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771
RAFFLES BULLETIN OF ZOOLOGY 2014
Taxonomy and distribution of four Cladoceran families (Branchiopoda:
Cladocera: Moinidae, Bosminidae, Chydoridae and Sididae) in
Philippine inland waters
Jhaydee Ann F. Pascual1, Eric Zeus C. Rizo1, Boping Han3, Henri J. Dumont3 & Rey Donne S. Papa1, 2*
Abstract. Cladocerans are one of the most abundant groups of zooplankton that serves as food for shes and as an
indicator of aquatic ecosystem health. This paper updates the diversity and distribution of cladocerans in freshwater
ecosystems in the Philippines. Samples from the University of Santo Tomas – Zooplankton Reference Collection
(UST-ZRC) (collected from 2006 to 2012) together with more recent collections (January to October 2013) were
examined and analysed. Results indicate the presence of 16 species from families Moinidae, Bosminidae, Sididae and
Chydoridae compared to the 56 species that were reported in previous studies. Moinids and Bosminids were mostly
encountered in the islands of Luzon and Mindanao but rarely found in the Visayas islands while family Sididae
and Chydoridae were more distributed throughout the archipelago. Family Sididae and Chydoridae are considered
as the most diverse and well-distributed of the four families inhabiting Luzon, Mindanao and some of the major
islands in Visayas. Though seven species of chydorids have been identied in this study, researchers observed very
limited number of individual chydorids in sampling locations where they have been collected. Recent alterations to
freshwater ecosystems may have also contributed to the disappearance of some species through changes in water
quality and the introduction of non-native species. Though no novel species have been identied from these four
families so far, our study provides a better understanding of distribution patterns among these taxa.
Key words. Zooplankton, species richness, biodiversity, freshwaters
RAFFLES BULLETIN OF ZOOLOGY 62: 771–794
Date of publication: 12 December 2014
http://zoobank.org/urn:lsid:zoobank.org:pub:291D68B9-3EB0-4E7D-B71A-AB4DCF3E4D17
© National University of Singapore
ISSN 2345-7600 (electronic) | ISSN 0217-2445 (print)
1The Graduate School University of Santo Tomas, Manila; Email: rspapa@mnl.ust.
edu.ph; reypaps@yahoo.com (*corresponding author)
2Research Center for the Natural and Applied Sciences and Department of Biological
Sciences, University of Santo Tomas, Manila
3Department of Ecology and Institute of Hydrobiology, Jinan University
Taxonomy & Systematics
INTRODUCTION
The Philippines (13.00 N, 122.00 E), located in Southeast
Asia between the West Philippine and Philippine Seas, is an
archipelago of 7,107 islands covering a total area of 300,000
square kilometres with three main island groups, namely:
Luzon, Visayas and Mindanao (Ong et al., 2002). The
Philippines is considered as a megadiverse country (Myers,
2000) with numerous endemic species . The archipelago has
immense natural resources that are a source of food, shelter
and livelihood that can cater its growing population. There
are about 211 lakes, 18 major river systems and 22 marshes,
swamps and reservoirs which provide energy, irrigation as
well as provide sources of sh either from open water sheries
or aquaculture (DENR, 2010). The rise of irresponsible
aquaculture practices in many Philippine lakes and other
inland waters has led to the decline of water quality and
has become a threat to the country’s freshwater biodiversity.
To avoid the possible misuse of the Philippines’ natural
resources including inland waters, several major freshwater
bodies such as Taal, Paoay, Lanao, and Naujan have been
declared as protected areas by the national government to
improve water quality monitoring and control the amount
of human activity in the lakes. However, much is yet to be
known about freshwater zooplankton (members of Class
Copepoda, Superorder Cladocera and Phylum Rotifera) in
many of these areas due to the lack of specialists and priority
by research funding institutions. Recent studies were focused
on identifying the composition and ecology of zooplankton
in several freshwater bodies in the Philippines, such as Lake
Taal (Papa et al., 2012) and Lake Paoay (Aquino et al., 2008).
Determining the distribution of freshwater zooplankton in
the Philippines has not been undergone since the work of
Petersen & Carlos (1984) and Mamaril (2001). The previous
studies of Brehm, (1938), Woltereck et al. (1941), Tsi-Chung
& Clemente (1954), Mamaril & Fernando (1978); Mamaril
(2001), Petersen & Carlos (1984); Korovchinsky (2000),
Aquino et al. (2008), Papa & Mamaril (2011), Papa &
Zafaralla (2011) and Papa et al. (2012) have identied almost
56 cladocerans in Philippine waters. There is a need to re-
analyse previous estimates of zooplankton species richness
and taxonomy given the recent advances in cladoceran
taxonomy and the development of more accurate microscopic
techniques. This, together with the more intensive use of
freshwaters for aquaculture and the decline of water quality
necessitate a re-analysis of previous estimates.
About 298 cladoceran species are currently known from
South East Asia, including 67 valid species, 68 that have
already been relegated as synonyms and 163 that represent
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Pascual et al.: Freshwater Cladocera of the Philippines
complexes of species (Korovchinsky, 2013). Recent advances
in microscopy and molecular phylogenetics may double
the number of species found worldwide. Identification
of specimens using other morphological characters then
considered irrelevant are now being studied through these
advances (Elmoor-Loureino, 2006). An estimation of the
number of taxa within families, genera and local faunas of
Cladocera reveals that only 129 species (17% of all known
species) may be considered as sufciently well described
(valid species) (Korovchinsky, 1996) With 28 genera present
in Philippine waters, endemicity is highly probable. Even
with this diversity, very few studies have been done in their
taxonomy and distribution in the country.
This overall lack of interest has become a handicap to both
their taxonomy and its use in sustainable aquaculture. Reports
on invasive zooplankton species which may eventually
become a threat to native species have already been reported
(Papa et al., 2012). Studies on the systematics, distribution,
and ecology of freshwater zooplankton can add information
needed to maintain a sustainable biodiversity for freshwater
habitats, as well as improve knowledge on their importance
to sheries.The research aimed to address these problems by
surveying the inland waters across the Philippine archipelago
for the diversity and distribution of zooplankton, specically
focused on four cladoceran families widely distributed in
the country and update on their taxonomy and geographical
distribution.
MATERIAL AND METHODS
Sampling. Zooplankton were collected from January to
October 2013 in six islands across the Philippines. Together
with previously collected samples (2006–2012) deposited
in the University of Santo Tomas - Zooplankton Reference
Collection (UST-ZRC), a total of 86 freshwater ecosystems
including 53 lakes, 17 rivers, 5 reservoirs and 11 other water
bodies such as swamps, ponds and temporary pools were
Fig. 1. Map of the Philippines indicating inland waters included
in this study. Refer to Table 1 for other pertinent details on the
sampling sites.
Fig. 2. Distribution of family Bosminidae based on recently collected
samples (2013) and stored samples in the UST ZRC.
analysed for cladocera (Fig. 1). All samples and pertinent
details regarding the various collecting sites are available
with the corresponding author. Plankton sampling was done
by towing 50, 80 and 100 μm mesh-size plankton nets from
several transects perpendicular to the lake shore. Littoral and
limnetic samples were stored separately unless there was no
clear demarcation between littoral and limnetic areas of the
lake (Papa & Hołyńska, 2013).
Materials examined. Samples from the UST-ZRC (Ref.
Nos. 0001–0105) and more recent collections (UST-ZRC
Ref. Nos. 0106–0309) were sorted, dissected and identied
using Olympus CX21 Compound Microscope and Swift
Stereomicroscope at the University of Santo Tomas (Manila,
Philippines). Additional analyses were also performed in Jinan
University (Guangzhou, China) using Zeiss Stereomicroscope
Discovery Version 2.0, Nikon Eclipse 80I, Zeiss Fluorescent
Microscope 8X10, Olympus CX41 Compound Microscope
and Hitachi TM300 Tabletop Microscope. Dissection
was done in a drop of glycerine in a glass slide and xed
permanently using nail varnish prior to examination under
light microscopes. For SEM viewing, specimens xed in
70% EtOH was transferred to 1ml of HMDS (Hexa-methyl-
disalizane) + 99% EtOH solution and kept for 15 minutes.
After which, one more change of 100% HMDS was done for
15 minutes and allowed to dry. Dried specimens were picked
up using ne dissecting needles and mounted on the stub
made of glass slide by using carbon tape. Gold coating was
carried out by the EMS 7620 Mini sputter coater attached to
the scanning electron microscope . SEM images were taken
using the Hitachi TM300 Tabletop Microscope.
Identication was aided by descriptions, taxonomic keys and
illustrations by Burckhardt (1924, 1941), Goulden (1968),
Fryer (1968), Kořínek et al., (1997, 1999, 2002), Mamaril
& Fernando (1978), Korovchinsky (1991, 1992, 1995, 1996,
1998, 2000), Dumont & Negrea (2002), Fernando (1980,
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RAFFLES BULLETIN OF ZOOLOGY 2014
2002), Smirnov (1996), Kotov et al., (2009), Petersen (2009)
and Papa et al. (2012).
Slides were deposited in the University of Santo Tomas –
Zooplankton Reference Collection (UST-ZRC) with accession
numbers 0072–0096A. The distribution of identied species
were noted and compared with those from previous studies.
Updated distribution maps were made using DIVA-GIS
Software Version 7.4.
RESULTS
Key to the species of the four Cladoceran families
1. Head with a protective head shield. Swimming antennae with
less than ten natatory setae ....................2 (Order Anomopoda)
– Head without a protective head shield. Swimming antennae
with more than ten natatory setae .............................................
..................................... 12 (Order Ctenopoda:Family Sididae)
2. Antennules fused with rostrum ........... 3 (Family Bosminidae)
– Antennules not fused with rostrum .........................................4
3 Distal part of the antennules diverging. Postabdomen cone-like
with rounded tip ...................................... Bosminopsis deitersi
– Distal part of the antennules parallel. Postabdomen rectangular
in shape with oblique anal opening .................Bosmina fatalis
4. Body laterally compressed. Rostrum present............................
..............................................................5 (Family Chydoridae)
– Body not laterally compressed. Rostrum absent .......................
............................................................... 11 (Family Moinidae)
5. Body more globular. Postabdomen short ..................................
........................................................ 6 (Subfamily Chydorinae)
– Body more oval. Postabdomen long ...8 (Subfamily Aloninae)
6. Rostrum blunt. Labrum with serrated teeth ..............................
........................................................... Ephemeroporus barroisi
– Rostrum pointed. Labrum without serrated teeth ...................7
7. Anal denticles gradually diminishing in size, valves without
polygons ...... Pleuroxus (Picripleuroxus) cf. quasidenticulatus
– Anal denticles almost equal in sizes, valves with polygons.....
........................................................... Chydorus cf. sphaericus
8. Ventral margin with long feathered setae. Postabdomen with
long terminal claw ...................................................................9
– Ventral margin with long naked setae. Postabdomen with short
terminal claw .........................................................................10
9. Antennae with two spines. Postabdomen terminal claws with
3 accessory spines ..................................Oxyurella singalensis
– Antennae with three spines. Postabdomen terminal claws with
2 accessory spines ........................Camptocercus cf. uncinatus
10. Two main head pores. Lateral head pores lobed ......................
............................................................................ Anthalona sp.
– Three main head pores. Lateral head pores circular .................
......................................................................... Alona pulchella
11. Ocellus absent. Head rounded with supraocular depression ....
.......................................................................... Moina micrura
– Ocellus present. Head triangular without supraocular
depression ...........................................Moinodaphnia macleayi
12. Head clearly delimited from the body. Antennule with a shorter
sensory seta than its basipodite, eye situated more ventrally ...
....................................................... 13 (Genus Diaphanosoma)
– Head not clearly delimited from the body. Antennule with a
longer sensory setae than its basipodite, eye situated more
dorsally .....................................................Latonopsis australis
13. Head cone shaped. Swimming antennae with hooked spines in
the apical ends of the rst and second segment of the antennal
branches .................................................................................14
– Head rectangular in shape. Swimming antennae with
straight spines in the apical ends of the first and second
segment of the antennal branches .........................................15
14. Head one half of the body size. Postabdomen without a strong
convex curve ........................................ Diaphanosoma dubium
– Head one third of the body size. Postabdomen with a strong
convex curve ......................................Diaphanosoma tropicum
15. Head large with a raised dorsal side. Carapace
with narrow free flap. Swimming antennae strong and
long ..................................................... Diaphanosoma excisum
– Head small with a sloping dorsal side. Carapace with
broad free flap. Swimming antennae weak and
short ......................................................... Diaphanosoma sarsi
Class Branchiopoda
Superorder Cladocera
Order Anomopoda
Family Bosminidae
Genus Bosmina Baird, 1846
The body is more or less globular ranging from 0.4–0.8 mm
in length. The head and the carapace have no demarcation.
Head shield has three types of pores (one dorsal, one
frontal and two paired lateral ones). The female rostrum
has an elongated conical extension where the antennules
are inserted. Antennules are elongated, recurved, immobile,
pluri-articulated and with sensory setae on the anterior
border. Ocellus is absent. The carapace has posterior border
that is more or less rectilinear ventrally ending in a mucro.
It has six pairs of trunk limbs but the last pair reduced to a
small, naked lobe. First limb of the male has a very distinct
copulatory hook. Second limb contains marginal setae of
endopodite. The exopod of the third limb has seven setae.
The fourth and fth limb has eight and ve setae respectively.
The postabdomen is rectangular in shape with anal opening
in apical position.
Bosmina fatalis Burckhardt, 1924
Specimen examined. 12 samples from Lake Taal, Lake
Buhi, Laguna de Bay, Lake Lumot-Mahipon and Pantabangan
Dam were examined and stored in UST-ZRC with reference
numbers 0083–0089, 0094A–0095A and 00134–00136.
Description. Body length ranges in 0.4–0.6 mm. Bosmina
fatalis has a long rostrum fused with antennules of variable
length and shape (Fig. 3B). B. fatalis in the study has an oval
shaped front head pore situated at the midpoint between the
two frontal setae that conforms to the description of (Fig.
4D) that conforms to the description of Kořínek et al. (1997).
A small and round median head pore (MHP) is also located
on the dorsal surface of the head which is connected to the
cord-like structure that is suggested to be a supraeosophageal
ganglion (Kořínek et al., 1997). Long mucro situated at
the distal end of the carapace. Setules are observed in the
postabdomen with a slightly convex ventral margin and an
anal opening at the lateral aspect. The postabdominal claw
consists of a row of 6–8 teeth with minute spines continuing
ventrally (Fig. 3C).
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Pascual et al.: Freshwater Cladocera of the Philippines
Distribution. Cagayan: Lake Bangalau, Benguet: Ambuklao
dam; Nueva Ecija: Pantabangan dam; Tarlac: Lake Tambo;
National Capital Region (NCR): Pasig River, Marikina River;
Laguna: Lake Palakpakin, Lake Kalibato, Lake Yambo, Lake
Tadlak, Lake Caliraya, Lake Lumot-Mahipon; Batangas: Lake
Taal; Camarines Sur: Lake Buhi, Lake Bato; Sorsogon: Lake
Bulusan; Oriental Mindoro: Lake Naujan; Negros Oriental:
Lake Danao, Lake Kabalin-an; Lanao del Norte: Agus IV
Dam; Agusan Del Norte: Lake Mainit; South Cotabato:
Lake Lahit, Lake Siloton, Lake Sebu; Lanao del Sur: Lake
Lanao (Fig. 2).
Remarks. B. fatalis collected in the Philippines were
compared to the same species sampled in Taihu Lake, China
(type locality). The species is reported in East Asian lakes and
reservoirs. B. fatalis can also be found from the temperate
zone in the north to the subtropical and tropical areas (Japan,
Philippines, China and Indonesia) (Fernando, 2002).
Superorder Cladocera
Order Anomopoda
Family Bosminidae
Genus Bosminopsis Richard, 1895
The head of Bosminopsis is relatively smaller than Bosmina.
The rostrum is fused with antennules that diverge at the distal
end. A cervical sinus separates the head from the carapace.
Postabdomen is cone-like narrowing to the obtusely rounded
tip. Short but robust postabdominal claws are distinguishable
on the tip of the postabdomen.
Bosminopsis deitersi Richard, 1895
Specimen examined. Three samples from Lake Bangalau
in Cagayan and Lake Naujan, Mindoro were stored in UST-
ZRC with reference numbers 0090–0092.
Fig. 3. Bosmina fatalis parthenogenic female. A, habitus; B,
antennule; C, terminal claws; D, postabdomen.
Description. Body length ranges in 0.1–0.4mm. Distributed
through the tropics and some localities in the temperate,
this species are characterised by relatively small with a
supraocular depressions or slight keels (Fig. 5A). The
antennule and the rostrum are merged (Fig. 5B). The distal
part of the antennules are diverging which makes Bosminopsis
deitersi unique from other species under Bosminidae (Fig.
5B). A cone-like postabdomen contains a rounded tip (Fig.
5C) that is mostly seen protruded outside of the carapace.
A short sharp postabdominal claw is not similar to most
Bosmina. It contains small teeth with small spines continuing
dorsally.
Distribution. Cagayan: Lake Bangalau; Camarines Sur: Lake
Bato, Lake Baao; Oriental Mindoro: Lake Naujan (Fig. 2)
Remarks. The species is reported in both tropics and
subtropics in the world with several localities in the north
temperate zone. Bosminopsis deitersi collected in the
Philippines was compared with the species in Taihu Lake,
China and revealed the same morphological characters in
exception of the setae at the tip of the antennule which are
not present in the samples coming from the Philippines
(Fernando, 2002).
Class Branchiopoda
Superorder Cladocera
Order Anomopoda
Family Chydoridae
Subfamily Chydorinae
Genus Chydorus Leach, 1816
Body globular with ventral margin with setae inserted on its
inner surface. Head shield with acute rostrum with two major
head pores. Have large labral plates. Antennules do not have
a peg. Swimming antennae usually with 8 setae. Species of
the genus are the commonest anomopods. Morphological
variability has made this genus a possible genus group.
Chydorus cf. sphaericus (Mueller, 1785)
Specimen examined. Six parthenogenic female.One mounted
specimens (UST-ZRC 0120-Lake Malbato, Palawan). Four
were not mounted due to rarity of specimens collected. One
specimen was damaged beyond repair.
Description. Chydoridae is the largest family in Anomopoda
and the largest family under Superorder Cladocera. It has
two recognised sub families: Chydorinae and Aloninae
(Dumont & Negrea, 2002). Chydorus sphaericus is one of
the most common species of chydorids found worldwide.
It is characterised by a subglobular body with valves
having polygons. It has a well pointed rostrum and a very
short antennule. Its labral plate is cuneiform and has an
elongated tip. It has a short post abdomen with 8–10 anal
teeth. It occurs in both the littoral and the limnetic zones. It
has limited distribution in the Philippines found mostly in
freshwater bodies with good macrophyte cover in the littoral
zones. (Fig. 8)
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RAFFLES BULLETIN OF ZOOLOGY 2014
Fig. 4. Scanning electron micrographs of Bosmina fatalis. A, habitus; B, frontal head pore (FHP); C, antennule showing the dimensions
of the rostrum; D, lateral head pore (LHP); E, postabdomen showing claw; F, inverted postabdominal claw showing pecten and setules.
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Pascual et al.: Freshwater Cladocera of the Philippines
Fig. 5. Bosminopsis deitersi. A, habitus; B, antennule; C,
postabdominal claw showing the post-anal extension; D, antennae.
Distribution. Palawan: Lake Malbato; Misamis Oriental:
Lake Danao; Bukidnon: Lake Pinamaloy (Fig. 7).
Remarks. It has a possible worldwide distribution but are
less morphologically variable in the tropics. Recognised as
a species complex and possibly be separated into several
genera and species. It occurs in both littoral and limnetic
zones. It is dominant in highly eutrophic waters with dense
macrophyte cover.
Genus Ephemeroporus Frey, 1982
Body globular to sub globular. Head shield without head
pores. Ventral margins with setae that are sub marginal.
Rostrum short and blunt. Labral plate with 1–4 serrated teeth.
Post abdomen is elongated with a strong sharp pre-anal angle;
post anal denticles marginal with 2–4 denticles longer than
the rest; terminal claw evenly curved with two basal spines.
Ephemeroporus barroisi (Richard, 1894)
Specimen examined. Six parthenogenic female (UST-ZRC
0126,0131-Lake Danao, Camotes Is., Cebu; 0127–0129- Lake
Mainit, Agusan Del Norte;0128- Lake Paoay, Ilocos Norte).
Description. Relocated from the genus Chydorus, this species
is easily identied by its serrated labrum with four teeth
(Frey, 1982). Its body is globular to subglobular like other
chydorids. It has short blunt rostrum in comparison with C.
sphaericus. Its ocellus is large but slightly smaller than the
eye. The post abdomen is ciliated along its entire length. It
is found only in the tropics. It is relatively well distributed
in the Philippines with specimens found from Luzon to
Mindanao (Figs. 9, 10).
Distribution. Ilocos Norte: Lake Paoay; Cagayan: A dam
in Binag, Lake Nalbuan, Lake Calig, Camarines Sur: Lake
Baao; Cebu: Lake Danao; Negros Oriental: Lake Kabalin-
an: Agusan Del Norte: Lake Mainit; South Cotabato: Lake
Lahit, Lake Siloton (Fig. 7)
Remarks. Possibly well distributed in the tropics and
sub tropics with specimens reported in Syria, Iran, India,
Sri Lanka, Australia, Nicaragua and North America. It
is considered as a species complex that can be separated
into several species. Fairly common in littoral zones and
sometimes present in low numbers in the limnetic and lotic
habitats.
Genus Pleuroxus Baird 1843
Body circular to oval. Ventral valves with setae attached to its
edges. Posterior dorsal angle of valve lower than the height
of the carapace. Antennules with peg. Swimming antennae
with spines. Post abdomen of average length; Terminal claw
with two basal spines. Several sub genera taken from this
genus are still in question and are usually used more often
as a sub genus rather than a genus.
Pleuroxus (Picripleuroxus) cf. quasidenticulatus
(Smirnov, 1996)
Specimen examined. Two parthenogenic female. One
mounted (UST-ZRC 0138-Lake Nalbuan, Cagayan). One
damaged beyond repair.
Description. The status of Picripleuroxus as a genus has
long been strongly argued. The latter has been re-established
as a subgenera in recent literatures. Carapace is oval with
no distinct characters in the posterior part of the dorsal
margin but rather high on its lateral part. Rostrum curved
and pointed. Ocellus situated near the tip of the rostrum.
Antennae with one spine on the apical ends of the antennal
segment. Post abdomen a little bit long and narrowing to
the terminal claw. Anal teeth size is decreasing across the
length of the post abdomen but variable across population.
Terminal claw with two accessory spines near its base. It
has only been observed in two lakes in Cagayan province
contrary to previous sightings (Fig. 11).
Distribution. Cagayan: Lake Nalbuan, Lake Cansiritan
(Fig. 7).
Remarks. Found also in Australia, Iraq, Argentina, Thailand,
Vietnam China and Far East Russia. Commonly found in
littoral zones of ponds and reservoirs and streams.
Subfamily Aloninae
Genus Alona Baird 1843
Body oval to elliptical curved to moderately convex; ventral
valve margins with rows of setae in similar sizes. Head
shield with two main head pores connected and circular
lateral pores. Ocellus smaller than the eye. Well-developed
blunt rostrum. Two antennular aesthetes projecting from
the rostrum. Labral plate variable Swimming antennae with
well- developed spines. Post abdomen narrow to broad with
several variably sized anal denticles with setules. Genus
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Fig. 6. Scanning electron micrograph of Bosminopsis deitersi. A, habitus; B, second antennae; C, postabdominal claw; D, mucro.
considered as a complex group and are still being separated
into several genera.
Alona pulchella King, 1853
Specimen examined. Ten parthenogenic female. Six mounted
specimens (UST ZRC 0121, 0122-Lake Danao, Camotes Is.,
Cebu;0123–0125-Lake Calig, Cagayan 0137-Lake Nalbuan,
Cagayan). Four damaged beyond repair.
Description. The genus Alona is the largest anomopod genus.
It represents almost half of the species of Chydoridae. The
genus itself has been broken down to several genera in recent
years. The species represent many synonyms and is recently
being separated into several species. In 2002, A. cambouei
was separated from this group. It has an oval body with
around 40–45 setae on the ventral margin. Its head shield is
elongated and has short and blunt rostrum. It has a narrow
post abdomen of moderate length. Found in several freshwater
bodies in all major regions of the Philippines from rivers to
lakes and reservoirs (Fig. 12)
Distribution. Cagayan: Lake Calig, Lake Nalbuan; Benguet:
Ambuklao Dam; Bontoc and Kalinga: Chico River; Laguna:
Lake Tadlak; Camarines Sur: Lake Baao; Cebu: Lake Danao;
Agusan Del Norte: Lake Mainit, Agusan River; Lanao Del
Norte: Agus IV Dam (Fig. 7)
Remarks. Considered as a species group with tropical to
sub-tropical distribution in Asia, Africa and Australia. Fairly
common in littoral zones of lakes, rivers and streams.
Anthalona sp. Van Damme, 2011
Specimen examined. 10 parthenogenic female. Three
mounted (two whole mounts (UST ZRC 0140, 0141, one
dissected UST ZRC 0142). Five subjected to SEM. Two
were damaged beyond repair.
Description. Recently established in 2011 by Van Damme,
the genus can easily be recognised by their ower-like (lobed)
lateral head pores. Body is oval with a prominent rostrum.
Has a characteristic connected main head pores and lobed
lateral head pores. Eye larger than its ocellus. Labral plate axe-
shaped. Post abdomen wide and short and with an S-shaped
appearance. Terminal claw long with one small accessory
basal spine (Fig. 13, 14). Specimen found in Lake Lanao
is morphologically similar to A. alonopsiformis described by
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Pascual et al.: Freshwater Cladocera of the Philippines
Fig. 7. Distribution of family Chydoridae based on recently collected
samples (2013) and stored samples in the UST ZRC.
Brehm in 1933 by its terminal claw but extensive analsysis
is still needed to elucidate its identity.
Distribution. Cagayan: Lake Calig; Mt. Province: Chico
River; Lanao del Sur: Lake Lanao (Fig. 7)
Remarks. The genus is mainly distributed in the southern
hemisphere with records in Central America, South America,
tropical Africa, Indonesia and Australia. They do not tolerate
colder waters, with the northern limit of the genus found in
the Mediterranean. It prefers shallow sandy or rocky littoral,
detritus ridden and heavily vegetated pools and swamps.
Genus Camptocercus Baird, 1843
Body ovoid with long feathered setae in its ventral margin
of its carapace. Head shield with three major head pores.
Fig. 8. Chydorus cf. sphaericus parthenogenic female. Lake Malbato,
Coron, Palawan. A, habitus; B, antennae; C, post abdomen.
Fig. 9. Ephemeroporus barroisi parthenogenic female. A, habitus;
B, labrum; C, post abdomen.
Fig. 10. Scanning electron micrographs of Ephemeroporus barriosi.
A, habitus; B, antennae.
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Fig. 11. Pleuroxus cf. quasidenticulatus parthenogenic female. A,
habitus; B, antennae; C, post abdomen.
Fig. 12. Alona pulchella parthenogenic female. A, habitus; B,
antennae; C, post abdomen.
Rostrum slightly pointed. Labral plate with widely rounded
apex and cuneiform in shape. Antennule usually reaching the
tip of the rostrum. Long but weak swimming antennae. Post
abdomen is long and narrow with 1–2 accessory spines on
its terminal claw; anal denticles pass over each other
Camptocercus cf. uncinatus Smirnov, 1971
Specimen examined. One parthenogenic female (UST-ZRC
0139-Dam in Binag, Cagayan).
Description. First described by Smirnov in 1971, it has a
worldwide distribution and is almost exclusively littoral as
most chydorids are. It has an oval body with several feathered
setae in its ventral margin. It has a very small ocellus situated
near its rostrum. It has long antennules almost reaching the
apex of the rostrum. Its labrum has an irregular convex
anterior margin and a few spines and setules on its ventral
margin. It has a very narrow distinct post abdomen with
14–16 anal denticles. A single specimen was observed in a
local reservoir in Cagayan province (Fig. 15).
Distribution. Cagayan: A dam in the town of Binag (Fig. 7).
Remarks. A widely distributed species, found in Romania,
southwest Siberia, Israel, Iraq, Ethiopia, Egypt, Mexico, and
Guatemala. Common in littoral zones of lakes and small
streams with thick macrophyte vegetation.
Genus Oxyurella Dybowski & Grochowski, 1894
Body ovoid with ventral margins having long feathered
setae. Head shield with two separate major head pores with
minor head pores in between and lateral to them. Rostrum
Fig. 13. Anthalona sp. parthenogenic female, Lake Lanao, Lanao
del Sur. A, habitus; B, antennae; C, postabdomen
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Fig. 14. Scanning electron micrographs of Anthalona sp. A, habitus; B, main and lateral head pores; C, antennae; D, post abdomen.
not elongated. Labral plate cuneiform and blunt. Antennule
with 8–9 sensory aesthetes inserted in the rostrum. Swimming
antennae with three spines. Post abdomen broad with large
anal denticles; terminal claw with 2–3 basal spines.
Oxyurella singalensis (Daday, 1898)
Specimen examined. Two parthenogenic female (UST ZRC-
0132-Lake Sampaloc, Laguna; 0133-Lake Calig, Laguna)
Description. It is common in stationary water bodies and
not widely observed in lotic ecosystems. Its body is oval-
oblong in shape. The carapace of the species does not have a
keel. Its antennule reaches the tip of its rostrum. It has short
swimming antennae with two strong well developed spines.
The ventral margin of its carapace has long feathered setae.
Relatively narrow, the post abdomen of O. singalensis has
a long basal spine with 3 accessory spines on its base. It
was found exclusively in Luzon in both littoral and limnetic
samples (Figs. 16, 17)
Distribution. Cagayan: A dam in the town of Binag, Lake
Nalbuan; Pampanga: Candaba swamp; Laguna: Lake
Sampaloc, Lake Kalibato (Fig. 7)
Remarks. Common in tropical and subtropical Asia, Africa
and Australia. Found in sandy and muddy sediments of littoral
zones of lakes and reservoirs.
Class Branchiopoda
Superorder Cladocera
Order Anomopoda
Family Moinidae
Genus Moina Baird, 1850
The moinids have a large fused compound eye but lack the
ocellus in the genus Moina, although it is present in one
species of Moina (reticulata). The head is frequently indented
above the eye (called the supra-ocular depression) by the
attachment of a muscle bundle to the inner surface of the
exible exoskeleton. The second antenna consists of a large
basiopod, an endopod with three segments and an exopod of
four segments. A third sensory seta is present on the distal
end of the basiopod between two rami. The endopod bears
one or more long swimming setae on its distal end. The
four-segmented exopod contains long swimming setae. The
exopod also bears inner row of short teeth that is suspected
to clean dentritus from the surface of the shell. Moinids
have ve thoracic limbs and each limb has been specically
modied towards greater efciency in its particular function.
The rst pair of limbs no longer functions for ltering. It has
reduced in size and fewer setae that found on the sidids. The
two distal setae in the second limbs are modied for grasping
food particles from the posterior limbs. The third and the
fourth limbs have large lter comb. The fth limb serves in
closing off the posterior end of the lter chamber forward
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though the lter comb of the fourth leg. Postabdomen has a
large post-anal extension which is conical and ends with the
distal claws. There is a row of lateral feathered teeth and a
single distal bident tooth on the postabdomen. The claw is
usually pectinate and has basal spines at the ventral base of
the claw that is also called as “Basaldorn”
Moina micrura Kurz, 1875
Specimen examined. 11 adult parthenogenic female
specimens from Lake Taal, Lake Sampaloc, and Candaba
Swamp were examined and stored in University of Santo
Tomas Zooplankton Reference Collection (UST-ZRC) with
reference numbers 0072–0081, and 0093.
Description. Moinidae is composed of only two genera:
Moina and Moinodaphnia. Moina are found from temporary
freshwater pools to brackish and saline lakes. Moina micrura
are one of the smallest species in Family Moinidae. Mean
length in all specimen examined is 1.1 mm. The head is
evidently characterised by a well-developed supraocular
depression and a large eye (Fig. 19A). The antennules of
M. micrura are situated in a knob-liked position just behind
the eye with two sensory setae in each located one-third to
one-half the distance from the head. The sensory papillae
are long and distinguishable. A distinct groove located
behind the second antennae separates the head and the body.
The exopod is composed of four-segmented ramus which
contains heavy setation that extends from second to the
fourth segment . Two short sensory setae are evident in the
second antennae as shown in Fig. 19E. The P1 has reduced
Fig. 15. Camptocercus cf. uncinatus parthenogenic female. A,
habitus; B, antennae and antennule; C, post abdomen
Fig. 16. Oxyurella singalensis parthenogenic female. A, Habitus;
B, Antennae; C, Post abdomen.
its ltering functions but have fewer setae compared to
other sidids (Fig. 19D). Carapace is round and surrounded
by 37–40 setae in the shell margin.The postabdomen is short
and slender. The distal conical portion composes one-fourth
of the total length. A sharp postabdominal claw contains three
to eleven feathered teeth (Fig. 19C). The number of teeth
varies on the body size of the specimen. Its dorsal margin
is characterised by having several numbers of setae extend
up to the sharp part of the claw. There is also a long bident
tooth present located at the base of the feathered teeth. The
long and sharply curved postabdominal claw contains four
to seven sharp teeth that are also called “basaldorn” that
extends towards the distal end (Goulden, 1968).
Distribution. Ilocos Norte: Lake Paoay; Cagayan: Lake
Nagatutuan; Benguet: Ambuklao Dam; Pangasinan:
Fish Pond in Burgos; Tarlac: Lake Tambo; Nueva Ecija:
Pantabangan Dam; Zambales: Lake Mapanuepe; Pampanga:
Candaba swamp; National Capital Region (NCR): Pasig
River, Marikina River; Laguna: Lake Bunot, Lake Palakpakin,
Lake Sampaloc, Lake Pandin, Lake Kalibato, Lake Tadlak,
Lake Caliraya, Lake Lumot-Mahipon, Laguna De Bay;
Batangas: Taal Lake; Camarines Sur: Lake Buhi; Oriental
Mindoro: Lake Naujan; Misamis Oriental: Lake Gumaod;
South Cotabato: Lake Siloton, Lake Lahit, Lake Sebu (Fig.
18)
Remarks. M. micrura can be distinguished from other
Moinids based from its small size and by the complete
absence of hairs on both the head and shell. The species is
reported in the Palearctic region and also from the tropics
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Pascual et al.: Freshwater Cladocera of the Philippines
and subtropics of America, Australia (2002) as well as in
Africa and Europe (Goulden, 1968). Goulden (1968) also
conrmed its presence in the Far East and Southeast Asia
specically in India, Indonesia and Taiwan. The species is
mostly found in temporary water bodies but is also common
in large plankton of large freshwater lakes (Fernando, 2002;
Goulden, 1968).
Genus Moinodaphnia Herrick, 1887
Moinodaphnia can easily be distinguished from Moina by
the presence of an ocellus. The antennules are long and
movable. The females are laterally attened, and the dorsal
margin of the shell has a sharp keel. The second antennae
are very characteristic because the distal segment of the
exopod has four rather short setae rather than three long
setae. Postabdomen is long. Only one sexual egg is normally
deposited in the ephippium.
Moinodaphnia macleayi King, 1853
Specimen examined. Two specimens were examined
and stored in UST-ZRC with reference number 0082 and
0133 collected from Lake Calig and a temporary pool in
Asibanglan, Pinukpuk, Kalinga.
Fig. 17. Scanning electron micrographs of Oxyurella singalensis. A, habitus; B, post abdomen; C, Dorsal margin; D, rostrum and antennule.
Description. The body length ranges in 0.9–1.3 mm. The
head is sub-triangular in shape. The large eye lls the tip
of the head with an ocellus present below the eye and is
located above the origin point of the antennules (Fig. 21D).
Supraocular depression is not present in Moinodaphnia
macleayi. The antennules are long and thin, lack the vertical
row of long hairs, and are ornamented only with horizontal
rows of short setae. The distal end of the antennule has nine
sensory papillae. One segment of the second antennae, the
basiopod has two sensory setae which originate from the
distal end of the rami (Fig. 12B). One of the four swimming
seta has been reduced in size for genus Moina but remains
its longer size in Moinodapnia. The valves of the carapace
joined together in the mid-line of the body axis and have a
row of small setae. The postabdomen is similar to the genus
Moina containing 10–11 feathered teeth and one long bident
tooth on the lateral side with pattern of ne setae in the dorsal
margin. (Fig. 21C).The claw lacks a pecten but with a row
of ne short hairs.
Distribution. Cagayan: Lake Calig; Kalinga: Temporary
pool in Asibanglan, Pinukpuk (Fig. 18).
Remarks. Moinodaphnia has a different distribution and
habitat from that of Moina. It prefers to be in small temporary
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ponds, small lakes, swamps and pools. The species lives near
the mud or in the weeds where it lters the water surrounding
the weeds. It is distributed throughout the humid tropics
specically in Africa, Australia, South America, Caribbean
Islands, Philippines and India (Fernando, 2002).
Class Branchiopoda
Superorder Cladocera
Order Ctenopoda
Family Sididae
Genus Diaphanosoma Fischer, 1850
Head of different forms and sizes, usually conical or
rectangular with noticeable dorsal impression in between the
head and the valves. Antennules small in size. Swimming
antennae are long and strong. Valves forms wide flap
inexions. Post abdomen without anal teeth, with terminal
claws with three basal spines.
Diaphanosoma dubium Manuilova, 1964
Specimen examined. Eight parthenogenic female (six
Philippine specimens and two Chinese specimens). Three
mounted Philippine specimens and one Chinese specimen
(UST-ZRC 0114-Lake Sampaloc, Laguna; 0115, 0116 - Lake
Tambo, Tarlac; Guangzhou City, Guangdong-0117). Four
specimens were damaged beyond repair.
Description. This Diaphanosoma species is characterised by
a very large cone-shaped head, almost ½ of its body size.
Spine on the apical end of the antennal branch can either
be curved or straight. Number of denticles in its carapace is
increasing with body size with thin setules in between them.
It commonly coexists with D. tropicum in tropical and sub-
Fig. 18. Distribution of family Moinidae based on recently collected
samples (2013) and stored samples in the UST ZRC.
Fig. 19. Moina micrura parthenogenic female. A, habitus; B,
antennule with exopodite; C, postabdomen; D, P1; E, antennae.
tropical Asia (Korovchinsky, 2000). In the Philippines it also
co-occurs with D. sarsi and D. excisum in several freshwater
bodies in Luzon (Fig. 24).
Distribution. Tarlac: Lake Tambo; Laguna: Lake Bunot,
Lake Yambo, Lake Mohicap, Lake Palakpakin, Lake Pandin,
Lake Sampaloc, Lake Caliraya, Laguna De Bay; Camarines
Sur: Lake Baao, Bicol River; Agusan Del Norte: Agusan
River (Fig. 23)
Remarks. Species can also be found in Russia, China,
Mongolia, Bangladesh, Thailand, Sri Lanka, Vietnam and
Malaysia. Comparative specimens collected from China
(Ming Lake, Jinan University, Guangzhou City) have strongly
curved apical antennal spine in comparison with Philippine
specimens with straight spines. The species is common in
lakes, reservoirs and sh ponds. It is primarily limnetic and
can co-exist with other sidids and limnetic cladocerans.
Species have bigger head in comparison with its two closely
related species D. tropicum and D. modigliani. It has a
variable spine on its antennal branches ranging from straight
to strongly curved.
Diaphanosoma excisum Sars, 1885
Specimen examined. Seven parthenogenic female (UST
ZRC 0105, 0107-Lake Palakpakin, Laguna; 0106-Lake Buhi,
Camarines Norte; 0108-Fish Pond, Laoagan Inn Resort,
Kalinga; 0109-Lake Taal, Batangas, 0110, 0111-Lake Cassily,
Cagayan).
Description. The species is common throughout the inland
waters of the country. Almost always co-existing with D.
sarsi, it can easily be differentiated from the latter with
its strongly-developed dorsal part on its head. This gives
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Fig. 20. Scanning electron micrographs of Moina micrura parthenogenic female SEM micrograph. A, habitus; B, antennae; C, antennule
with exopod; D, postabdominal claw.
its head a well large rectangular appearance. Eye ranges
from moderate to large. Its carapace oblong-oval in shape,
with ventral parts forming a narrow free ap with feathered
margins. Its ventral margins have large sharp denticles. It is
fairly common in the tropics and subtropics, It can survive
different environments from highly turbid to slightly saline
waters (Chatterji et al., 1995) (Figs. 25, 26).
Distribution. Ilocos Norte: Lake Paoay; Cagayan: Lake
Nalbuan, Lake Calig, Lake Nagatutuan, Lake Cansiritan,
Lake Cassily; Benguet: Ambuklao dam; Kalinga: Fish pond
in Laoagan Resort Inn in Tabuk; Pangasinan: Fish pond in
Burgos; Tarlac: Lake Tambo; Nueva Ecija: Pantabangan
Dam; Zambales: Lake Mapanuepe; National Capital Region
(NCR): Pasig River; Marikina River; Laguna: Lake Bunot,
Lake Yambo, Lake Mohicap, Lake Palakpakin, Lake Pandin,
Lake Sampaloc, Lake Tadlak, Lake Kalibato, Lake Lumot-
Mahipon, Laguna De Bay; Batangas: Lake Taal; Camarines
Sur: Lake Bato, Lake Baao, Lake Buhi; Oriental Mindoro:
Lake Naujan; Leyte: Lake Bito; Bohol: Malinao Dam;
Agusan del Norte: Lake Mainit; Lanao del Norte: Agus IV
Dam, Tubud-Mayahay River; Bukidnon: Lake Pulangi, Lake
Pinamaloy; South Cotabato: Lake Siloton (Fig. 23)
Remarks. D. excisum is also distributed in tropical and sub-
tropical Australia, India, Eastern China, Nepal and possibly
Northern Africa. One of the more adaptable sidids, it can
survive in acidic, turbid and even slightly brackish waters. It
is common to water bodies with aquaculture and co-existing
with other limnetic cladocerans. It can be easily differentiated
from D. sarsi with the strong dorsal portion of its head and
more massive antennae.
Diaphanosoma sarsi Richard, 1894
Specimen examined. Ten parthenogenic female, eight
mounted specimens (UST-ZRC 0097, 0099-Lake Paoay,
Ilocos Norte; 0098 - Lake Danao, Camotes Island, Cebu;
0100-Lake Buhi, Camarines Sur; 0101-Lake Tambo, Tarlac;
0102-Lake Nalbuan, Cagayan; 0103, 0104-Lake Sampaloc,
Laguna), two were damaged beyond repair.
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Description. Diaphanosoma sarsi is the most well-distributed
cladoceran found throughout the Philippines. It is found in
almost all types of water bodies from swamps to a few river
systems. This species is distinguished by a small roundish
rectangular head with sloping dorsal side. Its eye occupies
almost the whole area of the head. The carapace of D. sarsi
is situated high on its body with its ventral parts forming a
broad free ap. Its ventral margins have around 13–40 small
denticles. It is fairly common throughout inland water bodies
especially in lakes and reservoirs in the tropics except Africa
(Fernando, 2002). In the Philippines it co-exists with three
other species of Diaphanosoma (Figs. 27, 28).
Distribution. Ilocos Norte: Lake Paoay; Cagayan: Lake
Nalbuan, Lake Nagatutuan Lake, Lake Bangalau; Benguet:
Ambuklao dam; Kalinga: Fish pond in Laoagan Resort Inn;
Pampanga: Candaba swamp; Tarlac: Lake Tambo; Zambales:
Lake Mapanuepe; National Capital Region (NCR): Pasig
River, Marikina River; Laguna: Lake Bunot, Lake Yambo,
Lake Mohicap, Lake Pandin , Lake Sampaloc, Lake Tadlak,
Lake Kalibato, Lake Lumot-Mahipon, Lake Caliraya, Laguna
De Bay; Batangas: Lake Taal; Camarines Sur: Lake Baao,
Lake Buhi; Sorsogon: Lake Bulusan; Oriental Mindoro:
Fig. 21. Moinodaphnia macleayi parthenogenic female. A, habitus;
B, antennule with exopod; C, postabdomen; D, antennae.
Fig. 22. Scanning electron micrographs of Moinodaphnia macleayi parthenogenic female; A. habitus; B, head showing the antennule; C,
antennule; D, antennae showing bas, end, and exopod.
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Pascual et al.: Freshwater Cladocera of the Philippines
Fig. 23. Distribution of family Sididae based on recently collected samples (2013) and stored samples in the UST ZRC.
Fig. 24. Diaphamosoma dubium parthenogenic female. A, habitus;
B, postero-ventral valve margin; C, distal part of second segment
of upper 2-segmented antennal branch (Phil.); D, distal part of
second segment of upper 2-segmented antennal branch (China);
E, postabdomen.
Fig. 25. Diaphamosoma excisum, parthenogenic female. A, habitus;
B, postero-ventral valve margin; C, distal part of rst segment of
upper 2-segmented antennal branch; D, post abdomen.
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Lake Naujan; Negros Oriental: Lake Balinsasayao; Leyte:
Lake Danao, Lake Bito; Bohol: Malinao Dam; Agusan del
Norte: Agusan River; Lanao del Norte: Agus IV Dam, Tubud-
Mayahay River; Bukidnon: Lake Apo, Lake Pulangi, Lake
Pinamaloy; South Cotabato: Lake Lahit (Fig. 23)
Remarks. Species is found in tropical and sub-tropical
Asia. Populations have been found in Nepal, Eastern China,
Australia, New Guinea, New Caledonia, New Hebrides,
Guam and few sites in tropical Africa. It can be found in a
wide range of water bodies from large ancient lakes to rice
elds. Common sidid found in lotic ecosystems. Prefers
shallow waters with heavy vegetation but can also be observed
in the limnetic zones of lakes. They are more sensitive to
physico-chemical parameters in comparison with other
Diaphanosoma species. D. sarsi has a noticeable sloping
dorsal portion of its head that can easily differentiate it from
other closely related species.
Diaphanosoma tropicum Korovchinsky, 1998
Specimen examined. Five parthenogenic female, two
mounted (UST ZRC 0112-0113-Lake Pandin, Laguna), One
specimen with decapitated head with two damaged beyond
repair.
Fig. 26. Scanning electron micrographs of Diaphanosoma excisum. A, distal part of rst segment of upper 2-segmented antennal branch;
B, postero-ventral valve margin; C, ventral valve inexion; D, terminal claws of post abdomen.
Description. Separated from D. modigliani by (Korovchinsky,
1998) after re-assessing its occurrence outside Lake Toba in
Indonesia, it is characterized by a cone shaped head almost
1/3 of its body size. Its rst record in the Philippines is in
Lake Taal (Papa & Zafaralla, 2011). It exists in several lakes
in Luzon Island and co-existing with the more common
D. sarsi and D. excisum. Ends of the distal segment of its
swimming antennae have a noticeable curved spine often
having hook like appearance. The postero-ventral margin
of its carapace have noticeable differing denticles as they
progress dorsally and long thin setules between them. Its post
abdomen prominently dorsal and have a strong convex curve.
Outside of the country it is found in Malaysia, Thailand, India,
Sri Lanka and China (Fernando, 2002) (Fig. 29).
Distribution. Ilocos Norte: Lake Paoay; Tarlac: Lake Tambo;
Laguna: Lake Bunot, Lake Palakpakin, Lake Pandin, Lake
Lumot-Mahipon, Laguna De Bay; Batangas: Lake Taal;
Camarines Sur: Lake Bato, Lake Baao, Lake Buhi (Fig. 23).
Remarks. Recent re-evaluation of localities of D. modigliani
outside Lake Toba by Korovchinsky in 1998 where the
species is commonly associated with have shown that the
species is also present in India, Sri Lanka, China, Thailand
and Malaysia. Found in lakes and reservoirs co-existing
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Fig. 27. Diaphamosoma sarsi, parthenogenetic female; A, habitus;
B, postero-ventral valve margin; C, distal part of rst segment of
upper 2-segmented antennal branch; D, post abdomen.
Fig. 28. Scanning electron micrographs of Diaphanosoma sarsi.
A, habitus; B, antennule.
with other limnetic cladocerans with aquaculture. It is an
adaptive species that can survive slightly brackish water
and a wide range of varying physico-chemical parameters.
It is differentiated from D. modigliani and D. dubium from
its strongly curved antennal spine and prominently curved
post abdomen.
Genus Latonopsis Sars, 1888
Massive head without noticeable separation from the body.
Eye situated closer to the dorsal side of the body. Antennules
relatively short. Two of the ventral most setae on the antennal
branch larger than others with hook ends. Post abdomen with
a number of anal denticles with terminal claws having two
basal spines. Genus is widely distributed in the tropics and
subtropics. Few species are able to survive in the temperate
regions.
Latonopsis australis Sars, 1888
Specimen examined. Six parthenogenic female.One mounted
specimens (UST-ZRC 0120-Lake Malbato, Palawan). Four
were not mounted due to rarity of specimens collected. One
specimen was damaged beyond repair.
Description. Contrary to previous records, the species is
rare and very few individuals were observed in the collected
samples. It is easily identied by its body structure where
its head is not clearly delimited from the body. Its eye
positioned near the dorsal side of the head. Sensory seta of
the antennule of L. australis is longer than its basipodite. Its
post abdomen is relatively small with few anal spines and
has terminal claws with two basal spines. It is also a poorly
described species composed of several sub species. They are
found mainly in the littoral zone of lakes and reservoirs, in
ponds, swamps, and other temporary water bodies (Fig. 30).
Distribution. Cagayan: Lake Calig; Oriental Mindoro: Lake
Naujan; Leyte: Lake Danao (Fig. 23)
Remarks. L. australis are also found in tropical and sub-
tropical Australia, Asia, Africa and America. Temperate
localities have also been reported in Italy, Yugoslavia Bulgaria
and Northern America. They are found in littoral zones of
lakes, reservoirs, and other water bodies with good vegetation.
They also thrive in the limnetic zones of pools but prefer
bottom sediments for its habitat. It is a poorly described
species with a possibility of being a species group.
DISCUSSION
Changes in taxonomy and more intensive sampling effort
revealed improvements to previously existing knowledge
on the species distribution and richness for cladocera in the
Philippines. This veries the observations of Korovchinsky
(2013) where out of 298 species known from Southeast
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Fig. 29. Diaphamosoma tropicum parthenogenic female. A, habitus;
B, postero-ventral valve margin; C, distal part of second segment
of upper 2-segmented antennal branch; D, post abdomen.
Fig. 30. Latonopsis australis parthenogenic female. A, habitus; B,
postero-ventral valve margin; C, antennule; D, distal part of second
segment of upper 2-segmented antennal branch; E, postabdomen.
Asia, only 67 were valid species, 68 have been relegated as
synonyms while majority (163 species) represent complexes
of species. As such, this just goes to show how more intensive
studies on Philippine species revealed further changes in
known distribution patterns. Several species described from
family Moinidae such as M. makrophthalma, M.dubia, M.
dubia parva, M. weismanni and Moina sp. (Brehm, 1938;
Woltereck et al., 1941; Tsi-Chung & Clemente, 1954; Ueno,
1966) were re-identied as synonyms of M. micrura by
Goulden (1968). Meanwhile, M. brachiata (Mamaril &
Fernando, 1978; Mamaril 2001) was not encountered in the
any of the inland waters sampled. Both species should not
be confused with one another on an account that M. micrura
has a smaller body size of 0.05–1.2 mm, a different head
shape, and the absence of a large claw pecten. The absence
of M. brachiata in the Philippines was supported by Goulden
(1968) and Fernando (2002) mentioning its wide distribution
in the north temperate regions specically in most countries
in Europe and some localities in Northern Africa. Tsi-Chung
& Clemente (1954), Peterson & Carlos (1984), Mamaril
(2001) and Aquino et al. (2008) noted the presence of M.
macrocopa in several lakes and ponds in Luzon but the lack
of descriptions and illustrations of signicant taxonomic
characters may cause possible confusion in the identity of
other Moinids present in the Philippines. M. micrura was
widely distributed in 17 lakes, two rivers, one reservoir and
one swamp in the islands of Luzon and four freshwater lakes
in Mindanao. Most moinids encountered were from natural
lakes, rivers, temporary swamps and pools in the country. New
records for Moinodaphnia macleayi were noted in a small
lake and a temporary pool in the northern Philippines (Fig.
2) but not in Lake Taal as previously reported by Mamaril
(2001). This result veried that Moinodaphnia macleayi has
a completely different habitat compared to other moinids
where it preferred small swamps, pools and lakes (Goulden,
1968). The distribution of the moinids in this paper conrms
all previous records that no single species from family
Moinidae is present in the Visayas group of islands (Fig. 2).
The absence of Moinidae in this area veries the ndings
in past studies regarding the lack of Moina in the islands.
Comparison between the results of the present survey and
previous ones revealed that for Family Bosminidae, Bosmina
longirostris was not present in more recently collected
samples in spite of the fact that it was commonly recorded
in the previous studies. Morphological variations in some
populations of B. fatalis could have led to misidentication
of B. longirostris because most morphological characteristics
of the two species are similar like the shapes of their rostrum,
frontal head, and shell spines . Such variations may have led
to some confusion which resulted to past misidentications of
B. longirostris. To avoid this, Kořínek et al. (1997) proposed
the use of the lateral head pore (LHP) pattern as diagnostic
characters for species under the genus Bosmina. All species
of Bosmina found in 25 inland waters were subjected to
SEM to further analyse the LHP pattern which resulted in the
identication of B. fatalis. Only Bosmina fatalis was identied
from samples collected for this current study. It is one of
the well-distributed cladocerans in the Philippines including
many parts of Luzon, Mindoro, Negros and Mindanao (Fig.
3). The distribution of B. fatalis in the archipelago validates all
past literatures that the species is common in East Asian lakes
790
Pascual et al.: Freshwater Cladocera of the Philippines
Table 1. List of the different sampling localities included in this study.
Site UST ZRC Name Municipality/ Province Island
No. Ref. No. City
1 154–151 Lake Nalbuan Buguey Cagayan Luzon
2 152–158 Lake Calig Buguey Cagayan Luzon
3 159–162 Lake Bangalau Sta. Teresita Cagayan Luzon
4 163–165 Binag Dam La–lo Cagayan Luzon
5 171–172 Lake Cansiritan Cansiritan Cagayan Luzon
6 166–170 Lake Nagatutuan Gattaran Cagayan Luzon
7 173–175 Cagayan River Tuguegarao Cagayan Luzon
8 176–177 Lake Cassily Tuao Cagayan Luzon
9 178–180 Callao Caves Penablanca Cagayan Luzon
10 185–188 Chico River (Upstream) Pinukpuk Kalinga Luzon
11 189 Temporary Pool Asibanglan Kalinga Luzon
12 191–192 Limos Stream Limos Kalinga Luzon
13 193–194 Laoagan Resort Fish Pond Tabuk Kalinga Luzon
14 001–007 *Lake Paoay Paoay Ilocos Norte Luzon
15 198–200 Lake Danum Sagada Mt. Province Luzon
16 194–195 Underground River Sagada Mt. Province Luzon
17 196–197 Bokong Falls Sagada Mt. Province Luzon
18 203–204 Sumaging Cave Sagada Mt. Province Luzon
19 201–202 Chico River (Downstream) Bontoc Ifugao Luzon
20 181–184 Ambuklao Dam Bokod Benguet Luzon
21 114 Pangasinan Fish Pond Burgos Pangasinan Luzon
22 112–113 Lake Pantabangan Pantabangan Nueva Ecija Luzon
23 137–144 Lake Mapanuepe San Marcelino Zambales Luzon
24 133–136 Mt. Pinatubo Crater Lake Mt. Pinatubo Zambales Luzon
25 122–125 Lake Tambo Santa Juliana Tarlac Luzon
26 116–121 Candaba Swamp Pampanga Pampanga Luzon
27 115 Marikina River Marikina Metro Manila Luzon
28 32,38 *Pasig River Metro Manila Metro Manila Luzon
29 422–424 Imus River Imus Cavite Luzon
30 28 *Laguna de Bay Several municipalities Metro Manila, Luzon
& cities Laguna & Rizal
31 20–21 *Lake Tadlak Tadlak Laguna Luzon
32 25–27 *Lake Caliraya Lumban–Cavinti–Kalayaan Laguna Luzon
33 22–24 *Lake Lumot–Mahipon Lumban–Cavinti–Kalayaan Laguna Luzon
34 17–19 *Lake Sampalok San Pablo Laguna Luzon
35 08–10 *Lake Bunot San Pablo Laguna Luzon
36 11 *Lake Yambo San Pablo Laguna Luzon
37 12,13 *Lake Mohikap San Pablo Laguna Luzon
38 14–15 *Lake Palakpakin San Pablo Laguna Luzon
39 16 *Lake Pandin San Pablo Laguna Luzon
40 425–430 Lake Calibato San Pablo Laguna Luzon
41 229–235 *Lake Taal Several municipalities & cities Batangas Luzon
42 301–302 Daet River Daet Camarines Norte Luzon
43 302–303 Bicol River Naga Camarines Sur Luzon
44 63, 66, 59 *Lake Buhi Buhi Camarines Sur Luzon
45 61,62,65 *Lake Bato Iriga Camarines Sur Luzon
46 60 *Lake Baao Baao Camarines Sur Luzon
47 304–305 Lake Danao Albay Camarines Sur Luzon
48 42,64 *Lake Bulusan Mt. Bulusan Sorsogon Luzon
49 73–78 Lake Malbato Coron Palawan Luzon
50 79–81 Lake Kayangan Coron Palawan Luzon
51 67–72 Calbiga River Calbiga Eastern Samar Visayas
52 287 Lulugayan Falls Calbiga Eastern Samar Visayas
53 288–289 *Lake Danao Ormoc Leyte Visayas
54 53 Lake Bito McArthur Leyte Visayas
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RAFFLES BULLETIN OF ZOOLOGY 2014
and reservoirs including the Philippines (Fernando, 2002).
Previous and present studies highlight almost the same
distribution pattern of the species in Luzon, Visayas and
Mindanao (Fig. 3). This study also confirms previous
records of Bosminopsis deitersi in the islands of Luzon and
Mindoro. New records for Bosminopsis deitersi were noted
in Lakes Bangalau, Bato, and Baao. Sididae genera found in
past literatures in the Philippines include Diaphanosoma (at
least four species), Latonopsis (one species) and Pseudosida
(one species). Irregularities in the presence and distribution
of Sididae in past literatures have been observed in several
present collections. Recent researches in Lake Paoay (Aquino
et al., 2008) and Lake Taal (Papa et al., 2012) noted the
presence of Latonopsis and Pseudosida but were not present
in more recently examined samples. Four species from the
genus Diaphanosoma and one in the genus Latonopsis
have been found in the course of the study. Diaphanosoma
sarsi was the most common species under the family in the
Philippines that was found in several lakes and rivers in six
major islands including Luzon, Mindoro, Leyte, Negros,
Bohol and Mindanao. The distribution of D. sarsi conrms
the study of Korovchinsky (1992) noted that the species is
common in the tropics. D. modigliani which was previously
noted by Woltereck et al. (1941) has been re-identied as
D.orghidani transmurensis by Korovchincky (2000) from
the collected samples in Lake Lanao, Mindanao. It was
mentioned that D. orghidani was seen in the south of far
east Asia, but the species was not encountered in any of
the 86 inland waters that have been sampled in this study
especially in Lake Lanao. D. excisum was identified in
35 inland waters that are mostly found in Luzon (Fig. 4).
Similar to D. sarsi, D. excisum was mostly found in lakes,
some rivers, reservoirs and sh ponds. Thus, conrming the
studies of Mamaril & Fernando (1978), Mamaril (2001) and
Papa & Zafaralla (2011) on the presence of D. excisum in
Lakes Taal, Laguna de Bay, as well as in Buguey, Cagayan
and Ilocos Norte. Other studies suggest that most of the areas
where D. excisum was present have a high acidity, turbidity
or salinity content (Korovchinsky, 2000). Laguna de Bay is
highly turbid and often has saltwater intrusions related to
the inux of saltwater from Manila Bay while Lake Taal has
Table 1. Cont'd.
Site UST ZRC Name Municipality/ Province Island
No. Ref. No. City
55 290–293 *Lake Danao Danao Cebu Visayas
56 50,52,55 Malinao Dam Pilar Bohol Visayas
57 223–228 *Lake Danao Dumaguete Negros Oriental Visayas
58 53,43–45 *Lake Balinsasayaw Dumaguete Negros Oriental Visayas
59 54,49,36 *Lake Kabalin–an Mt.Talinis Negros Oriental Visayas
60 48 *Lake Naujan Naujan Oriental Mindoro Visayas
61 40–41 *Lake Mainit Mainit Agusan del Norte Mindanao
62 281 Fish Pond Magsaysay Misamis Oriental Mindanao
63 283 Agusan River Butuan Agusan del Norte Mindanao
64 277 Gingoong River Gingoong Misamis Oriental Mindanao
65 278 Cold Springs Albuig Misamis Oriental Mindanao
66 262–263 Cagayan River Kabugapit Misamis Oriental Mindanao
67 279 Rice Field Longalog Misamis Oriental Mindanao
68 264 Magkaambos Cave Magkaambos Misamis Oriental Mindanao
69 280 Kabulig River Kabulig Misamis Oriental Mindanao
70 271 Lake Danao Jasaan Misamis Oriental Mindanao
71 267–270 Lake Gumaod Jasaan Misamis Oriental Mindanao
72 266 Tagaloan River Tagaloan Misamis Oriental Mindanao
73 246 Tubod–Mayahay River Iligan City Lanao del Norte Mindanao
74 238 Tumuga Bridge Iligan City Lanao del Norte Mindanao
75 239–241 Ma. Cristina Falls Iligan City Lanao del Norte Mindanao
76 244–245 Agos IV Dam Iligan City Lanao del Norte Mindanao
77 236 Tinago Falls Iligan City Lanao del Norte Mindanao
78 73–105 Lake Lanao Marawi Lanao del Sur Mindanao
79 260–261 Lake Napalit Pangantucan Bukidnon Mindanao
80 247–250 Lake Apo Valencia Bukidnon Mindanao
81 251–254 Lake Pulangi Valencia Bukidnon Mindanao
82 258–259 Lake Tutay Don Carlos Bukidnon Mindanao
83 255–257 Lake Pinamaloy Don Carlos Bukidnon Mindanao
84 47 *Lake Siloton General Santos South Cotabato Mindanao
85 39 *Lake Lahit General Santos South Cotabato Mindanao
86 37 *Lake Sebu General Santos South Cotabato Mindanao
*Samples deposited in the UST–ZRC collected between 2006 and 2012
792
Pascual et al.: Freshwater Cladocera of the Philippines
high conductivity due to its volcanic nature (>1800 μS cm-1)
(Papa et al., 2011). Meanwhile, the town of Buguey (Cagayan)
has many brackishwater marshes owing to its proximity to
the coast. Interestingly, D. tropicum which was previously
described in Lake Taal (Papa & Zafaralla, 2011; Papa et al.,
2012) is now found in 11 other lakes in Luzon Island. D.
dubium which was only described in 2000 by Korovchinsky
from Isabela province is at present found in 10 lakes in
Luzon and one river in the island of Mindanao. The result
suggests new localities where most of the species from this
family were found (Fig. 4). The presence of D. dubium in
Luzon supports the record of Korovchinsky (2000). We also
conrm the presence of Latonopsis australis described by
Brehm (1938) and Mamaril & Fernando (1978) in the north,
central and southern Philippines (See Fig. 7–8). Among all
the three families in order Anomopoda, family Sididae has the
most number of species found in the Philippine archipelago.
The largest family of anomopods, Chydoridae includes
more than 200 described species. There are two subfamilies
recognised: Chydorinae and Aloninae. Family Chydoridae
includes the largest cladoceran genus Alona found in sub-
Antarctic to the tropics sometimes restricted to ancient
lakes and are often highly specialised. Among ctenopods,
chydorids show a wider range of morphological differences
between parthenogentic females and the rare male population
(Smirnov, 1967). Several species can even be found in cave
water springs and ground water. From a total of 28 species
that were reported in previous literatures by Brehm (1938),
Woltereck et al. (1941), Tsi-Chung & Clemente (1954),
Mamaril (1978, 2001), Petersen & Carlos (1984); Fernando
(1980), Korovchinsky (2000), Aquino et al. (2008), Papa
et al. (2011), Papa and Zafaralla (2011), Papa et al. (2012),
only seven species belonging to seven genera have been
encountered. This may be attributed to recent anthropogenic
impacts to the littoral zones of many freshwater ecosystems.
Littoral zones are the rst avenue of nutrient introduction from
various sources (Papa et al., 2008). Chydorids are sensitive
to physic-chemical parameters of water that they prone to
the effect of eutrophication. The clearing of macrophytes
that serve as habitat and protection from predators are being
cleared from several sampled inland water bodies. This
increases predation pressure on littoral zooplankton. Loss of
large chydorid species (Leydigia, Euryalona etc.) can also be
attributed to this. Introduction of aquaculture in lakes may
have also increased both predation pressure as well as the
overall water quality of freshwater ecosystems.
Zooplanktivorous shes both introduced and native have
larval stages that stay in the littoral zones thus increase
in their number will reduce their population (Papa et al.
2008). Several genera also have to be further re-assessed.
The genus Alona itself had been divided into several other
genera (Karualona, Coronatella, etc.) in the past few years.
The Anthalona sp. found in Lake Lanao, Lake Calig and
Chico River is probably an undescribed species due to
varying morphological characters such as the structure of its
P1 and other limb structuires but further analysis is needed
to determine its species Its terminal claw looks reminiscent
of a poorly described species, A. alonopsiformis by Brehm
in Lake Dagiangan in Lanao del Sur. Recent revisions on
their taxonomy and evaluation of morphological structures
by use of better microscopy may have led to a decline in
species richness as many species are already synonyms or
transferred to other genera (Frey 1982; Van Damme et al.,
2011). Still plenty of the genera are still questionable and
are still under a species group status like C. hermanni which
is considered as nearly invalid or vague (Korovchinsky,
2013). The confusion on the taxonomy of chydorids also
adds to the difculty in the mapping of their distribution.
The last published study on chydorids (Mamaril, 2001) has
only re-listed the species and no further re-evaluation of
morphological structures has been done. The lack of update
on the taxonomy and distribution among chydorids had
been a handicap in assessing the status of their presence in
Philippine inland waters (Papa et al., 2012).
Our results suggests that anthropogenic activities in inland
waters may serve as a major factor in the loss of its
zooplankton diversity since most freshwater ecosystems
sampled in the study were used for aquaculture. Furthermore,
selective predation by aquaculture shes may also affect the
diversity of large zooplankton (Cózar, 2003) like moinids
and sidids. Removal of macrophytes in the water may also a
reason of the diminished diversity of small cladocerans like
bosminids and chydorids in the Philippines.Cosmopolitan
records should be carefully examined because many of the
described taxa represent different species.
Our study presents an overview of cladoceran species
richness in inland waters of the three major island groups
in the Philippine archipelago. The 16 species identied from
more sampling sites compared to previous studies, is far less
than the previously reported 56 species. Less species were
encountered in most freshwaters in the major islands of
Visayas in comparison to the cladoceran species richness in
Luzon and Mindanao with families Sididae and Chydoridae
more distributed throughout the archipelago. Passive dispersal
of cladocerans through channels, rivers or through aquaculture
may also be one of the primary factors for higher number
cladoceran species in the Luzon and Mindanao islands.
Furthermore, freshwater aquaculture is also less practiced in
the Visayas due to its greater dependence on marine sheries
products. This may have inhibited the passive transport of
native and / introduced zooplankton from the larger islands
through aquaculture. Though no novel species have been
identied from these four families so far, our study already
provides a better understanding of distribution patterns as
we have collected samples from areas where no previous
records on freshwater zooplankton exist which led to the
addition of 78 new locality records for families Moinidae,
Bosminidae, Chydoridae and Sididae.
ACKNOWLEDGEMENTS
Funding for eld work was provided in part by the Research
Center for the Natural and Applied Sciences (RCNAS) of the
University of Santo Tomas and the Institute of Hydrobiology,
Jinan University. The scholarships and theses grants of
the rst two authors were provided by the Department of
793
RAFFLES BULLETIN OF ZOOLOGY 2014
Science and Technology – Acceleration of Science and
Technology Human Resource and Development Program
(DOST-ASTHRDP). The research visit of the first two
authors to the Institute of Hydrobiology, Jinan University
were also made possible through an additional grant from the
Institute of Hydrobiology, Jinan University. Samples stored
in the UST-ZRC were collected with the help of DMA Dela
Cruz-Papa, JCA Briones, DT Tordesillas, Huiming Li, Xiojun
Jiang and undergraduate Biology students of the University
of Santo Tomas under the supervision of the corresponding
author (RDS Papa).
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