The Philippines

15
Patrick C. Cabaitan, Timothy Joseph R. Quimpo,
Edwin E. Dumalagan Jr., Jeffrey Munar,
Mary Ann C. Calleja, Ronald Dionnie D. Olavides,
Kevin Go, Ritzelle Albelda, Dominic Cabactulan,
Erin Joy Capdos Tinacba, Ma. Angelique A. Doctor,
Cesar L.Villanoy, and Fernando P. Siringan

Abstract Keywords
The Philippines is situated within the Coral Triangle Mesophotic coral ecosystems · Biodiversity ·
marine biodiversity hotspot and supports highly diverse Macroinvertebrate · Megafauna · Reef fish
coral reef communities. However, Philippine reefs are
exposed to many natural and anthropogenic disturbances.
Consequently, mesophotic coral ecosystems (MCEs) 15.1 Introduction
have drawn increasing interest because of their potential
significance as refugia for many reef species. MCEs in the The Philippines is situated within the Coral Triangle, the epi-
Philippines occur in a variety of settings, reflecting the center of global marine biodiversity (Carpenter and Springer
wide diversity of reef habitats within the archipelago. 2005; Veron et  al. 2009; DeVantier and Turak 2017). The
MCEs remain poorly studied compared to shallow reefs, Philippine islands support a diverse range of different reef
but preliminary investigations show that MCE sites sup- habitats from turbid fringing reefs to oceanic atolls. At least
port diverse ecological communities. Here, we describe 468 identified species of scleractinian corals (Veron and
the physical features, biodiversity, and current condition Fenner 2002; Licuanan and Capili 2003, 2005), 736 reef
of three MCE sites in the Philippines that occur in differ- fishes (Allen 2002; Nañola et al. 2006), 648 mollusks (Wells
ent environmental settings: surge-exposed fringing reefs 2002), 16 seagrasses (Fortes and Santos 2004), and 820
at Patnanungan, turbid fringing reefs at Abra de Ilog, and marine algae (Trono 1999) have been recorded from the
the oceanic atolls of Apo Reef. Patnanungan is dominated Philippines. Shallow-water reefs (SWRs) in the country are
by rubble fields with high macroalgae cover, an indication exposed to both natural and anthropogenic disturbances such
of destructive fishing and overexploitation of herbivorous as destructive fishing and land-based runoff, and the increas-
fishes and invertebrates. Abra de Ilog is exposed to turbid ing frequency of disturbances is leading to declining reef
waters due to two nearby rivers, and as a result, hard cor- health in many areas (White et  al. 2000; Wilkinson 2008;
als are not found below 30 m. Apo Reef, an oceanic atoll Burke et al. 2012).
located within a marine park, supported higher abundance A nationwide assessment of SWRs in the 1990s high-
and diversity of benthic and mobile megafauna at meso- lighted the degraded state of many Philippine reefs (Gomez
photic depths than the other two sites. MCEs are probably et al. 1994), a finding supported by more recent assessments
common throughout the Philippines, and future research of reef fishes (Nañola et al. 2011; Go et al. 2015) and corals
should investigate additional sites, include long-term (Licuanan et  al. 2017). Continuing or accelerating habitat
monitoring of factors driving spatial and temporal pat- loss is increasing the extinction risk of important coral reef
terns of biodiversity, and investigate the potential impor- taxa such as fish (Graham et al. 2011) and corals (Carpenter
tance of Philippine MCEs as refugia. et  al. 2008). However, most coral reef research in the
Philippines has been conducted in relatively shallow waters
less than 10  m deep. The spatial extent, biodiversity, and
P. C. Cabaitan (*) · T. J. R. Quimpo · E. E. Dumalagan Jr. conservation status of mesophotic coral ecosystems (MCEs;
J. Munar · M. A. C. Calleja · R. D. D. Olavides · K. Go
30–150 m deep; Hinderstein et al. 2010) is largely unknown.
R. Albelda · D. Cabactulan · E. J. C. Tinacba · M. A. A. Doctor
C. L. Villanoy · F. P. Siringan Considering the high biodiversity of SWRs in the
The Marine Science Institute, College of Science, University of the Philippines and its location in the Coral Triangle, it has
Philippines, Diliman, Quezon City, Philippines strong potential for harboring significant biodiversity at
e-mail: [email protected]

© Springer Nature Switzerland AG 2019 265

Y. Loya et al. (eds.), Mesophotic Coral Ecosystems, Coral Reefs of the World 12,
https://doi.org/10.1007/978-3-319-92735-0_15
266 P. C. Cabaitan et al.

mesophotic depths. Indeed, in other areas of the Indo-Pacific, Analysis of data collected by GCM is still ongoing, but
MCEs exhibit high regional endemism compared to nearby detailed observations from three sites in different environ-
SWRs (e.g., Kane et al. 2014) and because they are poorly mental contexts are presented below. A brief summary of
documented, often support a high proportion of undescribed information on all 15 MCE sites examined is shown in
species. The recent discovery of at least two new species of Table 15.1.
coral reef fishes from Philippine MCEs (Anderson et  al. Data collected during the GCM includes geomorphologi-
2016; Rocha et al. 2017) highlights their likely biodiversity cal maps of the study sites and oceanographic data surveys of
value. Consequently, increased knowledge of MCEs is fish and benthic communities. Bathymetric maps of the sites
important for understanding biodiversity and conservation were collected using a multibeam echosounder and side-scan
status of coral reefs in the Philippines. sonar. The raw multibeam data were acquired and processed
Herein, we describe the physical features, biodiversity, through HYPACK and HYSWEEP software to produce the
and current condition of three MCE sites in the Philippines final gridded bathymetric datasets. Oceanographic data on
that occur in different environmental settings: surge-exposed spatial flow, temperature and salinity profiles were also col-
fringing reefs at Patnanungan, turbid fringing reefs at Abra lected. The degree of water column stratification can be esti-
de Ilog, and the oceanic atolls of Apo Reef. mated using the temperature and salinity profiles indicating
the magnitude of the vertical variations and the degree of
mixing in the water column. Biodiversity surveys of fish and
15.1.1 Research History benthic communities were limited to the upper mesophotic
zone (30–40 m depth) using SCUBA, while surveys greater
Coral reef research in the Philippines started four decades than 40 m were conducted using remotely deployed videos
ago and included the discovery and identification of a num- (RDV; 30–40  m) and remotely operated vehicles (ROVs;
ber of coral species in the Philippines by Francisco 50–200  m). For benthic diversity surveys, 1 × 1  m photo-
Nemenzo and his associates (Nemenzo 1986). Since then, quadrats were taken every 1 m along a 30 m transect (total
major efforts on coral reef research continued, spearheaded surveyed reef area per transect = 30 m2). To quantify benthic
by two prominent coral reef scientists, Edgardo D. Gomez community composition, a uniformly spaced grid of 25
and Angel C. Alcala (1979). Pioneering studies were con- points was overlaid onto each photoquadrat, and the benthic
ducted and influenced the establishment of marine pro- category that fell under each point was categorized. Fish
tected areas on coral reef fish communities (Alcala 1988). diversity surveys were conducted simultaneously with ben-
Also, one of the first countrywide coral reef assessments in thic surveys and involved a single diver noting counts and
the world was initiated (Gomez and Alcala 1979; Gomez size of all fishes encountered within 5 m on either side of the
et  al. 1994). The status of coral reef ecosystems in the 30  m belt transect (total surveyed reef area per
Philippines were monitored in the succeeding years (Yap transect = 300 m2).
and Gomez 1985; Gomez 1991; Gomez et al. 1994; Nañola
et  al. 2006, 2011; Go et  al. 2015; Licuanan et  al. 2017).
However, all of these studies had focused on SWRs with a 15.2 Environmental Setting
maximum depth of 10 m.
The earliest known study on MCEs in the Philippines The Philippines experiences a tropical climate, with sea-
was conducted by Ross and Hodgson (1981) to quantify sonal changes in temperature, humidity, and rainfall brought
coral diversity and percent cover in a depth range of 3–35 m about by monsoonal variations. The Philippines is exposed
in Apo Reef Natural Park, Philippines. While little is known to monsoonal climate and tides, responding to various phys-
about Philippine MCEs, significant efforts have been ical oceanographic factors (e.g., water and wind circulation)
recently made to address this research gap. The Geophysical from the South China Sea and the West Pacific that make the
Coral Reef Mapping (GCM) project of the Marine Science country prone to tropical cyclones (Gordon et al. 2011). The
Institute of the University of the Philippines, which includes archipelagic nature of the Philippines results mainly from
geological, oceanographic, and biological components, is its active tectonic setting evolving from a complex assem-
the first major MCE research project in the Philippines. blage of accreted island arcs, marginal basins, oceanic
Together with the MCE assessment efforts of other institu- crusts, and continental terraces (Karig 1983; McCabe et al.
tions in the Philippines (Abesamis et  al. 2017; Nacorda 1987).
et al. 2017), 15 MCE sites have been studied systematically Coral reefs in the Philippines are exposed to surface water
to date (Fig. 15.1), spanning a broad range of environmental current systems driven by monsoon winds. There are two
settings from coastal fringing reefs such as Abra de Ilog to main seasons: the northeast monsoon from November to
Benham Rise, a deep, submerged bank located far offshore. mid-April and southwest monsoon from July to mid-­October.
15  The Philippines 267

114° 0’ 0” E 117° 0’ 0” E 120° 0’ 0” E 123° 0’ 0” E 126° 0’ 0” E

0 70 140 280 420 560

Patnanungan
Kilometers

21° 0’ 0” N
Abra de Ilog

0 4
km

18° 0’ 0” N
0 14
km Benham Rise

Bolinao

Masinloc N

15° 0’ 0” N
Patnanungan

Calaguas
Calatagan
Abra de Ilog
West Apo Reef

12° 0’ 0” N
Philippine
Sea El Nido Santa Fe

Siargao
Tubbataha

9° 0’ 0” N
Reef
Apo Island

Apo Reef
6° 0’ 0” N

Mati

0 2 Described in this report

km
Not described in this report

Fig. 15.1  Locations of MCEs in the Philippines surveyed as part of the GCM project of the Marine Science Institute of the University of the
Philippines. Green circles show the location of sites described herein

During inter-monsoonal periods, which occur from March to

June and in October, there is no predominant wind direction. 15.3 Habitat Description
Initial observations indicate upwelling around Abra de Ilog
(Fig.  15.2) and Apo Reef (Fig.  15.3), which may enhance Most of the coral reefs in the Philippines consist of fringing
connectivity between MCEs and SWRs and contribute to the reefs that vary in size from tens of meters to up to 5 km (Gomez
productivity and diversity in these areas, although this et al. 1994). A few atoll reef systems have been studied and are
hypothesis remains to be tested. protected as natural parks, such as the Tubbataha Natural Park
268 P. C. Cabaitan et al.

Table 15.1  Summary of information of the MCE sites

Limits of
Continuity Geological Physical Biological coral Research
Site with SWRs characteristics characteristics characteristics dominance Methods conducted by
*Abra de Ilog Yes Fringing reefs; Upwelling area; Community 30 m Diver-­ UP, Marine
dominated by water movement structure of based Science
extensions of valleys influenced by corals and fishes; surveys Institute
and headlands; monsoons presence/absence and ROV
sediments were data on other surveys
mostly fine and benthic
muddy organisms
Apo Island Yes Fringing reefs Unknown Fish community Unknown BRUVS Silliman
structure University
*Apo Reef SWRs in Fringing and atoll General current Community 60–70 m Diver-­ UP, Marine
fringing reefs; terraces and direction from the structure of based Science
reefs are scarps present; south moving corals and fishes; surveys Institute
continuous carbonate sediments toward the lagoon presence/absence and ROV
with MCEs area; thermally data on other surveys
stratified water benthic
column organisms
Benham Rise No Atoll reefs Data currently Fish community Unknown BRUVS UP, Los
undergoing analyses structure; and ROV Baños; UP,
presence/absence surveys Marine
data on coral Science
genera and other Institute;
benthic OCEANA
organisms
Bolinao Yes Fringing reefs Data was not Presence/absence 40 m RDV and UP, Marine
collected of coral genera diver-­ Science
and fish species based Institute
surveys
Calaguas Yes Fringing reefs; gentle Data currently Community Unknown Diver-­ UP, Marine
slope with three undergoing analyses structure of based Science
terraces in the corals and fishes, surveys Institute
mesophotic zone and other benthic and ROV
organisms surveys
Calatagan Yes Fringing reefs; Temperature Presence/absence Unknown Drop UP, Marine
distant features such profiles indicate a of coral genera cameras Science
as pinnacles and uniform water and other benthic and Institute
mounds present column; salinity organisms RDVs
profiles indicated a
freshwater lens
occurs at 2–4 m
depth, near the
surface
El Nido Yes Fringing reefs Data was not Coral community Unknown Diver-­ UP, Marine
collected structure; based Science
presence/absence surveys Institute
of other benthic and drop
organisms cameras
Masinloc Yes Fringing reefs Data was not Community Unknown RDV and UP, Marine
collected structure of diver-­ Science
corals and fishes; based Institute
presence/absence surveys
data on other
benthic
organisms
Mati Yes Fringing reefs Data was not Coral community Unknown Diver-­ UP, Marine
collected structure and based Science
presence/absence surveys Institute
of other benthic and drop
organisms cameras
(continued)
15  The Philippines 269

Table 15.1 (continued)
Limits of
Continuity Geological Physical Biological coral Research
Site with SWRs characteristics characteristics characteristics dominance Methods conducted by
*Patnanungan Yes Fringing reefs; debris General current Community Dominated Diver-­ UP-Marine
lobes, paleochannels, direction is toward structure of by algae in based Science
smooth seafloor, and the island; small corals and fishes; SWRs and surveys Institute
terraces present; variation in the presence/absence MCEs and ROV
seafloor has temperature-salinity data on other surveys
carbonate and silicate profile, indicative of benthic
materials mixing during the organisms
ebb tide
Santa Fe Yes Fringing reefs Data was not Coral community Unknown RDV UP, Marine
collected structure; surveys Science
presence/absence Institute
of other benthic
organisms
Siargao Yes Fringing reefs Data was not Coral community Unknown Diver-­ UP, Marine
collected structure; based Science
presence/absence surveys Institute
of other benthic and drop
organisms cameras
Tubbataha No Atoll reefs Data currently Presence/absence 100 m ROV UP, Marine
Reefs undergoing analyses of coral genera surveys Science
and fish species Institute
West No Atoll reefs Data currently Community Unknown Diver-­ UP, Marine
Philippine undergoing analyses structure of based and Science
Sea corals and fishes, RDV Institute
and other benthic surveys
organisms
Sites marked with * are described in more detail in the text
BRUVS baited remote underwater video system, RDV remotely deployed videos, UP University of the Philippines

(Dygico et al. 2013) and Apo Reef Natural Park (Quimpo et al. Philippine Sea, some sites in Apo Reef, and Benham Rise.
2018a). A number of patch reefs are located in the western Benham Rise, recently renamed the Philippine Rise, is a 13
Philippines, particularly in the Spratly Islands, locally known million-hectare undersea plateau (Nacorda et al. 2017) which
as the Kalayaan Islands (Alino and Quibilan 2003). rises from 3000  m to ~50  m at its shallowest point and is
The high diversity of SWRs in the Philippines undoubt- known to support very high coral cover. Most of the other
edly extends into mesophotic depths, resulting in strong sites represent the lower reef slopes of SWRs. Here, we pres-
regional differences in MCE communities. The 15 sites ent a detailed description of the three main study sites of
examined during the GCM aimed to capture some of the GCM: Patnanungan, Abra de Ilog, and Apo Reef (Fig. 15.1).
variation in MCE habitats and span a diverse range of reef The three sites were selected because they differ greatly in
environments from the deep slopes of fringing reefs to sub- geological setting, environmental conditions, disturbance his-
merged banks. The geologic setting can strongly influence tory, and human population density (Fig. 15.4). These three
the characteristic geomorphology of each site. For example, sites, however, all had seafloors in the upper mesophotic zone
some fringing reefs support continuous reef habitat extend- (30–40  m) dominated by dead corals and abiotic features,
ing into mesophotic depths, while others exhibit a wide span including coral rubble, sand, and silt (Figs. 15.5 and 15.6).
of sandy area that separates MCEs from SWRs. In Apo Reef Patnanungan reef is a fringing reef situated on an uplifted
Natural Park, an oceanic atoll, SWRs are separated from island on the Pacific side of the Philippines underlain by
MCEs, which are concentrated on the edges of reef terraces siliciclastic sedimentary rocks within the Bicol Shelf (Karig
separated by steep scarps. Multibeam bathymetric maps 1983; Lewis and Hayes 1983). The shallow fringing reef at
have revealed the existence of these terraces and scarps at Patnanungan is exposed to strong surge and current from the
multiple depth intervals between 40 and 150  m around all Pacific (Carpenter 1998). High-resolution multibeam
three sites, interpreted as the crests of reefs that were drowned bathymetry shows submerged paleochannels at the shallow-
by postglacial sea level rise. Such terraces and reef scarps are est portion of the reef. Terraces also occur at depths of
common antecedent structures that constitute an important 20–30  m, 40  m, and 80–90  m and are separated by steep
substrate for MCEs. scarps where slumps and debris lobes were observed. The
Three sites surveyed during the GCM were submerged terraces in Patnanungan could be drowned coastal plains.
banks located offshore with no contiguous SWRs: in West Algal cover in these terraces are relatively high (59%) com-
270 P. C. Cabaitan et al.

a 30.00
VIP

14°N

2nd

Latitude
50
Transect 40
29.50
1st 30
Transect 20
36 48 121°E 12 24
Longitude
Latitude

13°N 0.3 m/s

29.00

28.50

28.00

48.00 48.50 120°E 49.50 50.00

Longitude

b
1 0.4
20
0.2
Depth (m)

40 offshore
0
60
-0.2
80
100 -0.4

13.479 13.48 13.481 13.482 13.483 13.484 13.485 13.486 13.487 13.488
Latitude
South North

0.4
2
20
offshore 0.2
Depth (m)

40

60
0
80
nearshore
100 -0.2

13.48 13.481 13.482 13.483 13.484 13.485 13.486 13.487 13.488

Latitude

Fig. 15.2  Surface current velocities (a) and indicators of upwelling around Abra de Ilog based on depth-latitudinal flow (b). In (b), color contour
represents meridional velocity of mean current for two transects (labeled 1 and 2); the red color indicates northward motion, while the blue color
indicates southward motion. Looking at the meridional component of flow, that is, the north-south component of flow, the upper 70 m of the water
column is predominantly moving to the north away from the coast, while the water column below is moving to the south or toward the coast. This
combination of flow is indicative of upwelling along the coast
 15  The Philippines 271

a 12.74
Flood Tide
Ebb Tide
12.72

12.7

Latitude
12.68

12.66

12.64

12.62

120.4 120.42 120.44 120.46 120.48 120.5 120.52 120.54 120.56

Longitude
b
44 44
Latitude (∞N)

Latitude (∞N)

43 43

12° 12°
42 42

41 41

26 27 28 120° 30 31 32 33 26 27 28 120° 30 31 32 33
Longitude (∞E) Longitude (∞E)
Salinity Salinity
0 34 0 34
a a
33.5 5 33.5
10
33 10
33
Depth

Depth

32.5 15
20 32.5
Ocean Data View/DIVA

Ocean Data View/DIVA

20
32
32
30 25
31.5
30 31.5
12.7°N 12.71°N 12.72°N 12.73°N 12.68°N 12.69°N 12.7°N 12.71°N 12.72°N 12.73°N 12.74°N
Latitude [degrees_North] Latitude [degrees_North]
Temperature Temperature
0 30 0
b b 30
5
10 29.5
10 29.5
Depth

Depth

29 15
20
29
Ocean Data View/DIVA

Ocean Data View/DIVA

20
28.5
30 25 28.5
28 30
12.7°N 12.71°N 12.72°N 12.73°N 12.68°N 12.69°N 12.7°N 12.71°N 12.72°N 12.73°N 12.74°N
Latitude [degrees_North] Latitude [degrees_North]

Fig. 15.3  Surface current velocities (a) and indicators of upwelling around Apo Reef based on salinity and temperature cross-sectional plots (b).
In (b), maps represent sampling locations. Temperature plots show that the colder water temperatures dome toward the surface, indicating the
presence of an upwelling
272 P. C. Cabaitan et al.

Fig. 15.4  Multibeam bathymetry maps of the study sites at Apo Reef (a), Abra de Ilog (b), and Patnanungan (c) showing the complexity of each
reef’s morphology. Geomorphic features such as terraces and scarps (white arrows), ridges, debris lobes, and paleochannels are indicated

prised mostly of turf algae, Padina, and Halimeda. Abundant in the Philippines (Carpenter and Springer 2005), is strongly
sea fans, such as Stylaster, are found in steep scarps. influenced by terrestrial runoff. SWRs in Abra de Ilog are
Abra de Ilog is located in the Verde Island Passage within not extensive and do not have a reef flat. The MCEs in Abra
the southern extent of the Macolod Corridor, a rift zone de Ilog receive sediment from two nearby rivers that drain
(Oles et al. 1991; Pubellier et al. 2000; Ohkura et al. 2001). the northern flanks of Mindoro Island. Consequently, the
The Verde Island Passage, the center of marine biodiversity occurrence of MCEs is patchy and appears to be dependent
15  The Philippines 273

Overall

Apo Reef

Abra de Ilog

Patnanungan

0% 20% 40% 60% 80% 100%

Coral Dead coral Other coral Octocoral

Invertebrates Algae Seagrass Abiotic

Fig. 15.5  Benthic community composition (% cover) in three MCE sites

Fig. 15.6  Sample snapshots at each surveyed MCE site across depths, illustrating changes in benthic community composition. (Photo credits:
Ronald Dionnie D. Olavides)

on rock outcroppings above the muddy siliciclastic sand. adjacent atolls and a smaller emergent limestone island sur-
Due to this, dead corals and abiotic features dominate the rounded by fringing reefs. The two atolls both contain a smaller
benthos. Sea fans, sea whips, soft corals, calcareous algae, limestone island and are separated by a channel which deepens
and sponges are observed, along with some encrusting and and opens to the west. Multiple terraces and ridges were gener-
foliose forms of Montipora and Porites, in patch reefs at ally observed between 40–50 m, 70–80 m, and ~150 m which
40 m depth. separate several reef communities from each other. These ter-
Apo Reef is located 15 km west of the island of Mindoro in races occur on the atoll walls, the broad shallow platforms on
the Mindoro Strait and is situated near a descending oceanic the west side of the channel, and the east side of Apo Reef. Due
plate toward the Manila Trench. Apo Reef is composed of two to its offshore location, Apo Reef receives little terrestrial sedi-
274 P. C. Cabaitan et al.

ment. Encrusting and foliose forms of corals and gorgonians Table 15.2 (continued)
dominate the reef walls and rocky transitions of these terraces. Family/genera Apo Reef Abra de Ilog Patnanungan
Beyond 60 m depth, few hydroids, sponges, and sea fans are Diploastreidae
conspicuous. Apo Reef is located in a national park and is a  Diploastrea x
nominated UNESCO World Heritage site. It is one of the larg- Euphylliidae
est ­“no-­take” marine protected areas in the country, and perma-  Catalaphyllia x
 Euphyllia x
nent human settlements are not permitted on the island (Weeks
 Galaxea x
et al. 2010; Tabaranza et al. 2014).
 Physogyra
 Plerogyra x x x
Faviidae
15.4 Biodiversity  Cyphastrea
 Favia x x
15.4.1 Macroalgae  Goniastrea x
 Montastrea x x
The biodiversity of algae in MCEs in the Philippines has not  Moseleya x
been explicitly surveyed. In our assessment of benthic habi- Fungiidae
 Ctenactis x
tats, few species of algae could be identified, but further
 Fungia x x x
work may uncover diversity of algae. Algae assemblage was  Heliofungia x
different across sites, with Apo Reef having the most diver-  Lithophyllon
sity, followed by Patnanungan and Abra de Ilog. In Apo  Podabacia
Reef, at least seven algae species were identified: Caulerpa Helioporidae
racemosa, Claudea sp., Halimeda macroloba, Halimeda  Helioporaa x
opuntia, Halymenia durvillei, Halymenia maculata, and Lobophylliidae
Portieria hornemannii. Four species were observed in  Lobophyllia x
Patnanungan at a depth range of 30–40 m, characterized by Merulinidae
 Echinopora x
thick beds of Sargassum along with other brown algal spe-
 Favites x x
cies such as Padina japonica, P. australis, and P. minor. In
 Merulina x x
Abra de Ilog, the algal assemblage was mostly filamentous  Oulophyllia x
brown algae that grew in coral debris. These algae were not  Platygyra x
identified taxonomically because of the lack of morphologi- Mussidae
cally distinct features.  Scolymia
 Symphyllia x
Pectiniidae
 Echinophyllia x x
 Mycedium x
Table 15.2  Coral genera recorded in three upper mesophotic (30–  Oxypora x x
40 m) sites in the Philippines Pocilloporidae
Family/genera Apo Reef Abra de Ilog Patnanungan  Pocillopora x
Acroporidae  Seriatopora x x
 Acropora x x x  Stylophora
 Anacropora x Poritidae
 Astreopora x  Alveopora x
 Isopora x  Goniopora
 Montipora x x  Porites x x x
Agariciidae Siderastreidae
 Gardineroseris x  Psammocora
 Leptoseris x x x Stylasteridae
 Pachyseris x x x  Stylastera x
 Pavona x Trachyphylliidae
 Pectinia x  Trachyphyllia x x
Dendrophylliidae Tubiporidae
 Duncanopsammia x  Tubiporaa x x
 Tubastrea x Total # of families (18) 16 6 13
 Turbinaria x Total # of genera (51) 38 8 19
(continued) Non-scleractinian coral
a
15  The Philippines 275

15.4.2 Anthozoans to massive and branching forms in the SWRs. Encrusting

and foliose corals were the most abundant growth forms at
At least 46 coral genera from 16 families were identified mesophotic depths at all sites, a response to the reduced
from the three upper mesophotic sites (30–40 m depth) from light irradiance in the mesophotic zone (Lesser et  al.
diver-based surveys: 38 genera in Apo Reef, 8  in Abra de 2009).
Ilog, and 19  in Patnanungan (Table  15.2). Porites was the Azooxanthellate corals were also observed in the study
most abundant genus at all sites, which comprised about sites; however, these were not taxonomically classified;
42% of all coral colonies recorded in surveyed photoquad- instead, they are described by their common names. All of
rats, followed by Acropora (10%) and Seriatopora (9%). the azooxanthellate corals (i.e., black corals, soft corals, sea
Similarly, coral density (individuals per m2) was highest at fans, sea whips, and gorgonians) were not abundant in all the
Apo Reef, followed by Abra de Ilog and Patnanungan. Other sites surveyed. They were particularly rare at a depth range
common corals at mesophotic depths were Mycedium, of 30–40 m; however, beyond 40 m up until a depth of 165 m,
Leptoseris, and the azooxanthellate Tubastraea micranthus. they were frequently observed with the ROV, although the
Seriatopora hystrix and Porites rus were both relatively maximum depth range that they occupied was highly site
common on both shallow and mesophotic reefs. specific. For example, their maximum depth range in Abra
The dominant morphology of Porites varied with depth, de Ilog was 47  m, 146  m for Apo Reef, and 165  m for
with encrusting and foliose forms in the MCEs compared Patnanungan (Fig. 15.7).

Fig. 15.7  Gorgonian coexisting with Sargassum algae at MCE in Patnanungan (a), soft coral beds (b) and Tubastrea micranthus at MCE in
Apo Reef (c), and gorgonian at MCE in Abra de Ilog. (Photo credits: Edwin E. Dumalagan Jr.)
276 P. C. Cabaitan et al.

15.4.3 Sponges (Sphyraena barracuda and Gymnosarda unicolor) had the

highest biomass. In Patnanungan, there was no discernible
Sponges were observed in MCEs in all the sites; however, differences in biomass across the trophic groups, probably
these were not taxonomically identified as observations were because fish biomass overall was low due to the degraded
done using ROV.  In all three sites, Apo Reef had the most state of the site and high fishing pressure.
diverse morphology of mesophotic sponges. We observed four
different morphologies: basket, vase, flat, and barrel sponges
(presumably of the genus Xestospongia). In Patnanungan, 15.4.5 Other Biotic Components
only two sponge morphologies were observed: epiphytic with
the macroalgae Sargassum and encrusting. In Abra de Ilog, Four species of macrobenthic mobile fauna were observed in
only encrusting sponges were observed; however, they were Apo Reef, including three species of Holothuroidea (sea
covered with silt. The maximum distribution of sponges was cucumbers Holothuria atra, H. ananas, and H. rubralineata)
highly site specific. For instance, the maximum distribution in and one species of sea star (Thromidia catalai). Most of
Abra de Ilog was 47 m, while in Apo Reef, it was 60 m. In these individuals were observed at a depth range of 30–40 m,
Patnanungan, sponges were only observed via SCUBA, which except for H. atra which was observed at 79 m.
was confined to a depth range of 30–40 m.

15.5 Ecology
15.4.4 Fishes
Coral cover was low (3–4%) in Patnanungan and Abra de
Based on diver-based visual censuses, fish species richness Ilog but was higher at Apo Reef (~25%). The coral cover in
generally decreased with depth across all sites, with 140 spe- these MCEs is comparable to most of the SWRs in the
cies from 23 families observed in 30–40 m depth and 35 spe- Philippines, where coral cover is below 50% (Licuanan et al.
cies in 17 families at 40–100  m (Table  15.3). The lower 2017). Algal cover was relatively high (59%) at Patnanungan,
species richness at depths >40 m may be attributed to a num- which may be considered a mesophotic algae reef system
ber of factors including lower sampling effort at these depths (Lesser and Slattery 2011). The depth at which benthic com-
where only RDVs and ROVs were used, vehicle avoidance, position transitions from being dominated by phototrophic
and time of sampling. Only two species were observed taxa such as corals to filter feeders (e.g., sponges or azooxan-
beyond 100 m—the pelagic thresher shark (Alopias pelagi- thellate soft corals) varied at the different sites and appeared
cus) and an unidentified cardinalfish (family Apogonidae). to be related to water clarity. The transition in Abra de Ilog,
Although 16 species of apogonids are known to occupy a site located between two rivers, occurred at a depth of
depths of 100–290 m in the Indo-Pacific (Froese and Pauly ~30 m, whereas in Apo Reef, the transition occurred at 60 m
2017), the unidentified cardinalfish may be a new species. (Fig. 15.6). There was no obvious transition observed in the
Unfortunately, the poor quality of the image obtained from MCEs at Patnanungan, with algae dominant on both SWRs
the ROV precluded accurate species-level identification. and MCEs, perhaps because of the high water clarity.
Species richness (Table 15.3) in the upper mesophotic Dynamite fishing, which is still rampant in the area, and the
zone (30–40  m) was highest at Apo Reef, followed by low abundance of herbivorous fishes also likely contribute to
Abra de Ilog and Patnanungan (Quimpo et  al. 2018b). the rubble fields and the proliferation of algae in Patnanungan.
Planktivorous fishes were the most abundant trophic Below 40 m depth, only qualitative descriptions based on
group across all sites (Fig.  15.8). The dominant plankti- ROV surveys were conducted because of the difficulty in
vores at Apo Reef were Chromis margaritifer, Acanthurus standardizing the sampling area (e.g., the ROV must be kept
thompsoni, and Dascyllus reticulatus (see Fig.  15.9 for at a minimum distance from the benthos to systematically
other species of planktivorous fishes). At Abra de Ilog, the sample a quadrat), and the resolution of ROV images was not
most abundant planktivores were Pomacentrus brachialis, sufficient to identify benthic organisms to species. Hence,
Amblyglyphidodon aureus, and A. leucogaster, while common names were used to describe the benthic fauna
Sufflamen chrysopterus, Pseudanthias pleurotaenia, and observed. In Abra de Ilog, taxa observed beyond 40  m
P. coelistes were the most dominant in Patnanungan. included sea fans, sea whips, soft corals, black corals, calcar-
On the other hand, the trophic group with the highest fish eous algae, and sponges. No sessile benthic fauna were
biomass varied in the different sites. In Abra de Ilog, omni- observed from 47 to 61  m, where the substrate was com-
vores had the highest fish biomass, attributed primarily to posed of fine sand with burrows. In Apo Reef, hard corals
two snapper species (Lutjanus lutjanus and Lutjanus sp.). In were observed up to a depth of 60–70 m. Some conspicuous
Apo Reef, the planktivores, benthic invertivores (Paracaesio coral genera (e.g., encrusting and foliose Montipora and
xanthura and Cheilinus undulatus), and piscivores Porites) could be identified from the ROV images. Beyond
15  The Philippines 277

Table 15.3  Fish species recorded at 30–40 m and 40–100 m depths in three mesophotic sites in the Philippines
Apo Reef Abra de Ilog Patnanungan
Families/species 30–40 40–100 30–40 40–100 30–40 40–100
Acanthuridae
 Acanthurus auranticavus x x
 Acanthurus japonicus x x
 Acanthurus pyroferus x x
 Acanthurus sp. x
 Acanthurus thompsoni x x
 Ctenochaetus binotatus x x x
 Ctenochaetus striatus x
 Naso hexacanthus x x
 Naso lituratus x
 Naso minor x
 Naso vlamingii x
 Unidentified x
 Zebrasoma scopas x
 Zebrasoma veliferum x
Alopiidae
 Alopias pelagicus x
Apogonidae
 Unidentified x
Balistidae
 Balistoides conspicillum x
 Odonus niger x x x
 Sufflamen bursa x x x
 Sufflamen chrysopterus x x
 Unidentified x
 Xanthichthys auromarginatus x
Caesionidae
 Caesio cuning x
 Caesio teres x x
 Pterocaesio pisang x x x
 Pterocaesio tile x x x
 Unidentified x
Carangidae
 Caranx lugubris x
 Caranx sexfasciatus x x
Chaetodontidae
 Chaetodon auriga x
 Chaetodon baronessa x
 Chaetodon burgessi x
 Chaetodon kleinii x x x x x
 Chaetodon lunulatus x
 Chaetodon melannotus x
 Chaetodon octofasciatus
 Chaetodon ornatissimus x x
 Chaetodon punctatofasciatus x x
 Chaetodon speculum x
 Chaetodon unimaculatus x
 Chaetodon xanthurus x
 Coradion melanopus x
 Forcipiger flavissimus x
 Forcipiger longirostris x x
 Hemitaurichthys polylepis x x
 Heniochus acuminatus x x x
(continued)
278 P. C. Cabaitan et al.

Table 15.3 (continued)
Apo Reef Abra de Ilog Patnanungan
Families/species 30–40 40–100 30–40 40–100 30–40 40–100
 Heniochus chrysostomus x
 Heniochus monoceros x
 Heniochus varius x
 Unidentified x
Dasyatidae
 Taeniura lymma x
Haemulidae
 Plectorhinchus lineatus x
Holocentridae
 Myripristis botche x
 Myripristis murdjan x
 Myripristis sp. x
 Sargocentron caudimaculatum x
 Sargocentron melanospilos x
 Sargocentron sp. x
Labridae
 Bodianus mesothorax x x
 Cheilinus fasciatus x
 Cheilinus undulatus x
 Choerodon fasciatus
 Cirrhilabrus cyanopleura x
 Coris aygula x
 Epibulus insidiator x
 Halichoeres chrysus x
 Halichoeres hortulanus x
 Hemigymnus fasciatus x
 Labrichthys unilineatus x
 Labroides dimidiatus x x x
 Labropsis manabei
 Oxycheilinus digramma x
 Oxycheilinus unifasciatus x
 Unidentified x
Lethrinidae
 Gnathodentex aureolineatus x
 Monotaxis grandoculis x
Lutjanidae
 Lutjanus bohar x
 Lutjanus decussatus x
 Lutjanus kasmira x x
 Lutjanus monostigma x
 Lutjanus russellii x
 Lutjanus sp. x
 Macolor macularis x x
 Paracaesio xanthura x
 Pinjalo lewisi x
 Unidentified x
Malacanthidae
 Hoplolatilus starcki x
Mullidae
 Parupeneus multifasciatus x x x
 Parupeneus trifasciatus x
 Unidentified x
(continued)
15  The Philippines 279

Table 15.3 (continued)
Apo Reef Abra de Ilog Patnanungan
Families/species 30–40 40–100 30–40 40–100 30–40 40–100
Nemipteridae
 Pentapodus bifasciatus x
 Pentapodus caninus
 Pentapodus paradiseus x
 Scolopsis bilineata x x
 Scolopsis ciliatus x
 Scolopsis margaritifera x
Pinguipedidae
 Parapercis cylindrica x
 Parapercis hexophtalma
Plotosidae
 Plotosus lineatus x
Pomacanthidae
 Apolemichthys trimaculatus x x
 Centropyge bicolor x x
 Centropyge flavissimus x
 Centropyge heraldi x
 Centropyge vroliki x
 Chaetodontoplus melanosome x
 Genicanthus lamarck x
 Pomacanthus imperator x x
 Pygoplites diacanthus x x
Pomacentridae
 Acanthochromis polyacanthus x
 Amblyglyphidodon aureus x x
 Amblyglyphidodon curacao x
 Amblyglyphidodon leucogaster x x
 Amphiprion frenatus x
 Amphiprion perideraion x
 Chromis alpha x
 Chromis amboinensis x
 Chromis stripes x
 Chromis flavipectoralis x x
 Chromis margaritifer x x
 Chromis ternatensis x
 Chromis viridis x
 Chromis xanthura x
 Dascyllus reticulatus x
 Dascyllus trimaculatus x
 Neoglyphidodon melas x
 Neoglyphidodon nigroris
 Neoglyphidodon thoracotaeniatus x
 Neopomacentrus azysron x
 Pomacentrus alexanderae x
 Pomacentrus amboinensis x
 Pomacentrus auriventris x
 Pomacentrus brachialis x x x
 Pomacentrus coelestis x
 Pomacentrus moluccensis x
 Pomacentrus nigromanus x
 Pomacentrus nigromarginatus x x
 Pomacentrus philippinus x
 Pomacentrus stigma x x
 Pomacentrus vaiuli x x
 Unidentified x
(continued)
Table 15.3 (continued)
Apo Reef Abra de Ilog Patnanungan
Families/species 30–40 40–100 30–40 40–100 30–40 40–100
Priacanthidae
 Priacanthus blochii x
Scaridae
 Chlorurus bowersi
 Scarus chameleon x
 Scarus forsteni x
 Scarus niger x
 Scarus sp. x
Scombridae
 Gymnosarda unicolor x
Serranidae
 Cephalopholis argus x
 Cephalopholis urodeta x x
 Epinephelus merra x
 Gracila albomarginata x
 Pseudanthias pleurotaenia x x x x x
 Pseudanthias sp. x
Siganidae
 Siganus vulpinus x
Synodontidae
 Synodus variegatus x x
Tetraodontidae
 Arothron nigropunctatus x x
Zanclidae
 Zanclus cornutus x x x x x
Total # of families (28) 20 9 16 4 11 13
Total # of species (148) 104 16 39 5 19 18

150 a b

15
Mean abundance of fishes per 1000 m2

Mean biomass of fishes in g per m2

Benthic Invertivore

100 Coralivore

Herbivore
10
Detritivore

Piscivore

Omnivore
50
Planktivore
5

0 0

Abra Apo Patnanungan Abra Apo Patnanungan

de Ilog Reef de Ilog Reef
Trophic group

Fig. 15.8  Mean abundance (a) and mean biomass (b) of reef fish trophic groups in the three upper mesophotic sites (30–40 m depth) obtained
using diver-based surveys
15  The Philippines 281

Fig. 15.9  ROV images of the planktivorous reef fishes observed in Apo Reef at a depth range of 30–50 m. (a) Pyramid butterflyfish (Hemitaurichthys
polylepis), (b) false fusilier (Paracaesio xanthurus), (c) blue-dash fusilier (Pterocaesio tile), (d) yellow-tail fusilier (Caesio teres), and (e) sleek
unicornfish (Naso hexacanthus). (Photo credits: Ronald Dionnie D. Olavides)
282 P. C. Cabaitan et al.

60 m, the benthic fauna were mostly dominated by hydroids, MCEs need to be protected for them to function as poten-
sponges (basket, vase, flat, and barrel of Xestospongia spp.), tial refugia. However, the protection of MCEs should focus
soft corals, gorgonians, sea whips, black corals, sea fans, on mitigating the local stressors within the area by designing
anemones, and crinoids. In Patnanungan, macroalgae genera appropriate management strategies. For example, dynamite
Halimeda, Sargassum, and Gracilaria were observed at a fishing should be banned in Patnanungan, whereas the runoff
depth of 50 m. Beyond 50 m, sea fans, soft corals, sea whips, from agriculture (e.g., pesticides and fertilizers) could be
and crinoids were observed to a depth of 65  m. At depths minimized in Abra de Ilog. In protecting MCEs, the
between 65 and 165 m in Apo Reef and Patnanungan, cri- ­communities may be allowed to recover from natural distur-
noids and whip corals were observed, respectively. bances, as exemplified by the Apo Reef National Park where
Megafauna (organisms with >50  cm body length) were juvenile corals are now observed in high abundance. The
observed only in Apo Reef, a potential indicator of reduced higher abundance of large and commercially important fish
fishing pressure and anthropogenic disturbance relative to species at Apo Reef (Quimpo et al. 2018a, b) compared to
other sites, and a more intact food web capable of supporting the other two sites also suggests that marine protected areas
larger-bodied organisms. The abundance of megafauna was can be effective for protecting large-bodied and commer-
higher in SWRs than in MCEs (Quimpo et  al. 2018a). cially important species from overexploitation. While marine
Megafauna observed in upper mesophotic depths (30–40 m) protected areas cannot guard against climate-related distur-
included hawksbill sea turtle (Eretmochelys imbricata), bances such as coral bleaching, evidence from Apo Reef
humphead wrasse (Cheilinus undulatus), and whitetip reef suggests that they can protect fish biomass and trophic struc-
shark (Triaenodon obesus), while the pelagic thresher shark ture, which in turn may assist recovery from disturbances.
(Alopias pelagicus) was observed at 113  m. Many marine
megafauna are known to have large foraging areas that may
include different depths (Papastiamatou et  al. 2015), and 15.7 Conclusions
MCEs may represent important habitats for these species.
Preliminary results confirm that MCEs in the Philippines
support diverse ecological communities, but there is a need
15.6 Threats and Conservation Issues to further assess other MCEs in different geographic regions
and environmental settings. Assessing representative MCEs
Similar to many of the SWRs in the Philippines (Nañola in different biogeographic regions will allow the identifica-
et al. 2011), some MCEs are under threat from human and tion of ecological, geological, and oceanographic factors that
natural disturbances. For instance, the high cover of mac- influence the biodiversity in MCEs at different spatial scales.
roalgae and low number of herbivores in Patnanungan are For example, understanding connectivity among MCEs and
indicative of overfishing. Although MCEs are perceived as between SWRs and adjacent MCEs is essential for under-
less likely to be impacted by disturbances than SWRs, MCEs standing the vulnerability of MCEs to disturbances and for
are not safe from overfishing. Spear fishing by hookah diving testing the “deep reef refuge” hypothesis (Bongaerts et  al.
and dragnet fishing can potentially exploit fish populations 2010). Identifying similarities in composition of coral reef
and damage habitat complexity on deeper reefs (Lindfield communities between SWRs and MCEs represents an initial
et al. 2014, 2016). Moreover, MCEs such as Abra de Ilog, step toward the goal of identifying depth refuge. Establishing
which are naturally exposed to turbid waters due to proxim- a long-term monitoring program would allow us to test
ity to a river may be vulnerable to sedimentation or reduc- whether reef communities that seem to be more connected
tions in light irradiance if river runoff is eutrophic or highly based on oceanographic and genetic evidence can indeed
sedimented from inland agriculture activities and coastal recover faster from disturbances than more isolated reefs.
development (Kahng et al. 2010). MCEs can also be affected Further studies should also focus on quantifying MCE biodi-
by thermal stress and associated coral bleaching and mortal- versity and taxonomic classification of many marine taxa
ity as was observed in Apo Reef. In 2016, qualitative obser- (e.g., sponges, soft corals), particularly since mesophotic
vations indicated that mesophotic corals bleached, with most biodiversity in the Coral Triangle is poorly known. Moreover,
of the affected colonies exhibiting a branching morphology. institutional linkages with other academic institutions, non-
However, the severity of the coral bleaching was lower in governmental organizations, and government agencies are
mesophotic depths compared to the SWRs of Apo Reef. warranted for the advancement of MCE research in the Coral
Philippine reefs are also subjected to typhoons that can cause Triangle.
physical destruction (Abesamis et al. 2017). The Philippines
has experienced a number of very severe typhoons in recent Acknowledgments We are grateful to the Biodiversity Management
years, a trend which is expected to increase due to climate Bureau of the Department of Environment and Natural Resources and
the Apo Reef Protected Area Management Board. We thank the
change.
15  The Philippines 283

Municipal Government of Sablayan, Albert Vallejo of Sablayan Tourism Hinderstein LM, Marr JCA, Martinez FA, Dowgiallo MJ, Puglise KA,
Office, Fernan Magno of DENR-PAMB, Mark Tolentino, and Frederico Pyle RL, Zawada DG, Appeldoorn R (2010) Theme section on
Sabban for their assistance in the field surveys. We are also grateful to “Mesophotic coral ecosystems: characterization, ecology, and man-
the three anonymous reviewers that provided inputs that improved the agement.” Coral Reefs 29(2):247–251
quality of the manuscript. This work was funded by the Philippine Kahng SE, García-Sais JR, Spalding HL, Brokovich E, Wagner D, Weil
Council for Agriculture, Aquatic and Natural Resources Research and E, Hinderstein L, Toonen RJ (2010) Community ecology of meso-
Development, Department of Science and Technology. photic coral reef ecosystems. Corals Reefs 29(2):255–275
Kane C, Kosaki RK, Wagner D (2014) High levels of mesophotic reef
fish endemism in the northwestern Hawaiian Islands. Bull Mar Sci
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