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Beneath 50 m of NW Pacific Water: Coral Reefs on the Benham Bank Seamount off the Philippine Sea

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The benthic habitats on the Philippine (Benham) Rise May 2014 when extensive coral reefs were discovered on the summit of the Benham Bank Seamount. Short observational surveys of five stations at depths up to 55 m revealed that the reefs were pristine and with excellent cover mostly by tiered, thick, rigid and foliose plate-forming Porites (Synaraea) rus. The voucher specimen collections indicated that there are at least 11 reef-building and two solitary coral species in the reef communities. The fish visual census and random hook-and-line fishing surveys recorded 62 species, 16 of which were reef health indicators and the rest were commercially exploited species. These short surveys yielded the first records of mesophotic coral reef biodiversity on the Benham Bank, albeit incomplete, and point to the inevitable requisite of further exploring these pristine reefs and their associated benthic habitats, since this Philippine natural heritage serves as an important area for fisheries.
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Journal of Environmental Science and Management 20-2:110-121 (December 2017) ISSN 0119-1144
Hildie Maria E. Nacorda1*
Romeo M. Dizon2
Lambert Anthony B. Meñez3
Cleto L. Nañola, Jr.4
Patrice Bianca L. Roa-Chio5
Diovanie O. De Jesus6
Homer B. Hernandez7
Fra-and Timothy R. Quimpo5
Wilfredo Roehl Y. Licuanan8
Porrio M. Aliño6
Cesar L. Villanoy6
1 School of Environmental Science
and Management, University of the
Philippines Los Baños (UPLB),
College, Laguna, 4031 Philippines
2 College of Science, UP Baguio,
Baguio City, 2600 Philippines
3 0272 Los Angeles, Brookside Hills,
Cainta, Rizal, 1900 Philippines
4 College of Science and Mathematics,
UP Mindanao, Davao City, 8022
Philippines
5 McKeough Marine Center, Xavier
University–Ateneo de Cagayan,
Cagayan de Oro City, 9000
Philippines
6 The Marine Science Institute,
UP Diliman, Quezon City, 1101
Philippines
7 85 V. Templo Avenue,
Mataasnakahoy, Batangas, 4223
Philippines
8 College of Science, De La Salle
University, Manila, 1004 Philippines
*Corresponding author:
henacorda@up.edu.ph
ABSTRACT
The benthic habitats on the Philippine (Benham) Rise were unknown until the
joint University of the Philippines Marine Science Institute (UPMSI)/University of
the Philippines Los Baños (UPLB)/Department of Agriculture-Bureau of Fisheries
and Aquatic Resources (DA-BFAR) cruise of May 2014 when extensive coral reefs
were discovered on the summit of the Benham Bank Seamount. Short observational
surveys of ve stations at depths up to 55 m revealed that the reefs were pristine and
with excellent cover mostly by tiered, thick, rigid and foliose plate-forming Porites
(Synaraea) rus. The voucher specimen collections indicated that there are at least 11
reef-building and two solitary coral species in the reef communities. The sh visual
census and random hook-and-line shing surveys recorded 62 species, 16 of which were
reef health indicators and the rest were commercially exploited species. These short
surveys yielded the rst records of mesophotic coral reef biodiversity on the Benham
Bank, albeit incomplete, and point to the inevitable requisite of further exploring
these pristine reefs and their associated benthic habitats, since this Philippine natural
heritage serves as an important area for sheries.
Key words: mesophotic coral reefs, reef sh, Benham Bank Seamount, Philippine
(Benham) Rise, Philippine Sea
INTRODUCTION
Seamounts are topographic rises of the ocean oor
that are ubiquitous but unevenly distributed among the
ocean basins (Wessel et al. 2010) and the large isolated
ones, i.e., with summits extending ≥1 km from the seaoor,
are mostly located in the Pacic basin (Kitchingman
JESAM
Beneath 50 m of NW Pacic Water: Coral Reefs on
the Benham Bank Seamount off the Philippine Sea
et al. 2007, Kim and Wessel 2011, Yesson et al. 2011).
Seamounts differ in characteristics—their bathymetry,
topography, substrates, and interactions with variable
background ow conditions result in a nutrient-enriched
water column and enhanced larval retention (Haney et
110
al. 1995, Richer de Forges et al. 2000, Koslow et al.
2001, Genin and Dower 2007, Pitcher and Bulman
2007, Lavelle and Mohn 2010), as well as curbed
local sedimentation (Boehlert and Genin 1987). These
conditions have concomitant inuence on standing stocks
and local distributions of benthic and pelagic organisms,
for example, seamount-aggregating sh and commercial
pelagic stocks (e.g., tuna species) (Uchida and Tagami
1984, Holland and Grubbs 2007, Morato and Clark
2007, Pitcher et al. 2010), pelagic sharks (Litvinov 2007),
marine mammals (Kaschner 2007), benthic algae (Littler
et al. 1986), and megafauna dominated by suspension
and/or lter-feeders, e.g., corals and sponges (Rogers
et al. 2007, Samadi et al. 2007, McClain et al. 2009).
The diversity of seamount assemblages may also be
relatively high at depths where the summit interacts with
the euphotic zone (Pitcher and Bulman 2007, Yesson et
al. 2011), where light-dependent benthic organisms such
as corals and macroalgae would recruit and establish.
Seamounts are also areas where seabirds aggregate
(Haney et al. 1995).
Deep sea environments of the Philippines have
been part of historic great expeditions, which revealed
the high diversity of the deep sea and resolved the old
perception that it was devoid of life. The collections of
these missions were deposited in various museums and
thereafter described, e.g., Philippine mollusks from the
Expedition of HMS Challenger (Watson 1879, 1881,
1882), various sh resources from the sheries steamer
Albatross (Smith and Williams 1999), and trawl fauna
of the Philippine Trench by the Galathea (Bruun 1951).
The same paradigm was supported by new species
discovered by more recent deep sea research projects,
e.g., PANGLAO 2005 (Richer de Forges and Ng 2007),
the Inner Space of the Celebes Sea (NOAA 2007, Osborn
et al. 2010), and the Philippine Biodiversity Expedition
(CAL 2011), among others. Subsequently, at least two
compendia of discoveries by the larger MUSORSTOM-
TDSB deep sea exploration program in the tropical Indo-
Pacic (1976-2012) also included rare and new species
of crustaceans from the Philippine archipelago (Richer
de Forges et al. 2014).
Discoveries of deep-water habitats are opportunities
to document biodiversity and understand its interactions
with the changing natural environment. Key populations
of shallow-water marine biodiversity hotspots may be
compared with counterparts in deep-waters to ascertain
habitat connectivity (e.g., Lesser et al. 2009) and
justify the establishment of, e.g., networks of marine
protected areas or large marine parks (e.g., Johnston and
Santillo 2004). For this study, the aim is to describe the
coral reef habitats discovered at summit depths of the
Benham Bank Seamount (BBS), a unique underwater
geological feature in the Philippine (Benham) Rise
Region off the Philippine Sea. The Bank may be a unique
biodiversity feature that allows for new perspectives on
the ecological and evolutionary signicance of isolated
seamounts. While the shallower biogeographic regions
have been proposed as the world's center of the center of
marine biodiversity (Carpenter and Springer 2005), the
eastern Philippine Sea occupies a considerable gap in the
explanation of the three dominant evolutionary theories,
i.e., center of origin, center of accumulation and overlap,
and the combined evolutionary ecology on the center
of refugia (Sanciangco et al. 2013). Understanding
the ecological and evolutionary explanations of the
phylogeography of the Benham Bank has profound
implications in unravelling its resiliency to the climate
change challenge, in addition to its implications on the
sustainability of the tuna stocks in the larger Philippine
(Benham) Rise Region. As this work jumpstarts further
assessments and monitoring of the marine biological
features of the Bank, essentially building up records of
Philippine deep-water biodiversity, the results may be
linked to the productivity of the Benham Rise Region,
where shing activities are known to occur. Finally,
this study consolidates the management and scientic
responsibility of the Philippines in this globally
signicant area.
MATERIALS AND METHODS
Study Area
The Philippine Rise (then referred to as the Benham
Rise) is one of several oceanic bathymetric highs that
currently impinge the Philippine archipelago (Yumul et
al. 2008). It is a submarine ridge of the West Philippine
Basin (Tetreault and Buiter 2014) and is a prominent
seaoor structure that is at least 49 Ma (Deschamps
and Lallemand 2002). The Benham Bank Seamount
(BBS), located at approximately 15°47'36.10"N and
124°17'47.65"E (Figure 1), rises from about 3,000
m of seaoor to its summit, which makes the area the
shallowest part of the Philippine Rise. The BBS is also
among the large seamounts in the Pacic basin (Yesson
et al. 2011). From earlier cruises, surface waters around
the BBS, often transported vigorously northward at ~2
knots, were found to mix with either the subtropical
western North Pacic water or the warmer North
Equatorial Current NEC) water when its bifurcation
latitude was located south (Gordon et al. 2014). These
strongly moving waters eventually feed into the waters
of the nascent Kuroshio Current. Cabrera et al. (2015)
111
Journal of Environmental Science and Management Vol. 20 No. 2 (December 2017)
subsequently found that chlorophyll a of surface waters
were also inuenced by the NEC’s bifurcation latitude,
so the typical oligotrophic waters of the Benham Bank
area may also occasionally exhibit high chlorophyll a,
i.e., when the NEC bifurcates at a southern latitude.
The UPMSI/UPLB/DA-BFAR Joint Cruise (Cruise
DY27 Leg 1), 3-18 May 2014
The Joint Cruise to the Benham Bank, carried out
aboard the research vessel M/V DA-BFAR (Figure 2)
by the University of the Philippines-Marine Science
Institute (UPMSI), UP Los Baños, through the School
of Environmental Science and Management (UPLB-
SESAM), and the Department of Agriculture-Bureau of
Fisheries and Aquatic Resources (DA-BFAR), departed
Pier 3 of the Manila North Harbor on 3 May 2014 and
arrived at the Benham Bank on the morning of 6 May.
From 6 to 16 May, diving activities were performed
on the summit and then oceanographic samplings were
conducted in the vicinity of the Bank (Benham Rise
Potential Productivity Project 2014). The cruise departed
the Bank on 16 May for the Port of Legazpi in Albay,
arrived on 17 May to disembark and load equipment and
samples in a hired truck; participants alighted the M/V
DA-BFAR on 18 May.
Coral Reef Assessment
Verifying Depth and Bottom Type. After the M/V
DA-BFAR anchored, underwater cameras (SplashCam
point of view camera and GoPro Hero cameras) and a
dive computer were lowered with a weighted line to lm
the bottom and conrm the depth reading at anchor point.
The short footages were reviewed after the cameras were
retrieved and then dives were thoroughly planned when
depth was dive-permitting and footages revealed a coral
reef bottom.
Diving. Divers used SCUBA for their bounce dives and
rapid underwater activities on the summit of the Benham
Bank. The divers were moved behind the ship’s stern
for the dives. Each diver was tethered to a weighted line
112 Coral reefs on the Benham Bank
Figure 1. Location of the seamount () on the Benham Bank, where coral reefs at mesophotic depths up to 55 m
were discovered and surveyed on 6-14 May 2014 during the UPMSI/UPLB/DA-BFAR Joint Cruise. The area
enclosed by the yellow polygon constitutes the extended continental shelf (ECS) of the Philippines, granted
in April 2012 by the UN Commission on the Limits of the Continental Shelf (CLCS) (Source of base map:
NAMRIA; http://www.namria.gov.ph/projects.aspx).
Figure 2. The marine vessel M/V DA-BFAR (DYCA) was
the research platform used during the UPMSI/
UPLB/DA-BFAR Joint Cruise, 6-14 May 2014.
The small rubber boats moved divers from
the starboard to behind the stern for diving
and underwater activities (Photo courtesy of
the Physical Oceanography Lab, UPMSI).
during descents to the bottom. The two most experienced
divers in the team served as tenders and deployed reserve
air tanks with regulators at three decompression depths
and then stayed tethered at the shallow decompression
depths. At the bottom, the divers worked within a radius
of ~10 m from the weighted line. The tenders sounded
off the signal when bottom time was reached (5-7 min),
hence, prompted divers to approach and then fasten their
carabiners to the weighted line prior to starting their ascent.
Whenever possible, three pairs of divers were
deployed sequentially to maximize the collection of data
from each station, allowing each pair to perform only
one specic task while at the bottom depth. Divers also
alternated assignments as deep divers and tenders between
dives. Safety procedures were followed in deploying
divers at the summit depths. The dives were scheduled
on mornings every other day from 6 to 14 May 2014.
Observational Surveys and Collections of Specimens.
At the ve dive stations, divers performed video- and
photographic documentation of the reef and associated
benthos. A rapid sh visual census (FVC) for 5 to 7 min
was done in Stations 1 and 2, in which reef sh species were
enumerated and sh total lengths (TL) were estimated. In
Stations 3, 4, and 5, two 5 x 10-m observation areas were
delimited for the rapid FVCs, where sh species were
also enumerated, sh individuals were counted, and TLs
were estimated, so that sh biomass can be calculated.
The divers also collected voucher specimens manually,
i.e., pieces of corals, conspicuous invertebrates (sponges,
soft corals), macroalgae, and samples of loose sediments.
Onboard Activities. All the samples collected were
photo-documented, i.e., the large ones were photographed
using digital SLRs while the smaller associated
organisms were photographed under M/V DA-BFAR’s
Nikon stereoscopic zoom microscope mounted with
a diascopic stand, a digital camera, and a stand-alone
monitor control unit. Thereafter, the coral specimens
were sun-dried, macroalgal samples were pressed onto
clean white paper and inserted into old newspapers, and
the macroinvertebrates (e.g., sponges and soft corals)
were xed in 70% ethanol in seawater. Subsamples for
microbial analyses were obtained from sediment samples
prior to preservation with 10% formalin in seawater.
Opportunistic shing by hook-and-line was carried
out by the crew of the M/V DA-BFAR in four stations
(Stations 2 to 5) to complement data from the FVCs.
These were done when the ship was still anchored and
divers were not underwater. The shes caught were
identied, measured, and photographed; some sh
individuals were frozen until preservation and archiving
at the UPMSI. Species lists from both rapid sh visual
censuses and hook-and- line shing events were merged
into one database and each species was classied as
either indicator or target species and as demersal or
pelagic species.
RESULTS AND DISCUSSION
The underwater activities were carried out at
veried depths from 51.3 (Station 2) to 55.1 m (Station
1) and with conrmed coral reef bottoms in ve stations
located on the eastern ridge of the Benham Bank summit
(Table 1).
Coral Reef Benthos
The coral reefs on the Benham Bank Seamount
were found to be in pristine condition and with excellent
benthic cover, i.e., 75 to 100% hard corals on the bottom
(Figure 3). The foliose plate-forming species, identied
as Porites (Synarea) rus (Figure 4), was found in all the
stations, particularly dominated the coral reef bottom
of Stations 1 and 2, and, at the same time, entirely
covered the substratum. Porites (Synarea) rus formed
circular tiers and gave an undulating appearance at
bottom depth; these were made up of heavy coralline
framework, indicating possible slow growth rates. The
reefscapes included other coral lifeforms, e.g., encrusting
Galaxea, encrusting and massive Porites, massive
Platygyra, branching Pocillopora, fungiids, and the blue
coral Heliopora coerulea. There were also occasional
tufts of the green algae Halimeda (Figure 5) observed,
including arborescent sponges and soft corals (Families
Alcyoniidae, Neptheidae, and Xeniidae). The divers also
noted sediments that became exposed as a result of the
ship's anchor being dragged at the bottom.
In Stations 3, 4, and 5, the reefs were more
heterogeneous (Figure 6) as other coral growth forms
(e.g., massive Porites and Astreopora, submassive
113
Journal of Environmental Science and Management Vol. 20 No. 2 (December 2017)
Table 1. Survey date, location, and depth of the ve coral
reef stations on the Benham Bank Seamount,
6-14 May 2014.
Date of
survey
(2014)
Station Location Depth
(m)
6 May
8 May
10 May
12 May
14 May
1
2
3
4
5
15o48.106’N, 124o18.160’E
15o48.060’N, 124o18.225’E
15o49.230’N, 124o18.210’E
15o49.214’N, 124o18.187’E
15o48.134’N, 124o18.258’E
55.1
51.3
54.8
54.4
53.8
Alveopora and Cyphastrea, branching Acropora and
Pocillopora) were observed as well as other conspicuous
lifeforms, i.e., soft corals (Neptheidae, Xenia sp.,
Sinularia sp., and Sarcophyton sp.), arborescent sponges,
and Halimeda. Coralline sand and calcied remains
of Halimeda constituted the loose surface sediments,
observed as large areas in between the coral communities
of Stations 3 and 4.
Based on examination and identication of
collected specimens, the following reef-building species
occurred on the Bank- Fungia scutaria, Cycloseris
cyclolites, Alveopora verrilliana, Cyphastrea calcidicum,
Cyphastrea. microphthalma, Galaxea astreata, G.
fascicularis, Pocillopora verrucosa, Acropora valida,
Porites lutea, Porites.murrayensis, Pavona minuta, and
the octocoral Heliopora coerulea, as previously noted
(Figure 7, a to m).
The thriving reefs were under clear waters that
allowed sunlight to reach their depths. Zooxanthellate
corals require sunlight in order for their symbiont algae
to carry out photosynthesis and, in turn, support coral
growth and survival. From a previous cruise in Lamon
Bay, light intensities down to 300 m were measured
(as PAR, photosynthetically active radiation) using a
sensor attached to the CTD (Cabrera et al. 2015). The
prole indicated that light received at the Bank summit
was only 12% of surface irradiance (~200 µE m-2 s-1),
which is quite low relative to light intensities received by
shallower reefs. Under such low light environment, only
particular taxa of certain growth forms have successfully
adapted and proliferated, thus, the diversity of coral
species and coral growth forms become limited. At ~125
m, light disappears and signals the depth at which the
aphotic zone in the Benham Bank begins (Cabrera et al.
2015).
114 Coral reefs on the Benham Bank
Figure 3. Foliose and plate-forming corals dominated the
reef bottoms of the Benham Bank Seamount
(photo grabbed from a video clip courtesy of
L.A.B. Meñez, 6 May 2014).
Figure 4. Fresh voucher specimens of Porites (Synarea)
rus, the dominant foliose plate-forming
scleractinian coral in the ve stations surveyed
on the summit of the Benham Bank Seamount,
6-14 May 2014 (scale bar: 2 cm) (photos
courtesy of G.S. Jacinto and D.O. De Jesus).
Figure 5. Tufts of the green algae Halimeda sp. with
hard corals on the Benham Bank Seamount.
Halimeda is a major contributor of loose
surface sediments on the Bank (photo
grabbed from a video clip courtesy of L.A.B.
Meñez, 6 May 2014).
Figure 6. Mix of coral growth forms in Station 4, Benham
Bank Seamount (photo grabbed from a video
clip courtesy of A.T. Yñiguez, 12 May 2014).
Reef-Associated and Pelagic Fishes
The combined data from underwater FVCs and
opportunistic hook and line shing listed a total of 61
bony shes and two cartilaginous sh, Triaenodon
obesus and Atelomycterus marmoratus, associated with
the coral reefs of the Benham Bank Seamount. Among
the reef shes, the most speciose families were the
Chaetodontidae (butteryshes, 10 species), Serranidae
(groupers, 9 species), and Acanthuridae (surgeonshes,
8 species) (Table 2). Five Families (16 species) were
known indicators of coral reef health- Chaetodontidae,
Pomacanthidae (angelshes, 2 species), Pomacentridae
(damselshes, 2 species), Ptereleotrididae (dart gobies, 1
species), and Zanclidae (Moorish idol, 1 species) (Table
2). Thirteen families (47 species) were commercially
important or food sh (Table 2). The schooling tunas
(Gymnosarda unicolor, made up of 5 to 15 individuals)
were the largest among the target pelagic species
observed, with an estimated total length (TL) of ~150
cm. The solitary rainbow runner (Elegatis bipinnulata,
Family Carangidae) observed was also large, sized ~70
cm TL. Only one individual was observed to be relatively
large among the demersal (coral reef-attached) species,
i.e., the footballer cod Plectropomus laevis (Family
Serranidae) at 80 cm TL (Table 2). The estimated pooled
115
Journal of Environmental Science and Management Vol. 20 No. 2 (December 2017)
Figure 7. Solitary corals (a, b) and reef-building species (c to m) on the Benham
Bank that occurred together with Porites (Synarea) rus, based on the
voucher specimens collected on 6-14 May 2014 (scale bar: 2 cm) (photos
courtesy of G.S. Jacinto and H.M.E. Nacorda).
116 Coral reefs on the Benham Bank
Table 2. Species, estimated sizes (in cm), and counts of demersal (site-attached) and pelagic shes recorded during
ve days of sh visual census (FVC) and four days of hook-and-line shing (*) on the summit of the Benham
Bank Seamount, 6-14 May 2014.
Family (Common Name)/ Species Size (cm) Classication Station
1 2 3 4 5
Chaetodontidae (butteryshes)
Chaetodon guentheri
Chaetodon kleinii
Chaetodon mertensii
Chaetodon punctatofasciatus
Chaetodon reticulatus
Chaetodon unimaculatus
Forcipiger longirostris
Hemitaurichthys polylepis
Heniochus monocerosa
Heniochus singularius
Pomacanthidae (angelshes)
Centropyge heraldi
Pomacanthus imperator
Pomacentridae (damselshes)
Chromis margaritifer
Dascyllus reticulatus
Ptereleotrididae (dart gobies)
Nemateleotris magnica
Zanclidae (Moorish idols)
Zanclus cornutus
Acanthuridae (surgeonshes)
Acanthurus nigricans
Acanthurus nigrofuscus
Acanthurus pyroferus
Acanthurus thompsoni
Naso brachycentron
Naso lopezi
Naso unicornis
Naso vlamingiia
Balistidae (triggershes)
Balistapus undulatus*
Balistoides conspicillium
Melichthys vidua*
Sufamen bursa*
Sufamen chrysoptera
Caesionidae (fusiliers)
Pterocaesio tile
Carangidae (trevallies)
Caranx sp.*
Elegatis bipinnulata
Holocentridae (squirrelshes)
Myripristis murdjan
Sargocentron caudimaculatum*
Sargocentron spiniferum*
Sargocentron tiere*
Labridae (wrasses)
Bodianus mesothorax
Cheilinus unifasciatus
8
8
8
10
12
8
10
12-14
16-25
16-18
8
25
5-6
6
7
12-14
10-12
10-12
12-14
12-15
40
30-50
40
30-50
23
15
22
16
10
14
43
70
16-18
17-19
32-35
27
14
12-16
Demersal; indicator
Demersal; indicator
Demersal; indicator
Demersal; indicator
Demersal; indicator
Demersal; indicator
Demersal; indicator
Demersal; indicator
Demersal; indicator
Demersal; indicator
Demersal; indicator
Demersal; indicator
Demersal; indicator
Demersal; indicator
Demersal; indicator
Demersal; indicator
Demersal; target
Demersal; target
Demersal; target
Demersal; target
Pelagic; target
Pelagic; target
Pelagic; target
Pelagic; target
Demersal; target
Demersal; target
Demersal; target
Demersal; target
Demersal; target
Pelagic, target
Pelagic, target
Pelagic, target
Demersal; target
Demersal; target
Demersal; target
Demersal; target
Demersal; target
Demersal; target
-
-
-
-
-
-
-
-
-
1
-
-
1
-
-
-
-
-
-
-
-
1
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
1
1
1
-
1
1
1
1
-
-
1
-
1
-
-
1
1
-
1
-
1
-
3
-
-
-
-
1
1
-
1
1
2
3
-
-
1
-
-
-
-
-
-
-
-
-
1
-
-
-
-
1
1
-
1
1
1
-
-
-
14
1
1
1
1
-
-
-
-
-
-
-
-
-
1
-
-
-
1
-
-
-
1
1
1
-
1
1
1
-
-
-
1
-
1
-
-
-
2
-
-
-
-
-
-
-
-
-
-
-
-
-
1
-
-
-
-
-
-
-
1
-
-
-
-
1
-
-
-
1
-
1
1
1
-
1
6
-
-
-
-
-
-
1
-
1
1
1
1
1
1
sh biomass, based on data of Stations 3 to 5, ranged from
17 to 102 mt km-2 (mean at 60 mt km-2), which is almost
threefold lower when compared to the mean biomass of
coral reef-associated shes in 10-m deep waters of the
protected Tubbataha Reefs National Park off the Sulu Sea
for the same year (~150 mt km-2; Dejucos et al. 2015).
This difference in sh biomass may be veried when the
communities are sampled through comparable methods,
or sufciently using underwater technologies for deep
water environments, with other representative reef areas
also covered during sampling.
There were 23 species (13 families) of sh from
hook- and-line shing, which frequently caught the
surgeonsh Naso vlamingii and wrasse Oxycheilinus
diagramma.Only six sh species were common in the
117
Journal of Environmental Science and Management Vol. 20 No. 2 (December 2017)
Table 2. Species, estimated sizes (in cm), and counts of demersal (site-attached) and pelagic shes recorded during
ve days of sh visual census (FVC) and four days of hook-and-line shing (*) on the summit of the Benham
Bank Seamount, 6-14 May 2014. (cont.)
Family (Common Name)/ Species Size (cm) Classication Station
1 2 3 4 5
Halichoeres chrysus
Labroides bicolour
Oxycheilinus diagramma*
Thalassoma lutescens
Lethrinidae (emperors)
Gnathodentex aureolineatus*
Monotaxis grandoculis
Lutjanidae (snappers)
Aphareus furca
Lutjanus bohar*
Macolor macularisa
Mullidae (goatshes)
Parupeneus multifasciatusa
Scaridae (parrotshes)
Scarus forsteni
Serranidae (groupers)
Cephalopholis spiloparaea*
Cephalopholis urodetaa
Epinephelus fasciatus*
Gracila albomarginata
Plectropomus areolatus
Plectropomus laevisa
Pseudanthias tuka
Variola albimarginata*
Variola louti*
Synodontidae (lizardshes)
Synodus variegatus*
Carcharhinidae (requiem sharks)
Triaenodon obesus
Scyliorhinidae (cat sharks)
Atelomycterus marmoratus*
Scombridae (tunas)
Gymnosarda unicolor
Rhincodontidae (whale sharks)
Rhincodon typusb
12
6
17-28
12-18
18-23
15
18
24
25-36
18-20
16-18
17-21
15-18
20-30
20
72
80
7
25-38
24-42
16
150
70
150
400
Demersal; target
Demersal; target
Demersal; target
Demersal; target
Demersal; target
Demersal; target
Demersal; target
Demersal; target
Demersal; target
Demersal; target
Demersal; target
Demersal; target
Demersal; target
Demersal; target
Demersal; target
Demersal; target
Demersal; target
Demersal; target
Demersal; target
Demersal; target
Demersal; target
Pelagic, target
Pelagic, target
Pelagic, target
Pelagic, target
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
1
-
-
-
-
-
-
1
-
1
1
1
1
-
-
-
-
1
-
1
4
1
2
1
-
-
1
-
-
-
-
1
1
1
-
-
8
-
6
1
-
1
-
-
-
4
1
-
-
-
-
-
-
-
-
1
-
-
-
-
-
5
-
-
-
1
-
1
-
1
2
1
12
-
1
-
-
2
2
-
-
-
-
-
-
-
8
1
1
-
-
-
1
1
1
3
1
-
-
-
-
-
1
1
1
-
-
1
-
Number of species recorded:
Cumulative number of species recorded:
Number of unique species:
Cumulative number of unique species:
5
5
5
5
35
40
32
37
19
59
11
48
21
80
8
56
27
107
7
63
a Species was recorded during both FVC and hook and line fishing
b Species was recorded during the verification of bottom type and not during FVC or through hook-and-line
species lists from the shing events and FVCs, i.e.,
Heniochus monoceros (Chaetodontidae), N. vlamingii
(Acanthuridae), Macolor macularis (Lutjanidae),
Parupeneus multifasciatus (Mullidae), Cephalopholis
urodeta, and Plectropomus laevis (both Serranidae)
(Table 2).
Overall, the important observations for the reefs
on summit depths of the Benham Bank Seamount
with respect to reef shes were: that the diversity of
butteryshes (Chaetodontidae) was high, which is
consistent with the high species diversity of butteryshes
per unit area of reef observed in surveys conducted along
the Pacic Seaboard between 2001 and 2003 (DOST
2004); that despite the depth, the majority of the reef
sh species observed were commercially important food
sh, represented by large species of snappers, emperors,
groupers, trevallies, and surgeonshes; and that majority
of the shes observed were mostly adults or large-sized
individuals.
CONCLUSION AND RECOMMENDATION
The rst observations of the newly discovered
mesophotic reefs on the offshore underwater seamount
of the Benham Bank are presented here. The rapid
observational surveys revealed the pristine state of
the reefscapes, composed of mostly the foliose plate-
forming Porites (Synaraea) rus and at least 11 reef-
building and two solitary coral species that covered the
bottom, together with 63 reef-associated sh species,
conspicuous soft corals, arborescent sponges, and the
small-sized Halimeda. The apparent diversity observed
(corals, sh, macroalgae, and other invertebrates),
however, seemed lower than those in the nearshore and
shallower nging reefs (cf. Carpenter and Springer 2005,
Sanciangco et al. 2013). Further investigations on the
reefs’ biodiversity need to be performed to understand
its functioning and dynamics as an offshore habitat, and
to ascertain its connectivity with nearshore reefs (e.g.,
Van Oppen et al. 2011, Kahng et al. 2014, Thurber et
al. 2014, Abesamis et al. 2017). The Bank’s summit
depth at 50 m is currently used for anchorage during a
limited window (April to June each year) and may be
vulnerable to further shing pressure (cf. Koslow et al.
2000, Morato et al. 2006, Clark et al. 2010, Pitcher et al.
2010, Koslow and Couture 2013). Its sheries potential
must also be examined (White et al. 2007, Taranto et al.
2012) if shers were to be weaned from nearshore shing
or if the Bank (or part thereof) would be endorsed as a
no-take marine reserve and contribute to food security.
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ACKNOWLEDGMENT
The Department of Science and Technology (DOST)
and the Philippine Council for Agriculture, Aquatic
and Natural Resources Research and Development
(PCAARRD) funded the research program Exploration
and Assessment of Deep Water Areas in which the Joint
Cruise was a major activity. Thanks to the Department
of Agriculture-Bureau of Fisheries and Aquatic
Resources (DA-BFAR) for facilitating the approval
of the Gratuitous Permit and the opportunity to work
in the research platform M/V DA-BFAR with full
and steadfast support from the Captain, his crew, and
the research team. Sincere gratitude to the Marine
Science Institute for the diving and support equipment
provided; and the Marine Environment and Resources
Foundation (MERF), Inc. for nancial assistance. V.
Olivar inspected the air compressor units that served
the joint cruise.J. Rengel, Dr. A.T. Yniguez, and Dr.
O.C. Cabrera were volunteer divers; K.C.E. Pardo, R.
Arriesgado, and J.O.O. Nacorda assisted all divers upon
arrival onboard the M/V DA-BFAR; Dr. G.S. Jacinto
provided expert photos of the fresh specimens collected
and the Physical Oceanography Lab shared photos
taken by the kite. Thanks to K. Luzon who took whole
specimen and close-up photos of the bleached corals
collected, with assistance from J.R. Lapresca and T.J.
Estrella; Prof. N.B. Armada and Dr. S. Arakaki shared
fruitful discussions with Dr. C.L. Nanola, Jr. regarding
the identity of the pelagic tuna. Sincere gratitude to G.
Mapacpac, R. Coladilla, J. Mendoza, then Dean Dr.
L.M. Florece, and A. Dominguita for the administrative
support during the preparations for the cruise. Finally,
the Project acknowledges National Scientist and Prof.
Emeritus A.C. Alcala and Dr. S. Vergara for their review
and technical inputs in the research proposal.
121
Journal of Environmental Science and Management Vol. 20 No. 2 (December 2017)
... and 124°17=47.65ЉE (1). It is covered in clear waters, allowing sunlight to reach its depth, and was previously shown to host a diverse community of hard corals with associated reef fishes, sponges, and algae (1). ...
... (1). It is covered in clear waters, allowing sunlight to reach its depth, and was previously shown to host a diverse community of hard corals with associated reef fishes, sponges, and algae (1). Except for fishing within the narrow window of March to June every year, this unique offshore coral reef environment is naturally protected from further anthropogenic activities. ...
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We report here the draft genome sequences of six bacteria isolated from the near-bottom waters and surface sediments of the Benham Bank, Philippine Rise, Philippines. These genome sequences represent candidate novel species and/or strains from the families Flavobacteriaceae and Dermacoccaceae and the genera Idiomarina , Bacillus , and Vibrio .
... The Philippines is located within the Coral Triangle (CT), the area of highest coral reef biodiversity (Randall 1998;Roberts et al. 2002;Mora et al. 2003;Bellwood and Meyer 2009), and areas around its center are often identified as having the highest number of marine-fish species on Earth based on range-overlap analyses (Carpenter and Springer 2005;Allen 2008;Nañola et al. 2011;Sanciangco et al. 2013). Scientists have conducted fish collections of the shallow-water ichthyofauna in the Philippines for over a century (Herre 1953;Smith and Williams 1999), and extensive information about commercial fisheries and coral reef fish biodiversity for the region is widely available Niem 1998-2001;Randall 1998; Allen and Erdmann 2012; Motomura et al. 2017). Many hypotheses Topic Editor Morgan S. Pratchett Electronic supplementary material The online version of this article (https://doi.org/10.1007/s00338-019-01825-5) ...
... Recent studies in the Philippines have shown the importance of connectivity among different habitats for fishes that undergo ontogenetic habitat shifts and the fisheries that they support (Honda et al. 2013;Ramos et al. 2015). However, deeper ecosystems (depths [ 30 m) in this region have received less attention and very little information about the biodiversity of coral reefs at these depths is available (Nacorda et al. 2017;Turner et al. 2017;Abesamis et al. 2018;Joseph Quimpo et al. 2018). ...
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The Philippines is often highlighted as the globalepicenter of marine biodiversity, yet surveys of reef-asso-ciated fishes in this region rarely extend beyond shallowhabitats. Here, we improve the understanding of fish spe-cies diversity and distribution patterns in the Philippines byanalyzing data from mesophotic coral ecosystems (MCEs;30–150 m depth) obtained via mixed-gas rebreather divingand baited remote underwater video surveys. A total of 277fish species from 50 families was documented, whichincludes thirteen newly discovered and undescribed spe-cies. There were 27 new records for the Philippines and110 depth range extensions, indicating that many reeffishes have a broader geographic distribution and greaterdepth limits than previously reported. High taxonomicbeta-diversity, mainly associated with family and genusturnover with depth, and significant effects of traits such asspecies body size, mobility and geographic range withmaximum recorded depth, were observed. These resultssuggest that MCEs are characterized by unique assem-blages with distinct ecological and biogeographic traits. Ahigh proportion (60.5%) of the fish species are targeted byfishing, suggesting that Philippine MCEs are as vulnerableto overfishing as shallow reefs. Our findings support callsto expand conservation efforts beyond shallow reefs anddraw attention to the need to explicitly include deep reefsin marine protected areas to help preserve the unique bio-diversity of MCEs in the Philippines.
... The Philippines is located within the Coral Triangle (CT), the area of highest coral reef biodiversity (Randall 1998;Roberts et al. 2002;Mora et al. 2003;Bellwood and Meyer 2009), and areas around its center are often identified as having the highest number of marine-fish species on Earth based on range-overlap analyses (Carpenter and Springer 2005;Allen 2008;Nañola et al. 2011;Sanciangco et al. 2013). Scientists have conducted fish collections of the shallow-water ichthyofauna in the Philippines for over a century (Herre 1953;Smith and Williams 1999), and extensive information about commercial fisheries and coral reef fish biodiversity for the region is widely available Niem 1998-2001;Randall 1998; Allen and Erdmann 2012; Motomura et al. 2017). Many hypotheses Topic Editor Morgan S. Pratchett Electronic supplementary material The online version of this article (https://doi.org/10.1007/s00338-019-01825-5) ...
... Recent studies in the Philippines have shown the importance of connectivity among different habitats for fishes that undergo ontogenetic habitat shifts and the fisheries that they support (Honda et al. 2013;Ramos et al. 2015). However, deeper ecosystems (depths [ 30 m) in this region have received less attention and very little information about the biodiversity of coral reefs at these depths is available (Nacorda et al. 2017;Turner et al. 2017;Abesamis et al. 2018;Joseph Quimpo et al. 2018). ...
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The Philippines is often highlighted as the global epicenter of marine biodiversity, yet surveys of reef-associated fishes in this region rarely extend beyond shallow habitats. Here, we improve the understanding of fish species diversity and distribution patterns in the Philippines by analyzing data from mesophotic coral ecosystems (MCEs; 30–150 m depth) obtained via mixed-gas rebreather diving and baited remote underwater video surveys. A total of 277 fish species from 50 families was documented, which includes thirteen newly discovered and undescribed species. There were 27 new records for the Philippines and 110 depth range extensions, indicating that many reef fishes have a broader geographic distribution and greater depth limits than previously reported. High taxonomic beta-diversity, mainly associated with family and genus turnover with depth, and significant effects of traits such as species body size, mobility and geographic range with maximum recorded depth, were observed. These results suggest that MCEs are characterized by unique assemblages with distinct ecological and biogeographic traits. A high proportion (60.5%) of the fish species are targeted by fishing, suggesting that Philippine MCEs are as vulnerable to overfishing as shallow reefs. Our findings support calls to expand conservation efforts beyond shallow reefs and draw attention to the need to explicitly include deep reefs in marine protected areas to help preserve the unique biodiversity of MCEs in the Philippines.
... Although substantial work has been conducted in many parts of the world (reviewed in Turner et al., 2017), MCEs in the biodiverse Indo-Pacific are still under-studied (Kahng et al., 2010). Currently, there are only a handful of MCE studies in the Philippines (Ross & Hodgson, 1981;Abesamis et al., 2017;Nacorda et al., 2017;Quimpo et al., 2018aQuimpo et al., , 2018bCabaitan et al., 2019), the centre of marine biodiversity (Carpenter & Springer, 2005;Veron et al., 2009), with only three studies conducted in a few locations in the WPS biogeographic region (Ross & Hodgson, 1981;Quimpo et al., 2018aQuimpo et al., , 2018b, though none have focused specifically on comparing multiple locations within the WPS. ...
... Although substantial work has been conducted in many parts of the world (reviewed in Turner et al., 2017), MCEs in the biodiverse Indo-Pacific are still under-studied ( Kahng et al., 2010). Currently, there are only a handful of MCE studies in the Philippines (Ross & Hodgson, 1981;Abesamis et al., 2017;Nacorda et al., 2017;Quimpo et al., 2018aQuimpo et al., , 2018bCabaitan et al., 2019), the centre of marine biodiversity (Carpenter & Springer, 2005;Veron et al., 2009), with only three studies conducted in a few locations in the WPS biogeographic region (Ross & Hodgson, 1981;Quimpo et al., 2018aQuimpo et al., , 2018b, though none have focused specifically on comparing multiple locations within the WPS. ...
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The South China Sea (SCS) is a biodiversity hotspot, however, most biodiversity surveys inthe region are confined to shallow water reefs. Here, we studied the benthic habitat andfish assemblages in the upper mesophotic coral ecosystems (MCEs; 30–40 m) and SWRs(8–22 m) at three geographic locations (Luzon Strait; Palawan; and the Kalayaan Group ofIslands) in the eastern SCS (also called the West Philippine Sea) using diver-based surveymethods. Mean coral genera and fish species richness ranged from 17–25 (per 25 m2) and11–17 (per 250 m2) in MCEs, respectively; although none of these were novel genera/species.Coral and fish assemblages were structured more strongly by location than by depth. Locationdifferences were associated with the variability in benthic composition, wherein locations withhigher hard coral cover had higher coral genera richness and abundance. Locations withhigher algae and sand cover had higher diversity and density of fish herbivores and benthicinvertivores. Fishing efforts may also have contributed to among-location differences as thehighly exploited location had the lowest fish biomass. The low variation between depthsmay be attributed to the similar benthic composition at each location, the interconnectivitybetween depths due to hydrological conditions, fish motility, and the common fishinggears used in the Philippines that can likely extend beyond SWRs. Results imply that local-scale factors and anthropogenic disturbances probably dampen across-depth structuring incoral genera and fish species assemblages.
... MCEs may represent deep extensions of shallow reef systems, such as shelf-margin reefs around the Caribbean ( Goreau and Goreau 1973;Goreau and Land 1974;James and Ginsburg 1979;Liddell et al. 1997; Liddell and Ohlhorst 1988;Sherman et al. 2010). Alternatively, they may occupy more isolated banks with no direct connection to shallower systems, such as MCEs in the Gulf of Mexico ( Hine et al. 2008;Locker et al. 2010Locker et al. , 2016, Great Barrier Reef (GBR; Harris et al. 2013;Bridge et al. 2019), Northwest Australia ( Heyward and Radford 2019), and the Philippine Sea ( Nacorda et al. 2017). MCEs are often dynamic transition zones between shallow shelves and deep ocean basins in terms of biological communities, sedimentary facies, and physical oceanographic processes. ...
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Geomorphology and geological processes exert fundamental controls on the occurrence, distribution, and makeup of mesophotic coral ecosystems (MCEs). Two broad geomorphic categories are shelves and slopes. Shelves include outer portions of continental and insular shelves that dip gently into mesophotic depths before reaching the shelf break and have very low gradients (<1°). Other low-gradient habitats include tops of isolated banks. Slope habitats extend from platform breaks down into adjacent basins and can be divided into low-gradient slopes (<30°), steep slopes (~30 to 70°), and walls (>70°). On shelves, MCEs are best developed on positive relief features elevated above the surrounding seafloor. In slope settings, MCE development is typically favored on steep irregular slopes, where coral cover is concentrated on steep-sided buttresses and sediment is channelized into narrow chutes. Relict features related to past sea levels are critically important MCE habitats on both shelves and slopes. Coral and coralline algae remain the primary frame builders in MCEs. However, accretion at mesophotic depths is likely very slow, such that they form only thin biostromal veneers over relict substrates. Sediments in MCEs are dominantly autochthonous skeletal sands and gravels. Although fluxes of sediments to the seafloor in MCEs are typically lower than in shallow reefs, sedimentary dynamics still play an important role. Low-gradient seafloor has an increased potential for accumulation of sediment detrimental to MCEs. In slope settings, downslope bed-load transport of sediment can be orders of magnitude higher than vertical fluxes and likely exerts an important influence on MCEs.
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The study of age and growth of fish is essential for understanding their biology and population dynamics. Age and growth of Sargocentron spiniferum, from the Egyptian Red Sea at Shalateen fishing area were studied, depending on the otoliths’ readings using a non-linear backcalculation method. A total of 685 specimens (17.7–45.8 cm in TL) of S. spiniferum were aged and their maximum life span was 7 years. The Von Bertalanffy growth parameters for the investigated S. spiniferum specimens were: L∞ = 53.25 cm, K = 0.23 y-1, to = -0.66 years and the computed index of growth performance (Ø') was 2.81. The mortality rates were 1.04, 0.47 and 0.57 y-1 for total, natural and fishing mortality rates, respectively. The spiny squirrelfish fishery in Shalateen fishing area is working around its optimum situation, where the estimated exploitation ratio is 0.55. The obtained data from this study are the basic inputs of the analytical models used to achieve the wise management of this potential fishery.
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The study of age and growth of fish is essential for understanding their biology and population dynamics. Age and growth of Sargocentron spiniferum, from the Egyptian Red Sea at Shalateen fishing area were studied, depending on the otoliths’ readings using a non-linear backcalculation method. A total of 685 specimens (17.7–45.8 cm in TL) of S. spiniferum were aged and their maximum life span was 7 years. The Von Bertalanffy growth parameters for the investigated S. spiniferum specimens were: L∞ = 53.25 cm, K = 0.23 y-1 , to = -0.66 years and the computed index of growth performance (Ø' ) was 2.81. The mortality rates were 1.04, 0.47 and 0.57 y-1 for total, natural and fishing mortality rates, respectively. The spiny squirrelfish fishery in Shalateen fishing area is working around its optimum situation, where the estimated exploitation ratio is 0.55. The obtained data from this study are the basic inputs of the analytical models used to achieve the wise management of this potential fishery.
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Mesophotic coral ecosystems (MCEs) have received increasing attention in recent years in recognition of their unique biodiversity and also their potential importance as refuges from disturbance events. However, knowledge of the composition of MCEs and how they vary in space is lacking in many regions, particularly the Coral Triangle biodiversity hotspot. Here, we compared the benthic components and coral genera composition between shallow-water reefs (SWRs, 8–13 m depth) and upper MCEs (30–40 m) in four locations in the Philippines that are exposed to differing environmental conditions. Coral cover, abundance, and generic diversity were lower in MCEs than SWRs at three of the four locations. Benthic composition and coral generic composition also varied significantly among locations for both shallow and deep sites. Differences in benthic composition among sites was due primarily to variation in hard corals, macroalgae, sand and silt, while variation in coral assemblage was due to differences in abundance of encrusting Porites, branching Acropora, branching Seriatopora. Our results showed that the composition of MCE communities varied significantly from adjacent shallow reefs, but also among MCEs in differing geographic locations. Furthermore, our results suggest disturbances affecting shallow-water reefs, particularly sedimentation, also negatively impact MCEs, and that depth therefore provides no potential refuge from these disturbances. We recommend that conservation of MCEs consider spatial variability in community composition among sites, and urge further research to better understand the spatial variation in the composition of MCE communities in the Philippines.
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Baseline ecological studies of mesophotic coral ecosystems are lacking in the equatorial Indo-West Pacific region where coral reefs are highly threatened by anthropogenic and climate-induced disturbances. Here, we used baited remote underwater video to describe benthic habitat and fish assemblage structure from 10 to 80 m depth at Apo Island, a well-managed marine protected area in the Philippines. We conducted surveys 2 yr after two storms (in 2011 and 2012) caused severe damage to shallow coral communities within the no-take marine reserve (NTMR) of Apo Island, which led to declines in fish populations that had built up over three decades. We found that hard coral cover was restricted to < 40 m deep in the storm-impacted NTMR and a nearby fished area not impacted by storms. Benthic cover at mesophotic depths (> 30 m) was dominated by sand/rubble and rock (dead coral) with low cover of soft corals, sponges and macroalgae. Storm damage appeared to have reached the deepest limit of the fringing reef (40 m) and reduced variability in benthic structure within the NTMR. Species richness and/or abundance of most trophic groups of fish declined with increasing depth regardless of storm damage. There were differences in taxonomic and trophic structure and degree of targeting by fisheries between shallow and mesophotic fish assemblages. Threatened shark species and a fish species previously unreported in the Philippines were recorded at mesophotic depths. Our findings provide a first glimpse of the benthic and fish assemblage structure of Philippine coral reef ecosystems across a wide depth gradient. This work also underscores how a combination of limited coral reef development at mesophotic depths close to shallow reefs and severe habitat loss caused by storms would result in minimal depth refuge for reef fish populations.
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Recent collections of deep-sea majid crabs from the South Pacific Ocean and Taiwan provide new records of five species of Cyrtomaia Miers, 1886, and a new species from French Polynesia, C. polynesica n. sp. The news species is most similar to the recently described C. micronesica Richer de Forges & Ng, 2007, but differs from this species in the morphology of its carapace and pereopods.
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New collections of deep-sea crabs from the Bohol Sea in Central Philippines have obtained a large series of specimens of the deep-sea spider crabs of the genus Cyrtomaia (Majidae), of which one is here recognised as new, C. largoi, new species. Three other species: C murrayi Miers, 1886, C. horrida Rathbun, 1916, and C. echinata Rathbun, 1916, are all represented by an extensive series of specimens, allowing invaluable insights into their difficult taxonomy and ecology. One nominal subspecies, Cyrtomaia horrida pilosa Ihle and Ihle-Landenberg, 1931, is synonymised with C. horrida.