Mar. Freshwater Res.

, 2002, 53, 207–213

Long-term trends in yield and catch rates of the coral reef fishery
at Apo Island, central Philippines

A. P. MaypaAB, G. R. RussC, A. C. AlcalaA and H. P. CalumpongD

A
Silliman University, Angelo King Center for Research and Environmental Management,
Dumaguete City 6200, Philippines
B
Present address: Project Seahorse–Haribon Foundation, No. 12 Espina Compound, B. Rodriguez,
Cebu City 6000, Philippines. email: aimaypa@yahoo.com
C
School of Marine Biology and Aquaculture, James Cook University, Townsville, Qld 4811, Australia
D
Silliman University Marine Laboratory, Dumaguete City 6200, Philippines
eAYMtF.ia0Pl1Mad.a34yndpcatch ratesofacoar-lre fishery
A. PMaypa, G. R. us, A. C. Alacla nd H. PCalum pong

Abstract. Fish yields and catch rates recorded in the 1980s were compared with daily roving creel surveys
carried out in 1997/98, 2000 and 2001 at Apo Island. Total annual fish yields were measured six times over
the period 1980–2001. Total fish yield was 19–25 t km–2 year–1, with reef and reef-associated fish accounting for
15–20 t km–2 year–1, for five measurements. A sixth measurement, made in 1986, estimated 36.7 t km–2 year–1.
Annual yield remained stable over the study period. Carangidae and Acanthuridae accounted for 26–47% and
16–27% of the catch, respectively. Non-reef catches declined over time, from 6.21 t year–1 in 1980/81 to 1–2 t year–1
in 2000 and 2001. Estimates of annual hook and line catch per unit effort (CPUE) increased from 0.13–0.17 kg
man–1 h–1 in 1980/81, to 1–2 kg man–1 h–1 in 1997–2001. For target families, hook and line CPUE was consistently
higher in 1997–2001 than in 1980–86. However, hook and line CPUE for Carangidae and Acanthuridae declined
significantly between 1997 and 2001. Possible reasons for the long-term patterns of fish yields and catch rates are
discussed. Differences in methods used in estimates, and changes in gears and fishing effort over the years, make
comparisons difficult.

Introduction coralline shelf rather than small areas of actively growing

coral reef (Russ 1991). Early studies of reef-fish yields
In recent years, declines and collapses of marine fisheries have carried out in American Samoa (Hill 1978; Wass 1982) and
been attributed in large part to overfishing (FAO 1995; Pauly the Philippines (Alcala 1981; Alcala and Luchavez 1981),
and Christensen 1995; Vitousek et al.1997; Watson and Pauly documented yields in the range 8–27 t km–2 year–1 for
2001) and changes in the marine environment (Lauck et al. intensively fished, small areas of coral reef. The
1998). Fishing is the most important human activity sustainability of such yields was viewed with some doubt,
exploiting the resources of coral reefs (Russ 1991). Fish yields despite the study of Alcala (1981) at Sumilon Island,
decline in heavily exploited coral-reef areas (Munro 1983; Philippines, demonstrating consistently high yields for
Koslow et al. 1988; Russ 1991; Polunin and Roberts 1996). 5 years (1976–80). That study was expanded to demonstrate
Populations of large predatory fish have been depleted on a consistently high reef-fish yield over a period of 9 years
coral reefs owing to selective fishing (Munro 1983; Koslow (1976–85) (Alcala and Russ 1990). To our knowledge, no
et al. 1988; Russ and Alcala 1996a), and destructive fishing other long-term studies of reef-fish yields from individual
practices have caused severe damage to coral-reef habitats coral reefs have been published since. Thus, there are few
(Alcala and Gomez 1987; Gomez et al. 1987; McManus long-term data series of yield and catch rates from reef
1997). As a result, coral-reef fisheries worldwide are being fisheries of individual coral reefs.
studied in detail with resource management as a goal. In the Philippines, coastal communities rely on fish as a
Yields of coral-reef fisheries have been reviewed many source of animal protein and income, and 10–25% of the total
times (see Munro and Williams 1985; Russ 1991; Polunin fish biomass can be harvested from coral reefs annually
and Roberts 1996). Early reviews by Stevenson and (McManus 1988; Alcala and Russ 1990; Russ and Alcala
Marshall (1974) and Marshall (1980) concluded that yields 1998). Two of the highest reef-fishery yields recorded in the
ranged from 0.8 to 5 metric t km–2 year–1. Many of the data world come from Philippine coral islands, Sumilon and Apo,
from these reviews were based on studies from large areas of with annual yields of the order of 15–36 t km–2 year–1 (Alcala

© CSIRO 2002 10.1071/MF01134 1323-1650/02/020207

208
and Luchavez 1981; White and Savina 1987; Bellwood 1988;
Alcala and Russ 1990; Russ 1991). This paper reports on the
long-term trends of fish yields, catch composition and catch
rates of Apo Island. In addition, the paper documents yield,
catch rate and catch composition trends, before and over a long
period of establishment, of a successful marine reserve on Apo
Island (Russ and Alcala 1999). This ‘no take’ reserve, ~10%
of the coral-reef fishing area of Apo Island, was set up in 1982.
Materials and methods
Study site
The fish catch estimates were made at Apo Island, central Philippines
(9o4′N,123o16′E; Fig. 1) from 1980 to 2001. The island has an area of
0.74 km2, with 1.06 km2 of coral reef to the 60 m isobath (Russ and
Alcala 1996b). Only traditional, non-destructive, fishing methods are
allowed on the island: hook and line, gill-nets, spears, and bamboo
traps. A ‘no take’ marine reserve 400 m long lies along the south-
eastern portion of the island. It occupies ~10% of the coral-reef area
(Russ and Alcala 1999). This reserve has been protected since 1982
and Apo Island is known for its strict implementation of reserve
regulations (Russ and Alcala 1999).
Trends in yield and catch rates
Annual fish yields from various gear types and mean annual hook and
line catch per unit effort (CPUE) were estimated six times between
1980/81 and 2001. All fishers used hook and line, and this gear
Fig. 1. Apo Island, showing the fishing grounds and major fish
landing sites.
A. P. Maypa et al.
accounted for most of the catch during the period. Data for 1980/1981
were taken from Alcala and Luchavez (1981). These 1980/81 data were
recalculated, correcting for more accurate estimates of reef area and
mean days fished than were available in 1981. This allowed the data to
be compared more reliably with data from later studies. The CPUE for
1980/81 was the median of the reported range for hook and line
CPUE of Alcala and Luchavez (1981), since no mean was reported.
Data for 1985/86 were taken from White and Savina (1987). A
0.83 kg man–1 h–1 annual hook and line CPUE was reported, but no
standard error was provided. Fish yield for 1986 was based on
Bellwood (1988), but CPUE was not estimated in his study.
Fish yield estimates for 1997/98, 2000 and 2001 were based on data
gathered daily through roving creel surveys. Data for 1997/98 were
collected by field observers from June 1997 to September 1998, a total
of only 10 months. These data were extrapolated to an annual estimate,
and may have underestimated total yield. For the years 2000 and 2001,
field observers collected data from January to December. Catches of
reef fish and reef-associated fish from within the 60 m-isobath line
were recorded. Weight measurements of fresh fish were made by a
researcher (A.M.) and a fish dealer living on the island, trained to
identify most of the fish species using local names. Fish landed were
weighed on commercial weighing scales to the nearest 10 grams. When
fishers did not allow either length or weight measurements of their
catch to be taken, the fisher’s estimate of fish weight was recorded.
Catch surveys from 1997/98 to 2001 used fish landing forms which
recorded the following: name of fisher, date, fishing hours, fishing
ground, manpower, gear used, and fish species caught and the
corresponding weights. At Apo Island, fishers land their catch in many
different sites, making it difficult to record every fish landing. Three
major landing sites (Fig. 1) were therefore monitored. The estimated
annual yields for 1997/98, 2000 and 2001 were adjusted by adding 10%
to the actual values for both reef/reef-associated and non-reef catches to
account for this. This figure of 10% was based on the mean figure
given by fishers through interviews as the amount of catch not recorded
in surveys.
Identification of fish species followed that of Masuda et al. (1984),
Smith and Heemstra (1986), Randall et al. (1997) and Fish Base 2000
(Froese and Pauly 2000). Major fish groupings followed Bellwood
(1988).
Hook and line CPUE data in 1997/98 were gathered daily per
landing site at Apo Island. In 2000 and 2001 hook and line fishing data
were collected for three randomly chosen days in every week of the year
by drawing lots for three sampling days each week
Hook and line catch per unit effort (CPUE) at the family level
Hook and line is the dominant gear used in Apo Island. Since the
1980s, >90% of the Apo fishers have used hook and line, and >50%
of each reported total annual yield is caught by this gear. Thus, CPUE
of fish families targeted by fishers (mostly large predatory species)
using hook and line data were compared.CPUE data at the species level
were not available from 1980/81 (Alcala and Luchavez 1981), 1985/86
(White and Savina 1987) and 1986 (Bellwood 1988). Thus, only data
from 1997/98, 2000 and 2001 were analysed. A one-way ANOVA was
used to test for significant differences in CPUE between years at the
level of fish family. These families were Carangidae, Lethrinidae,
Lutjanidae and Serranidae (Epinephelinae) and Acanthuridae.
Although most acanthurids are planktivores, their contribution to the
annual yield was as high as 30% of the catch. All data were tested for
normality and homoscedasticity using Kolmogorov’s test and Levene’s
test, respectively (Velleman 1997). A Box–Cox transformation
normalized and homogenized the following data sets: Carangidae
(exponent, e = – 0.0287), Acanthuridae (e = – 0.0402), Lethrinidae
(e = – 0.5115). Data sets for Lutjanidae and Serranidae required no
transformations. A Bonferroni test was used for post hoc analysis.
Yield and catch rates of a coral-reef fishery
Results
Yield and catch composition
From 1980 to 2001, three forms of fishing gears were used
consistently at Apo Island: hook and line, gill-nets and
spears. Bamboo fish traps were also popular over the study
period, but were not used in 2000, because tourist divers had
deliberately damaged traps. By 2001, 12 traps were again
deployed in the Apo Island fishing grounds. For >20 years,
Apo Island has maintained fish yields of 19–37 t km–2 year–1,
of which 15–30 t km–2 year–1 were reef and reef-associated
fish (Fig. 2). However, for five of the six measurements of
yield (excluding that of White and Savina 1987), total yield
ranged from 19 to 25 t km–2 year–1, with reef and reef-
associated fish accounting for 15–20 t km–2 year–1. There
was a decline of non-reef catch late in the study from
6.21 t km–2 year–1 (25%) in 1980/81 to 1–2 t year–1 (9%) in
the years 2000 and 2001 (Fig. 2, Table 1).
The catch composition of the Apo Island fishery is
dominated by a few fish families (Table 1), with five families
accounting for about 50–90% of the catch. Reef and reef-
associated species became more dominant in the catch over
the years. In particular, an increasing proportion of
carangids and acanthurids was observed in the catch over
time. The proportion of each of these families in the total
40
Total annual yield
t km-2 year -1
30
reef/reef-associated
non-reef
20
10
0 two decades. The reef-fish component has consistently been
2 CPUE
kg man -1 h-1
1.5
1 yields from coral reefs (Munro and Williams 1985; Russ
0.5
0 made by Alcala (1981) and Alcala and Russ (1990), who
1980/1981 85/86 97/98 2000
Year
Fig. 2. Trends in total annual fish yields and annual hook and line
catch per unit effort (CPUE: 1981 = median; 1985/86 = mean, no
s.e.; 1997/98, 2000, 2001 = mean ± s.e.) from Apo Island, central
Philippines (data plotted from 1980/81, 1985/86 and 1986 are from
Alcala and Luchavez 1981, White and Savina 1987 and Bellwood
1988, respectively).
209
catch almost doubled over the study period (Carangidae:
26% in 1980/81 to 47% in 2000/2001, Acanthuridae: 16% in
1980/81 to 27% in 2000/2001). About 90% of the carangid
catch at Apo (Caranx ignobilis, C. melampygus,
C. sexfasciatus, Elagatis bipinnulata) were caught by hook
and line gear. Larger acanthurids (e.g. Acanthurus bleekeri,
Naso hexacanthus, N. lopezi, N. vlamingii) were also caught
by hook and line. Those species with smaller-bodied
individuals (mostly Naso minor, N. thynnoides) were caught
by gill-nets.
Catch rates
Annual catch per unit effort (CPUE) for hook and line
increased from 0.13–0.17 kg man–1 h–1 (0.15 kg man–1 h–1,
median) in 1980/81, to 1–2 kg man–1 h–1 in 1997–2001
(Fig. 2). Statistical comparisons were not possible, because
raw data were unavailable from the earlier studies of Alcala
and Luchavez (1981), White and Savina (1987) and
Bellwood (1988).
Hook and line CPUE values for Carangidae and
Acanthuridae in 1997/98 were significantly higher than
those for 2000 and 2001 (ANOVA: PCarangidae <0.001,
PAcanthuridae <0.01; Table 2, Fig. 3). CPUE declined over the
period 1997–2001 for both families. In 1997/98, hook and
line CPUE of Carangidae was 2.06 ± 0.01 kg man–1 h–1. This
had declined significantly (ANOVA: P <0.001) in 2001 to
1.34 ± 0.12 kg man–1 h–1. Similarly, hook and line CPUE of
Acanthuridae declined significantly (ANOVA: P <0.01)
from 1.64 ± 0.29 kg man–1 h–1 in 1997/98 to 1.05 ± 0.13 kg
man–1 h–1 in 2001. No significant differences were detected
between years for hook and line CPUE of Lethrinidae,
Lutjanidae and Serranidae (Epinephelinae).
Discussion
This study has demonstrated that fish yields of 19–25 t km–2
year–1 have been maintained at Apo Island, Philippines, for
15–20 t km–2 year–1. One measurement (White and Savina
1987) was considerably higher (total yield 36.7 t km–2 year–1,
with reef fish yield of 31.8 t km–2 year–1) than the other five
made during the study. The data from Apo Island over the
period 1980–2001 are some of the highest recorded fishery
1991; Polunin and Roberts 1996). Such high yields of reef
fish from small areas of actively growing coral reef are
clearly sustainable in the long term. The same point was
reported that Sumilon Island, a coral reef near Apo Island,
yielded 15–36 t km–2 year–1 of reef/reef-associated fish
during 1976–86. Hill (1978) and Wass (1982) also reported
yields of 8.6–40.4 t km–2 year–1 of fish from reefs around
American Samoa. The high yields from Apo and Sumilon
Islands consist of large proportions of planktivorous fish or
fish that prey heavily on planktivorous fish (e.g. Carangidae
210 A. P. Maypa et al.

Table 1. Major fish families comprising the annual catch caught by all gear types at Apo Island, central Philippines from 1980/81 to 2001
TY, total yield; TSY, total sample yield; NR, not reported. 1980/81, 1985/86 and 1986 data are from Alcala and Luchavez (1981), White and Savina
(1987) and Bellwood (1988), respectively

Family (major family/ 1980/81 1985/86 1986 1997/98 2000 2001

genus) %TY TSY %TY TSY %TY TSY %TY TSY %TY TSY %TY TSY
(kg) (kg) (kg) (kg) (kg) (kg)

Reef/Reef associated
Carangidae 26.1 828 23.32 3783 39 9690 29.6 6460.22 46.31 9025.8 46.64 8596.73
(Caranx)
Acanthuridae 16.4 519 23.83 3866 23.6 5870 27.4 5963.63 29.43 5737.1 24.53 4522.82
(Naso)
Caesionidae 15.2 483 9.97 1617 3.46 860 1.91 416.09 8.91 1737 3.95 728.78
(Caesio)
Cephalopoda 3.72 118 2.47 400 2.98 740 0.55 120.2 1.0 195.9 1.48 272.9
(Octopus)
Scaridae 2.85 90.6 2.79 453 3.34 830 0.18 40.28 3.44 671.06 6.68 1230.43
(Scarus)
Lethrinidae 1.2 38 0.1 17 NR 0.11 24.2 0.46 89.85 0.21 38.1
(Lethrinus)
Lutjanidae 0.65 20.5 2.62 425 5.75 1430 3.95 858.27 2.89 561.27 2.33 428.93
(Lutjanus)
Serranidae,Epinephelinae 0.36 11.2 0.57 92 NR 0.21 45.89 0.89 174.29 0.5 92.12
(Cephalopholis/
Epinephelus)
Others 7.7 244 4.2 682 1.17 290 1.89 412.2 1.76 343.13 0.93 169.78
Total (kg) 2352 11335 19710 14341 18535 16080.59
Total (t km-2) 2.39 10.7 18.59 13.53 17.48 15.17
Estimated annual yield 17.86 31.8 19.72 14.88 19.23 16.69
(t km–2 year–1)
Non-reef 25.8 819 30.13 4886 20.7 5150 34.2 7460.51 4.91 956.44 12.75 2350.8
(Scombridae)
Total (t) 0.82 4.9 5.15 7.46 0.96 2.35
Estimated annual yield 6.21 4.9 5.15 8.2 1.05 2.4
(t km–2 year–1)

at Apo). Thus, the high yields may be sustainable in the long studies of reef fish yields from Sumilon and Apo Islands are
term, particularly if there is likely to be substantial input of some of the longest time-series of fisheries yield and catch
plankton onto the reef from outside. Since strong currents rates for any coral reefs in the world.
flow onto both islands from the north, this is likely. These Over the period 1980–2001 the fisheries yield of both
reef/reef-associated fish and non-reef fish from Apo Island
has remained stable, particularly if the very high estimate of
Table 2. Results of one-way ANOVA of hook and line catch per White and Savina (1987) is excluded. The composition of
unit effort of reef/reef-associated target families of fish between the catch has changed slightly, with reef fish such as
years (1997/98–2001) at Apo Island, central Philippines Carangidae and Acanthuridae both almost doubling as a
NS, not significant proportion of the catch, and fewer non-reef fish
Family One-way ANOVA Bonferroni post hoc (Scombridae, Elopidae) landed in 2000/01. In addition, the
(α = 0.05) catch per unit effort of hook and line fishing, the main gear
F P used at the island over the two decades, has increased
Carangidae 19.68 ≤0.00015 1997/98 > 2000 > 2001
substantially between the 1980s and the period 1997–2001.
Acanthuridae 6.10 0.0027 1997/98 > 2000 = 2001 With catch stable, and catch rate of the main gear increasing
Lutjanidae 2.59 0.0824 NS substantially, fishing effort of the fishing community has
Serranidae 1.36 0.2783 NS clearly declined over the two decades.
(Epinephelinae) The trends in yield and catch rates reported here are
Lethrinidae 0.70 0.5184 NS subject to a range of potential limitations and influences.
Yield and catch rates of a coral-reef fishery
4 do not preclude useful inferences about temporal changes in
Acanthuridae
Carangidae
Lutjanidae
Serranidae
Lethrinidae
kg man -1 h -1
3
2 consequence of negative impacts of tourism. Fishers at Apo
1 using traps in the year 2000 because diving tourists were
0
1997/98 2000 2001
Year
Fig. 3. Trends in catch per unit effort of target families of fish at
Apo Island, central Philippines.
Fishing effort by the fishers has declined over the study,
influencing comparisons of catch over time. Fishing gears
used by fishers have changed slightly over the period,
although hook and line fishing has remained the principal
fishing gear. The decline of non-reef catch from 1980/81 to
2000/01 coincided with less frequent use of drift gill-nets
(locally known as ‘pamo’) which target non-reef species such
as elopids and scombrids. Such drift gill-nets were not used
after 1999/2000 (L. Pascobello-Rhodes, Apo Island resident,
personal communication). Fish traps, a common fishing gear
in the period 1980–86 (Alcala and Luchavez 1981; White and
Savina 1987; Bellwood 1988) were not used at all during
2000, but were reintroduced into the fishery in 2001.
Furthermore, slight variations in methods used to collect
data in 1980/81 (Alcala and Luchavez 1981), 1985/86 (White
and Savina 1987), 1986 (Bellwood 1988), 1997/98
(Calumpong, present paper) and 2000/2001 (Maypa, Alcala,
Russ, present paper) may have influenced comparisons of
yield and catch rates. The yield estimate of White and Savina
(1987) for 1985/86 was 65% higher than the mean of the
other five measurements made over the study period. This
may represent natural variation from year to year in yields,
although the study of Bellwood (1988) also collected data in
1986, and large variations were not observed between years
for the other five measurements. White and Savina (1987)
recorded 11.33 t of reef fish landed over a year, but expanded
this by 200% (to 31.8 t) to account for local consumption and
the practice of drying fish for sale/consumption. It is possible
that they overestimated the significance of local consumption
and fish drying. For example, Alcala and Luchavez (1981)
estimated that local consumption accounted for 8% of yield.
All studies except that of Bellwood (1988) used fisher
interviews and a creel survey approach on the island.
Bellwood estimated fish catch from records of weekly fish
landings at a fish market on the mainland, near Apo Island.
Finally, annual yields and catch rates will clearly vary with
environmental changes. However, these variations over time
211
fishery yields and CPUE.
The significant decline in hook and line CPUE of
carangids and acanthurids from 1997/98 to 2001 may
indicate a decrease in fish standing stocks and/or may be a
Island have suggested recently that tourists are disturbing
their fishing by diving on fishing grounds. Fishers ceased
deliberately damaging traps. Bernardo (2001) recorded
several indicators that suggested that Apo Island has
exceeded its carrying capacity in terms of tourism, and he
recommended the implementation of open and closed
seasons for diving in fishing grounds.
This study documents fishery yields and catch rates
before, during and almost two decades after establishment of
one of the most successful small ‘no-take’ marine reserves in
the world (Russ and Alcala 1996a, 1996b, 1998, 1999). One
expectation of marine reserves as fisheries management
tools is that they may eventually become net exporters of
adult fish biomass (Russ 2002), the so-called ‘spillover
effect’ (Alcala and Russ 1990; Russ and Alcala 1996b;
McClanahan and Mangi 2000; Roberts et al.2001). The
evidence for spillover from marine reserves has been
reviewed recently by Russ (2002). The results presented in
this paper are consistent with, but insufficient to establish
unequivocally, spillover. The results suggest that the fishing
community on Apo Island has benefited from the presence
of the marine reserve despite exclusion of fishers from 10%
of the fishing grounds around Apo Island in 1982. First, the
local economy has benefited, perhaps by US$500 per hectare
of reef per year (Alcala 1998), from tourism generated by the
reserve, with two tourist resorts established on Apo Island
during the 1990s (Russ and Alcala 1999). Secondly, fishing
effort has declined, perhaps because of higher catch rates and
additional income from tourism (Russ and Alcala 1999).
Thirdly, fishers appear to have changed their fishing
patterns. They travel off-reef to catch scombrids and elopids
with drift gill-nets far less frequently because reef fish may
be far more available. The increase in both acanthurids and
carangids d from 40% to >70% of the catch between 1980/
81 and 2000/01 supports this contention.
A large proportion of fish are captured from a small area of
reef in the Cogon area of Apo Island, a considerable distance
from the reserve. This is, a region of considerable current,
and probably considerable input of plankton (Bellwood
1988). The availability of Cogon as a fishing site (during
May–October, the south-west monsoon) appears to be an
important determinant of the size of fish yields at Apo Island
(Bellwood 1988), which makes interpretation of the role of
the reserve in affecting yields more problematic.
Other evidence is needed to make a convincing case that
the Apo reserve has benefited the local fishery by spillover.
Firstly, evidence that reef fish species that have increased
212
proportionally in the catch over the study period
(e.g. acanthurids, carangids) have increased in abundance in
the reserve over the period. Secondly, that these species have
also increased in abundance in those fished areas closest to
the reserve (see Russ and Alcala 1996b). Thirdly, that catch
rates of such species are higher closer to the reserve than
further away. Fourthly, that such species have a net
movement out of the reserve (measured by tagging studies).
For example, it is possible that mobile predators such as
carangids are attracted to the reserve by high abundances of
reef-fish prey. In this sense, the reserve could act as an
attractor, somewhat like an artificial reef, and enhance catch
rates of carangids near the reserve boundary.
Acknowledgments
The financial support for this study was provided by the Pew
Fellows Marine Conservation Program to A.C.A. and G.R.R.
USAID through Silliman University COE also provided
funds for the 1997/98 study. Special thanks to J. L. Maypa
for generating the map. We are grateful to B. Abrenica,
R. Aldeon and family, L. Rhodes, P. Rhodes and all the Apo
Island people for making this study possible. Thanks also to
L. J. Raymundo, whose comments improved this paper.
Presentation of this study to the 6th Indo–Pacific Conference
was funded by Pascual Foundation, DA-Bureau of
Agricultural Reasearch, Philippines through E. Ponce and
UP Los Baños Foundation and the 6th IPFC funders through
L. Beckley. The help of M. Maguyon in facilitating the
release of travel funds is appreciated.
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