Ctenochaetus striatus 12.58 Acanthuridae 26 medium Top 5: 61.6 Epinephelus merra 10.70 Serranidae 31 large Others: 38.4 Halichoeres hortulanus 8.09 Labridae 27 medium Chaetodon baronessa 8.02 Chaetodontidae 16 medium 4.03 Plectorhinchus chaetodonoides 12.89 Haemulidae 72 large Similarity:13.0 Hemiglyphidodon plagiometopon 9.65 Pomacentridae 18 medium Top 5: 44.3 Dischistodus prosopotaenia 7.69 Pomacentridae 17 medium Others: 55.7 Acanthurus auranticavus 7.38 Acanthuridae 35 large Pentapodus bifasciatus 6.72 Nemipteridae 18 medium 4.04 Ctenochaetus striatus 15.95 Acanthuridae 26 medium Similarity:18.0 Thalassoma hardwicke 11.24 Labridae 20 medium Top 5: 44.0 Thalassoma lunare 6.69 Labridae 25 medium Others: 56.0 Chlorurus sordidus 5.87 Scaridae 40 large Lutjanus decussatus 4.29 Lutjanidae 35 large JA Anticamara and others 47 Append ix 3B Table showing the top 5 dominant species in terms of biomass, based on Bray-Curtis SIMPER analysis within municipalities, for each municipality, arranged from north to south of the Philippines. Names of each municipality and biogeographic region can be found in Appendix 1. Also shown are the average within-municipality similarity % (“Similarity”), the total contributory % of the top 5 dominant species for each municipality (“Top 5”), and the total contributory % of all other species not included in the top 5 (“Others”) (cont’n.) Municipality Code Species Contributory % Family Max TL (cm) Body Size 4.05 Ctenochaetus striatus 30.01 Acanthuridae 26 medium Similarity:13.1 Chaetodon lunulatus 12.9 Chaetodontidae 14 medium Top 5: 59.1 Zebrasoma scopas 6.64 Acanthuridae 20 medium Others: 40.9 Balistapus undulatus 5.85 Balistidae 30 medium Pomacentrus moluccensis 3.74 Pomacentridae 9 small 4.06 Ctenochaetus striatus 49.75 Acanthuridae 26 medium Similarity:18.2 Scolopsis bilineatus 19.54 Nemipteridae 23 medium Top 5: 89.0 Chaetodon kleinii 8.7 Chaetodontidae 15 medium Others: 11.0 Thalassoma lunare 7.24 Labridae 25 medium Centropyge bicolor 3.72 Pomacanthidae 15 medium 5.01 Pomacentrus chrysurus 17.35 Pomacentridae 9 small Similarity:15.7 Pomacentrus moluccensis 16.27 Pomacentridae 9 small Top 5: 64.9 Thalassoma lunare 11.6 Labridae 25 medium Others: 35.1 Neoglyphidodon nigroris 10.22 Pomacentridae 13 medium Chaetodon octofasciatus 9.49 Chaetodontidae 12 medium 5.02 Balistapus undulatus 15.29 Balistidae 30 medium Similarity:20.9 Ctenochaetus striatus 12.64 Chaetodontidae 26 medium Top 5: 43.7 Thalassoma hardwicke 6.5 Labridae 20 medium Others: 56.3 Parupeneus multifasciatus 4.79 Mullidae 35 large Naso lituratus 4.52 Acanthuridae 46 large 6.01 Chlorurus sordidus 55.2 Scaridae 40 large Similarity:28.7 Thalassoma lunare 22.12 Labridae 25 medium Top 5: 95.6 Ctenochaetus striatus 11.16 Acanthuridae 26 medium Others: 4.4 Cephalopholis argus 5 Serranidae 60 large Chaetodon octofasciatus 2.17 Chaetodontidae 12 medium 6.02 Ctenochaetus striatus 37.91 Acanthuridae 26 medium Similarity:22.3 Balistapus undulatus 6.7 Balistidae 30 medium Top 5: 60.4 Thalassoma hardwicke 5.67 Labridae 20 medium Others: 39.6 Plectroglyphidodon dickii 5.38 Pomacentridae 11 medium Plectroglyphidodon lacrymatus 4.74 Pomacentridae 10 small 6.03 Ctenochaetus striatus 22.99 Acanthuridae 26 medium Similarity: 29.0 Balistapus undulatus 12.09 Balistidae 30 medium Top 5: 56.6 Parupeneus multifasciatus 8.28 Mullidae 35 large Others: 43.4 Zanclus cornutus 7.16 Zanclidae 23 medium Thalassoma hardwicke 6.07 Labridae 20 medium