Philippine Journal of Science
141 (2): 243-252, December 2012
ISSN 0031 - 7683
Date Received: 03 May 2012
An Updated Taxonomic Account of Limnetic Crustacean
Zooplankton in Lake Taal, Philippines
Rey Donne S. Papa1, 2, 3* Dino T. Tordesillas2, 5, and Augustus C. Mamaril, Sr.4
1
Department of Biological Sciences, 2Graduate School and
3
Research Center for the Natural and Applied Sciences,
University of Santo Tomas, Manila, Philippines
4
Institute of Biology, University of the Philippines - Diliman, Quezon City, Philippines
5
Department of Biology, St. Paul University, Quezon City, Philippines
Limnetic crustacean zooplankton are the preferred prey of the economically important and
endemic zooplanktivore Sardinella tawilis (Clupeidae) of Lake Taal. In this paper, we update
the species composition, morphology and distribution of limnetic crustacean zooplankton in
Lake Taal based on samples collected from 2008 through 2010. A total of nine species belonging
to Copepoda (3 spp.) and Cladocera (6 spp.) have been documented including Arctodiaptomus
dorsalis, a Neotropical species and Pseudodiaptomus brehmi, which was previously thought to
be restricted to Lake Naujan, Mindoro Is. Information on these relatively understudied taxa
is an important contribution to the on-going re-assessment of Lake Taal biodiversity in the
light of aquaculture and eutrophication impacts, as well as the presence of introduced species.
Key Words: Crustacean Zooplankton, Freshwater, Southeast Asia, Systematics, Tropical caldera lakes,
INTRODUCTION
Studies on Philippine zooplankton were formally started
by the description of Trochosphaera aequatorialis in 1872
based on samples collected from a rice field in Mindanao
Island in southern Philippines (De Elera 1895; Mamaril Sr. &
Fernando 1978; Petersen & Carlos 1984). This was followed
by the first accounts of zooplankton species richness during
the Wallacea expedition (Brehm 1938; Brehm 1942;
Hauer 1941; Kiefer 1930; Kiefer 1939a; Kiefer 1939b;
Woltereck et al. 1941). This likewise included surveys of
lakes such as Laguna de Bay, Taal, Balinsasayao, Buhi,
Calibato, Danao, Lanao, Mainit, and Naujan. Measurements
on some basic physico-chemical and morphological
characteristics of the aforementioned lakes were also
carried out. New species of crustacean zooplankton such
as Alona pseudoanodonta Brehm, Diaptomus insulanus
Wright and its synonym D. sensibilis Kiefer, D. vexillifer
*Corresponding author: [email protected];
[email protected]
Brehm, Mesocyclops hyalinus Kiefer, Pseudodiaptomus
brehmi Kiefer, Tropodiaptomus gigantoviger Brehm, T.
prasinus Brehm, and Thermocyclops wolterecki Kiefer
have been described based on the samples collected from
the expedition. New records have also been established
for many cladoceran species.
The most comprehensive publication on Philippine
freshwater zooplankton systematics reported 125
zooplankton species including 49 cladocerans and nine
copepods collected from the littoral and limnetic zones
of lakes and reservoirs as well as smaller water bodies
in the Philippines (Mamaril Sr. & Fernando 1978). This
together with two more papers by Mamaril Sr. (1986;
2001) became the most utilized references on Philippine
freshwater zooplankton. At present, a website maintained
by F. Petersen of Denmark also serves as an invaluable
on-line reference for those who want to study Philippine
freshwater zooplankton taxonomy (Petersen 2009). Other
significant studies on Philippine freshwater zooplankton
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Philippine Journal of Science
Vol. 141 No. 2, December 2012
Papa et al.: Taxonomic Account of Limnetic
Zooplankton in Lake Taal, Philippines
Figure 1. Line drawings of limnetic crustacean zooplankton species in Lake Taal A) Diaphanosoma tropicum B) Diaphanosoma excisum
C) Diaphanosoma sarsi D) Ceriodaphnia cornuta E) Moina micrura F) Bosmina fatalis G) Pseudodiaptomus brehmi (female)
H) Arctodiaptomus dorsalis (female) I) Thermocyclops crassus (female). Scale bars: A-C, E, G, H (200 μm); D, F & I (20 μm).
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Papa et al.: Taxonomic Account of Limnetic
Zooplankton in Lake Taal, Philippines
Philippine Journal of Science
Vol. 141 No. 2, December 2012
4 μm
60 μm
Figure 2. Scanning electron micrographs of selected morphological characters of female Thermocyclops crassus A) Cephalon showing
the antennae and mouthparts B) Right fifth thoracic leg (P5). Legend: A1 – antennule, A2 – antenna, R - rostrum La – labrum,
Md – mandible, Mxl – maxillule, Mx – maxilla, Mxp – maxilliped.
were done by Cheng & Clemente (1954); Ueno (1966),
Lewis (1979); Lai et al. (1979); Petersen & Carlos (1984);
Tuyor & Segers (1999); Tuyor & Baay (2001); WALTER
et al. (2006); Aquino et al. (2008); Lazo et al. (2009); and
Papa et al. (Papa & Zafaralla 2011; PAPA et al. 2011;
Papa et al. 2012). They have either discussed taxonomic
revisions, new records or ecological aspects.
Lake Taal (13°55’-14°05'N, 120°55’-121°105'E; Altitude:
2.5 masl) has long caught the attention of scientists for
its unique geological origin, geographical location and
limnological characteristics. The lake is a 236.9 km2
caldera with 90.4 m mean and 198 m maximum depths,
respectively. It was formed after a series of volcanic
explosions separated the lake from the rest of the South
China Sea (Ramos 2002) with a shallow north and a
deeper south basin partially separated by an active volcano
island in the middle. Since 1975, fish cage aquaculture
has proliferated in the north basin which has begun to
take its toll on water quality (Aypa et al. 2008). The
decline in ecosystem health due to this has prompted
renewed interest in assessing the lake’s biodiversity (TvplPamb 2009). As an important component of freshwater
ecosystems, zooplankton diversity needs to be updated to
keep up with the recent developments in Systematics. The
zooplankton composition of Lake Taal was first studied
during the Wallacea expedition (Woltereck et al. 1941)
which was later updated by Ueno (1966) and Mamaril
Sr. (2001). A report on the limnological status of Lake
Taal also presented some data on zooplankton species
composition during the late 1980s (Zafaralla et al. 1992;
Zafaralla et al. 1989). This paper updates the crustacean
zooplankton species richness in Lake Taal using specimens
collected during limnetic plankton surveys from 2008
to 2010. We describe key morphological features by
examining collected samples using a combination of
light and scanning electron microscopy. We also included
observations on their abundance and distribution.
MATERIALS AND METHODS
Plankton were collected from six sites in Lake Taal by
making replicate vertical tows from a depth of 40 m using
an 80 µm mesh size conical plankton net (diameter = 30
cm). Samplings were held monthly in 2008, four times
in 2009 (May, June, August, and November) and twice in
2010 (April). Samples were fixed in 5 % formalin with
Rose Bengal dye. Density was determined for each species
by taking average counts of three 1 mL Sedgwick Rafter
sub samples per replicate. Specimens were sorted and
dissected on slides with glycerine. These were examined
using an Olympus CX21 compound microscope and drawn
with the aid of an Olympus U-DA drawing attachment.
All measurements were made using a calibrated ocular
micrometer and presented in millimetre(s). Representative
individuals from the copepod were dehydrated in ethanol
series, critical point dried in a BALTEC CPD 030, fixed
on SEM aluminum stubs, sputter coated with 5 nm Au/Pd
in a BALTEC SCD 030 and viewed in a Philips SEM 505.
Pictures were taken with a DISS (Digital Image Scanning
System) from Point Electronics. Body and appendage
terminology was based on Dussart and Defaye (2001)
and Korinek (2002). Specimens (sorted or otherwise) are
deposited in the Zooplankton Reference Collection of the
University of Santo Tomas.
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Philippine Journal of Science
Vol. 141 No. 2, December 2012
Papa et al.: Taxonomic Account of Limnetic
Zooplankton in Lake Taal, Philippines
Figure 3. Scanning electron micrographs of selected morphological characters of male (A & B) and female (C & D) Arctodiaptomus dorsalis.
A) Terminal segment of right antennule B) Exopod of fifth leg (P5) C) Right leg (P5) D) Endopod of P5.
RESULTS
Class Maxillopoda
Order Cyclopoida
Family Cyclopidae
Thermocyclops crassus (Fischer, 1853)
This is the type species for the genus Thermocyclops. Of
the more than 50 recognized species under this genus, only
T. crassus has a cosmopolitan distribution. Though several
studies have pointed out the presence of at least two cyclopoid
species in Lake Taal, our collections have only yielded this
species, at least for the limnetic zone. Female T. crassus has a
hammer-shaped seminal receptacle with a short and wide saclike handle. It may be distinguished from the morphologically
similar T. decipiens by its straight spine in the inner distal
segment of the P4 endopod, as well as the presence of hairs
instead of spinules on the medial margin of the P4 basipodite
246
(Alekseev 2002; Dussart & Defaye 2001). The female P5
has an apical seta and a setiform spine inserted apically (Fig.
2B). Previous studies on Lake Taal zooplankton have listed T.
crassus as T. hyalinus (Kiefer 1930; Woltereck et al. 1941).
At present, T. crassus dominated the two other copepod
species in Lake Taal and is also common to other sampling
localities in the Philippines. Mean lengths of T. crassus was
0.49 mm for males and 0.58 mm for females. Specimens of
adult, copepodite and nauplii T. crassus were observed from
samples collected throughout the year.
Order Calanoida
Family Diaptomidae
Arctodiaptomus dorsalis (Marsh, 1907)
A. dorsalis is the largest copepod species in Lake Taal
with average length of 0.97 mm for females and 0.82 mm
for males. One key characteristic of female A. dorsalis is
Philippine Journal of Science
Vol. 141 No. 2, December 2012
Papa et al.: Taxonomic Account of Limnetic
Zooplankton in Lake Taal, Philippines
Figure 4. Scanning electron micrographs of selected morphological characters of female (A-C) and male (D) Pseudodiaptomus brehmi
from Lake Taal A) Seminal receptacle B) Last two thoracic segments (T4 and T5) C) Last two urosome segments (Ur4 and Ur5)
D) Terminal segments of right antennule.
that its P5 has an endopod that is nearly as long as the
exopodite 1 (Fig. 3C & D) while for males, the P5 has a
lateral spine inserted proximal to the middle of the right
of exopodite 2 where the said spine is longer than the
exopodite 2 (Fig. 3B).
The Wallacea papers did not mention any calanoid
copepods in Lake Taal. The occurrence of a calanoid
initially identified as Tropodiaptomus vicinus (Zafaralla
et al. 1989; Zafaralla 1992 cf Mamaril Sr. 2001) was
reported in Lake Taal and later mentioned in Amarasinghe
et al. (2008), Vijverberg et al. (2008) and Papa and
Zafaralla (2011). However, a more detailed examination
of specimens collected from 2008 to 2010 revealed that
the diaptomid copepods in Lake Taal were A. dorsalis and
not T. vicinus. This was also mentioned by PAPA et al.
(2012) in their report on the invasion of A. dorsalis which
they found in18 of 27 lakes throughout the archipelago.
Similar to T. crassus, adults, copepodites and nauplii of A.
dorsalis were also observed all year round in Lake Taal.
The adults were not as abundant as those of T. crassus as
calanoid copepods are known to be less numerous than
cyclopoids in eutrophic lakes.
Family Pseudodiaptomidae
Pseudodiaptomus brehmi Kiefer, 1939
Pseudodiaptomus is classified as a highly diversified
demersal marine genus in the Indo-Pacific region with
77 species, 14 of which are recorded from coastal
environments in the Philippines (Walter et al. 2006).
P. brehmi was first described by Kiefer in 1939 based
on samples collected from Lake Naujan (Mindoro Is.,
Philippines). At present, P. brehmi is listed as a marine
species (Walter 2009) and until this study, no record of P.
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Philippine Journal of Science
Vol. 141 No. 2, December 2012
brehmi in another freshwater environment exists. Petersen
has previously classified it as endemic to Lake Naujan
(Petersen 2009). Further studies have to be undertaken
to ascertain its distribution. Female specimens are
characterized by the absence of spines in the first urosome
segment (Fig. 4B) while the male P5 has no medial pointed
process in its left Re1. Specimens from Lake Taal are
rare even for samples collected from deeper locations
for almost the entire sampling period except for those
collected in April 2010 when they were fairly common.
Class Branchiopoda
Order Cladocera
Family Bosminidae
Bosmina fatalis Burkhardt 1924
This species is common in lakes and reservoirs in East
(including far eastern Russia) and Southeast Asia. In the
Philippines, it has been reported from Luzon Is. including
La Mesa reservoir and Laguna de Bay and in Mindanao
Is. in Lakes Lanao (Mamaril Sr. 1986) and Sebu (Papa
unpublished). Bosmina spp., (B. fatalis and B. longirostris)
were first reported from Lake Taal in 2001 (Mamaril Sr.
2001). Another study only found B. fatalis in this lake
(Vijverberg et al. 2008). Research on the prey preference
of S. tawilis from 2004 to 2005 showed that Bosmina spp.
featured prominently in its diet during the northeast monsoon
coinciding with a drop in copepod abundance (Papa et al.
2008). B. fatalis is known for its rounded carapace with
a long rostrum with a pair of fixed antennules of variable
length and shape curving ventrally. Lake Taal specimens
had mean lengths of 0.39 mm. Monthly plankton surveys
in 2008 revealed that B. fatalis was second to Ceriodaphnia
cornuta in terms of abundance and were present throughout
the entire year with highest densities in January.
Family Moinidae
Moina micrura Kurz, 1874
The cosmopolitan Moina micrura is found in a variety of
habitats from ponds, rice fields, reservoirs and lakes (Mamaril
Sr. 1986). Species from this genus together with those from
Diaphanosoma are considered as the replacement of Daphnia
in the tropics and are the preyed upon by invertebrate
predators (Mamaril Sr. 2001). M. micrura was first reported
from Lake Taal in 1938 as M. dubia (Brehm 1938) which is
now considered as its junior synonym. Known for having
a large head with a well-developed supraocular depression,
it can be differentiated from other members of this genus
by the absence of hairs on the ventral surface of its head
(Korinek 2002). In terms of body size, M. micrura would be
the second largest cladoceran genus in the lake at 0.56 mm
total length. M. micrura were commonly encountered from
the samples collected in Lake Taal though never as abundant
as C. cornuta or B. fatalis.
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Papa et al.: Taxonomic Account of Limnetic
Zooplankton in Lake Taal, Philippines
Family Daphniidae
Ceriodaphnia cornuta Sars, 1885
This cladoceran is common to tropical and subtropical
regions and is usually found in both the littoral and
limnetic areas of lakes but is absent in running waters. The
first recorded occurrence of C. cornuta in the Philippines
was from the Wallacea papers which included Lake Taal
where it was listed as C. rigaudi. It is characterized by
its pointed rostrum and small head which sometimes
contains a horn-like projection. The carapace is nearly
circular and has a deep cervical sinus (Korinek 2002).
It is a highly polymorphic species, wherein the hornless
and horned forms appear depending on lake productivity
and intensive predation by fish, respectively (Zaret 1972).
Lake Taal specimens had average lengths of 0.38 mm. The
specimens did not possess horns, which may be due to
the high productivity of the lake from the excess nutrient
inputs from aquaculture as well as natural nutrient sources
from the volcanic sediments. Though found in the diets of
S. tawilis it is not considered as a preferred prey item in
spite of its high densities in the lake (PAPA et al. 2008).
It was present in samples collected from all months.
Family Sididae
Diaphanosoma sarsi Richard, 1894
D. sarsi is very common to lakes and reservoirs in tropical
regions except in the African continent. Philippine
collection localities extend from as far north as Buguey,
Cagayan in northern Philippines to Lake Lanao and
Surigao City in the southern Philippines. The genus
Diaphanosoma includes the largest limnetic cladocerans
collected from the Philippines where together with
Ceriodaphnia are considered as species replacements
of Daphnia in the tropics. It is known for its small head
with a sloping dorsal side where the eye occupies the
entire head. The ventral margin of its valves also has a
duplicature that is rounded at the distal end. The posteroventral margin of its valve has 13-40 small spines which
diminish dorsally and has two thin posterior dorsal spines.
Collected specimens had average lengths of 0.65 mm.
D. sarsi was first reported from Lake Taal during the
Wallacea expedition where until 1998, it was thought to
have been the only species from this genus to be in this
lake. They are usually found in low numbers where they
have similar frequencies of occurrence and abundance
with D. excisum.
Diaphanosoma excisum Sars, 1885
Korovchinsky (1992) and Korinek (2002) remarked that
D. excisum is common in tropical and subtropical regions
where they inhabit a wide range of environments from
acidic, to turbid, to slightly brackish lakes. In the country,
Philippine Journal of Science
Vol. 141 No. 2, December 2012
it has only recorded from a few localities which led to the
belief that it is rare in this part of the tropics. Among the
major Philippine lakes, D. excisum has only been reported
from Lake Laguna de Bay (Mamaril Sr. 2001; Mamaril
Sr. & Fernando 1978). It is characterized by its large head
with a well-developed dorsal part and a moderate to large
eye. The ventral margin of the carapace valves has a flap
that joins the margin almost at a right-angle. Samples of
D. excisum collected from the lake had mean lengths of
0.65 mm. Similar to the other two Diaphanosoma spp. in
Lake Taal, it occurred in low densities in 2008 with highest
recorded densities at < 30 ind. / l (April).
Diaphanosoma tropicum Korovchinsky, 1998
This species has only been recently separated from D.
modigliani. The revision was done based on samples
collected from Sri Lanka, Malaysia, Thailand, China and
the Philippines. D. modigliani is believed to be endemic
to Indonesian lakes Toba and Tempe (Korinek 2002;
Korovchinsky 1998). The Philippine specimens were
collected from Talisay, Batangas which is situated on
the northern shores of Lake Taal. Further examination of
samples collected from our survey validated the existence
of D. tropicum in the lake. It is characterized by its large
head with a protruding dorsal part and moderate to large
eyes. It may be differentiated from D. modigliani by its
strongly curved spine on the end of the upper antennal
branch, long ventral valve inflexion, large and fewer
denticles in the ventro-posterior valve margin, among
others (Korovchinsky 1998). It is the largest of the three
Diaphanosoma spp. in the lake at 0.67 mm mean length.
D. tropicum were observed to occur in similar densities
in the north and south basins of the lake and was least
abundant of all the cladoceran species present.
DISCUSSION
A total of nine species (six cladocera, three copepoda)
of crustacean zooplankton were recorded from Lake
Taal (Fig. 1). Seven genera from seven families were
represented by the following number of species:
Cyclopidae (1); Diaptomidae (1); Pseudodiaptomidae (1);
Bosminidae (1); Daphniidae (1); Moinidae (1); Sididae
(3). The species composition of zooplankton in Lake
Taal has been described as typically tropical, with low
diversity for limnetic zooplankton, the absence of large
cladocerans such as Daphnia and the existence of smallbodied species (Mamaril Sr. 2001). Predation pressure
from zooplanktivores such as Sardinella tawilis may have
led to the present size structure of limnetic zooplankton
in the lake (Fernando 2002; Papa et al. 2008). The
declining populations of this zooplanktivore in Lake Taal
may however trigger a shift in zooplankton community
Papa et al.: Taxonomic Account of Limnetic
Zooplankton in Lake Taal, Philippines
structure. Furthermore, the culture of herbivorous fish
such as tilapia in floating fish cages and eutrophication
may in the long-term cause changes in the phytoplankton
community which will later have an impact on the
zooplankton community as well.
This is also the first time that scanning electron
micrographs have been used in the Philippines to augment
light microscopy in validating micro-characters present
in selected species. SEM has been widely used by
other scientists to study zooplankton micro-characters,
especially minute details or complex three-dimensional
structures (Dussart & Defaye 2001) such as appendages,
setae and setule patterns, spine and spinule ornamentation
etc. Many publications have already began to include
scanning electron micrographs with traditional line
drawings in the description of novel zooplankton species
(Dussart & Defaye 2001; Suarez-Morales & EliasGutierrez 2001; Walter et al. 2006). In this study, the use
of SEM was useful in validating the presence or absence
of certain structures or minute details such as spinule
patterns in some appendages that were otherwise more
difficult to observe using conventional light microscopy.
Some examples are the spinules at the terminal portion of
the P5 endopod of A. dorsalis (Fig. 3D), the gonophore
and spinule ornamentation in the last thoracic segment of
female P. brehmi (Fig. 4A & B). The utilization of SEM
in systematic studies of crustacean zooplankton taxa
has been very useful in verifying certain structures that
were otherwise more difficult to examine in detail using
conventional microscopy. Although there are still a few
SEM facilities in the country, the availability of cheaper
table-top SEMs that require minimal specimen preparation
nowadays will probably lead to its wider utilization in
verifying taxonomically important micro-characters in
zooplankton in the Philippines. Future applications of
this technique in identifying zooplankton from other areas
will surely lead to a faster and more accurate evaluation
of zooplankton biodiversity.
In spite of the cosmopolitan nature of many of the species
encountered, our study yielded two interesting species for
Lake Taal which may be due to the increased sampling
effort put in this study compared to the past surveys and
the introduction of non-indigenous species into the lake.
The presence of Pseudodiaptomus brehmi in Lake Taal
may be considered as one of the few occasions that a
Pseudodiaptomid has been recorded in a freshwater
environment in the Philippines (Kiefer 1939a; Petersen
& Carlos 1984). This is made more significant by the fact
that they have not been found in more recent collections
from the two other lakes where they have previously been
recorded (Laguna de Bay and Lake Naujan). An earlier
paper by the senior author has listed P. brehmi as a new
record for Lake Taal (Papa & Zafaralla 2011). The other
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Zooplankton in Lake Taal, Philippines
Philippine Journal of Science
Vol. 141 No. 2, December 2012
interesting species found in the lake is Arctodiaptomus
dorsalis (Marsh 1907), a Neotropical invasive species. The
presence of A. dorsalis confirms the role of aquaculture
in the introduction of non-indigenous zooplankton, as
the spread of A. dorsalis has been linked to the presence
of tilapia cage culture (Papa et al. 2012). This may
also be the reason behind the introduction and spread
of the Neotropical rotifer Brachionus havanaensis in
many freshwater ecosystems (including Lake Taal) in
the Philippines (Papa et al. 2011). Aquaculture in Lake
Taal started as a pilot project in 1975 (Aypa et al. 2008),
modelled after the success of aquaculture in nearby Lake
Laguna de Bay. A. dorsalis was first recorded in the
Philippines from Laguna de Bay in 1991 (Tuyor & Baay
2001) and since Laguna de Bay serves as the source of
tilapia fingerlings that are used to stock other Philippine
lakes, it could have easily dispersed A. dorsalis to the
other lakes as well. Calanoid copepods were first recorded
in the lake in August 1989 (unpublished data of Mamaril
Sr. 2001). As under-regulated aquaculture practices and
the introduction of alien species continue in many lakes
in the archipelago, more alterations to the native limnetic
zooplankton fauna may occur.
Although this paper only presents the species composition
of limnetic zooplankton and does not give a complete
picture of the overall biodiversity for zooplankton in Lake
Taal as littoral areas were not sampled, it more or less
gives a complete account for the limnetic zone based on
the 18 sampling trips in six sampling sites that covered
both open water and aquaculture areas in a span of two
years (2008-2010). The difference in sampling strategies
that has to be employed for littoral zooplankton also
entails a separate analysis for this group (MatsumaraTundisi & Tundisi 2005). It is expected that the species
richness of zooplankton in Lake Taal will increase after
the inclusion of littoral species, and further new records
may be established given that it has been 26 years since
a comprehensive listing of littoral species (Mamaril Sr.
1986) and 11 years since a review of zooplankton species
diversity in Philippine lakes were published (Mamaril
Sr. 2001). The littoral zooplankton community is highly
diverse, and support species that are not usually found in
the pelagic zone (Fernando 2002; Peters & Lodge 2010).
Since these species serve as prey for many littoral fishes,
including larval fish, it is likewise important to update
their species composition. The impact of increasing
nutrient levels in the lake can also greatly influence
the community structure of the littoral zone. Given the
impacts of eutrophication on the pelagic zooplankton
community of Lake Taal (Papa et al. 2011), it would be
equally important to document its impact on the littoral
zooplankton community. It is also high time that a reevaluation of littoral zooplankton species for Lake Taal
(and the entire Philippines) be done in the light of recent
250
alterations on the littoral zones of lakes brought about
by human-mediated changes, such as the impact of
aquaculture on limnetic zooplankton composition. Other
smaller freshwater ecosystems must also be sampled given
how deteriorated some lake ecosystems are at present as
these places may reveal higher diversity compared to the
limnetic zones of tropical lakes.
ACKNOWLEDGMENTS
We would like to thank R. Eckmann and J. Hentschel
(University of Konstanz, Germany) for the use of the
SEM; T. Chad Walter, H. Dumont, and M. Holynska for
verifying our identification of P. brehmi, A. dorsalis and T.
crassus, respectively; and Aissa Domingo for the scientific
illustrations. The SEMs were taken during the DAAD
Research Fellowship of the first author. The samplings
were funded by the Tonolli Fund Postgraduate Fellowship
of the International Society of Limnology, the Philippine
Commission on Higher Education, and the UST Research
Center for the Natural and Applied Sciences.
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