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 in Taal,

Rey Donne S. Papa1, 2, 3* Dino T. Tordesillas2, 5, and Augustus C. Mamaril, Sr.4

1Department of Biological Sciences, 2Graduate School and 3Research Center for the Natural and Applied Sciences, University of Santo Tomas, Manila, Philippines 4Institute of Biology, University of the Philippines - Diliman, Quezon City, Philippines 5Department 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 and eutrophication impacts, as well as the presence of introduced species.

Key Words: Crustacean Zooplankton, Freshwater, Southeast Asia, Systematics, Tropical caldera ,

INTRODUCTION Brehm, Mesocyclops hyalinus Kiefer, Pseudodiaptomus brehmi Kiefer, Tropodiaptomus gigantoviger Brehm, T. Studies on Philippine zooplankton were formally started prasinus Brehm, and Thermocyclops wolterecki Kiefer by the description of Trochosphaera aequatorialis in 1872 have been described based on the samples collected from based on samples collected from a rice field in Mindanao the expedition. New records have also been established Island in southern Philippines (De Elera 1895; Mamaril Sr. & for many cladoceran species. Fernando 1978; Petersen & Carlos 1984). This was followed by the first accounts of zooplankton species richness during The most comprehensive publication on Philippine the Wallacea expedition (Brehm 1938; Brehm 1942; freshwater zooplankton systematics reported 125 Hauer 1941; Kiefer 1930; Kiefer 1939a; Kiefer 1939b; zooplankton species including 49 cladocerans and nine Woltereck et al. 1941). This likewise included surveys of copepods collected from the littoral and limnetic zones lakes such as , Taal, Balinsasayao, Buhi, of lakes and reservoirs as well as smaller water bodies Calibato, Danao, Lanao, Mainit, and Naujan. Measurements in the Philippines (Mamaril Sr. & Fernando 1978). This on some basic physico-chemical and morphological together with two more papers by Mamaril Sr. (1986; characteristics of the aforementioned lakes were also 2001) became the most utilized references on Philippine carried out. New species of crustacean zooplankton such freshwater zooplankton. At present, a website maintained as Alona pseudoanodonta Brehm, Diaptomus insulanus by F. Petersen of Denmark also serves as an invaluable Wright and its synonym D. sensibilis Kiefer, D. vexillifer on-line reference for those who want to study Philippine freshwater zooplankton taxonomy (Petersen 2009). Other *Corresponding author: [email protected]; significant studies on Philippine freshwater zooplankton [email protected]

243 Philippine Journal of Science Papa et al.: Taxonomic Account of Limnetic Vol. 141 No. 2, December 2012 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).

244 Philippine Journal of Science Papa et al.: Taxonomic Account of Limnetic Vol. 141 No. 2, December 2012 Zooplankton in Lake Taal, Philippines

60 μm 4 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), to 2010. We describe key morphological features by Lewis (1979); Lai et al. (1979); Petersen & Carlos (1984); examining collected samples using a combination of Tuyor & Segers (1999); Tuyor & Baay (2001); WALTER light and scanning electron microscopy. We also included et al. (2006); Aquino et al. (2008); Lazo et al. (2009); and observations on their abundance and distribution. 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. MATERIALS AND METHODS Lake Taal (13°55’-14°05'N, 120°55’-121°105'E; Altitude: 2.5 masl) has long caught the attention of scientists for Plankton were collected from six sites in Lake Taal by its unique geological origin, geographical location and making replicate vertical tows from a depth of 40 m using limnological characteristics. The lake is a 236.9 km2 an 80 µm mesh size conical plankton net (diameter = 30 caldera with 90.4 m mean and 198 m maximum depths, cm). Samplings were held monthly in 2008, four times respectively. It was formed after a series of volcanic in 2009 (May, June, August, and November) and twice in explosions separated the lake from the rest of the South 2010 (April). Samples were fixed in 5 % formalin with China Sea (Ramos 2002) with a shallow north and a Rose Bengal dye. Density was determined for each species deeper south basin partially separated by an active volcano by taking average counts of three 1 mL Sedgwick Rafter island in the middle. Since 1975, fish cage aquaculture sub samples per replicate. Specimens were sorted and has proliferated in the north basin which has begun to dissected on slides with glycerine. These were examined take its toll on water quality (Aypa et al. 2008). The using an Olympus CX21 compound microscope and drawn decline in ecosystem health due to this has prompted with the aid of an Olympus U-DA drawing attachment. renewed interest in assessing the lake’s biodiversity (Tvpl- All measurements were made using a calibrated ocular Pamb 2009). As an important component of freshwater micrometer and presented in millimetre(s). Representative ecosystems, zooplankton diversity needs to be updated to individuals from the copepod were dehydrated in ethanol keep up with the recent developments in Systematics. The series, critical point dried in a BALTEC CPD 030, fixed zooplankton composition of Lake Taal was first studied on SEM aluminum stubs, sputter coated with 5 nm Au/Pd during the Wallacea expedition (Woltereck et al. 1941) in a BALTEC SCD 030 and viewed in a Philips SEM 505. which was later updated by Ueno (1966) and Mamaril Pictures were taken with a DISS (Digital Image Scanning Sr. (2001). A report on the limnological status of Lake System) from Point Electronics. Body and appendage Taal also presented some data on zooplankton species terminology was based on Dussart and Defaye (2001) composition during the late 1980s (Zafaralla et al. 1992; and Korinek (2002). Specimens (sorted or otherwise) are Zafaralla et al. 1989). This paper updates the crustacean deposited in the Zooplankton Reference Collection of the zooplankton species richness in Lake Taal using specimens University of Santo Tomas. collected during limnetic plankton surveys from 2008

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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 (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. Class Maxillopoda crassus as T. hyalinus (Kiefer 1930; Woltereck et al. 1941). Order Cyclopoida At present, T. crassus dominated the two other copepod Family Cyclopidae species in Lake Taal and is also common to other sampling Thermocyclops crassus (Fischer, 1853) localities in the Philippines. Mean lengths of T. crassus was 0.49 mm for males and 0.58 mm for females. Specimens of This is the type species for the genus Thermocyclops. Of adult, copepodite and nauplii T. crassus were observed from the more than 50 recognized species under this genus, only samples collected throughout the year. 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 Order Calanoida species, at least for the limnetic zone. Female T. crassus has a Family Diaptomidae hammer-shaped seminal receptacle with a short and wide sac- Arctodiaptomus dorsalis (Marsh, 1907) like handle. It may be distinguished from the morphologically A. dorsalis is the largest copepod species in Lake Taal similar T. decipiens by its straight spine in the inner distal with average length of 0.97 mm for females and 0.82 mm segment of the P4 endopod, as well as the presence of hairs for males. One key characteristic of female A. dorsalis is instead of spinules on the medial margin of the P4 basipodite

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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 Similar to T. crassus, adults, copepodites and nauplii of A. exopodite 1 (Fig. 3C & D) while for males, the P5 has a dorsalis were also observed all year round in Lake Taal. lateral spine inserted proximal to the middle of the right The adults were not as abundant as those of T. crassus as of exopodite 2 where the said spine is longer than the calanoid copepods are known to be less numerous than exopodite 2 (Fig. 3B). cyclopoids in eutrophic lakes. The Wallacea papers did not mention any calanoid copepods in Lake Taal. The occurrence of a calanoid Family Pseudodiaptomidae initially identified as Tropodiaptomus vicinus (Zafaralla Pseudodiaptomus brehmi Kiefer, 1939 et al. 1989; Zafaralla 1992 cf Mamaril Sr. 2001) was Pseudodiaptomus is classified as a highly diversified reported in Lake Taal and later mentioned in Amarasinghe demersal marine genus in the Indo-Pacific region with et al. (2008), Vijverberg et al. (2008) and Papa and 77 species, 14 of which are recorded from coastal Zafaralla (2011). However, a more detailed examination environments in the Philippines (Walter et al. 2006). of specimens collected from 2008 to 2010 revealed that P. brehmi was first described by Kiefer in 1939 based the diaptomid copepods in Lake Taal were A. dorsalis and on samples collected from Lake Naujan (Mindoro Is., not T. vicinus. This was also mentioned by PAPA et al. Philippines). At present, P. brehmi is listed as a marine (2012) in their report on the invasion of A. dorsalis which species (Walter 2009) and until this study, no record of P. they found in18 of 27 lakes throughout the archipelago.

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brehmi in another freshwater environment exists. Petersen Family Daphniidae has previously classified it as endemic to Lake Naujan Ceriodaphnia cornuta Sars, 1885 (Petersen 2009). Further studies have to be undertaken to ascertain its distribution. Female specimens are This cladoceran is common to tropical and subtropical characterized by the absence of spines in the first urosome regions and is usually found in both the littoral and segment (Fig. 4B) while the male P5 has no medial pointed limnetic areas of lakes but is absent in running waters. The process in its left Re1. Specimens from Lake Taal are first recorded occurrence of C. cornuta in the Philippines rare even for samples collected from deeper locations was from the Wallacea papers which included Lake Taal for almost the entire sampling period except for those where it was listed as C. rigaudi. It is characterized by collected in April 2010 when they were fairly common. 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). Class Branchiopoda It is a highly polymorphic species, wherein the hornless Order Cladocera and horned forms appear depending on lake productivity Family Bosminidae and intensive predation by fish, respectively (Zaret 1972). Bosmina fatalis Burkhardt 1924 Lake Taal specimens had average lengths of 0.38 mm. The specimens did not possess horns, which may be due to This species is common in lakes and reservoirs in East the high productivity of the lake from the excess nutrient (including far eastern Russia) and Southeast Asia. In the inputs from aquaculture as well as natural nutrient sources Philippines, it has been reported from Is. including from the volcanic sediments. Though found in the diets of La Mesa reservoir and Laguna de Bay and in Mindanao S. tawilis it is not considered as a preferred prey item in Is. in Lakes Lanao (Mamaril Sr. 1986) and Sebu (Papa spite of its high densities in the lake (PAPA et al. 2008). unpublished). Bosmina spp., (B. fatalis and B. longirostris) It was present in samples collected from all months. 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 Family Sididae of S. tawilis from 2004 to 2005 showed that Bosmina spp. Diaphanosoma sarsi Richard, 1894 featured prominently in its diet during the northeast monsoon coinciding with a drop in copepod abundance (Papa et al. D. sarsi is very common to lakes and reservoirs in tropical 2008). B. fatalis is known for its rounded carapace with regions except in the African continent. Philippine a long rostrum with a pair of fixed antennules of variable collection localities extend from as far north as Buguey, length and shape curving ventrally. Lake Taal specimens Cagayan in northern Philippines to and had mean lengths of 0.39 mm. Monthly plankton surveys Surigao City in the southern Philippines. The genus in 2008 revealed that B. fatalis was second to Ceriodaphnia Diaphanosoma includes the largest limnetic cladocerans cornuta in terms of abundance and were present throughout collected from the Philippines where together with the entire year with highest densities in January. 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 Family Moinidae entire head. The ventral margin of its valves also has a Moina micrura Kurz, 1874 duplicature that is rounded at the distal end. The postero- ventral margin of its valve has 13-40 small spines which The cosmopolitan Moina micrura is found in a variety of diminish dorsally and has two thin posterior dorsal spines. habitats from ponds, rice fields, reservoirs and lakes (Mamaril Collected specimens had average lengths of 0.65 mm. Sr. 1986). Species from this genus together with those from D. sarsi was first reported from Lake Taal during the Diaphanosoma are considered as the replacement of Daphnia Wallacea expedition where until 1998, it was thought to in the tropics and are the preyed upon by invertebrate have been the only species from this genus to be in this predators (Mamaril Sr. 2001). M. micrura was first reported lake. They are usually found in low numbers where they from Lake Taal in 1938 as M. dubia (Brehm 1938) which is have similar frequencies of occurrence and abundance now considered as its junior synonym. Known for having with D. excisum. 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 Diaphanosoma excisum Sars, 1885 (Korinek 2002). In terms of body size, M. micrura would be Korovchinsky (1992) and Korinek (2002) remarked that the second largest cladoceran genus in the lake at 0.56 mm D. excisum is common in tropical and subtropical regions total length. M. micrura were commonly encountered from where they inhabit a wide range of environments from the samples collected in Lake Taal though never as abundant acidic, to turbid, to slightly brackish lakes. In the country, as C. cornuta or B. fatalis.

248 Philippine Journal of Science Papa et al.: Taxonomic Account of Limnetic Vol. 141 No. 2, December 2012 Zooplankton in Lake Taal, Philippines it has only recorded from a few localities which led to the structure. Furthermore, the culture of herbivorous fish belief that it is rare in this part of the tropics. Among the such as tilapia in floating fish cages and eutrophication major Philippine lakes, D. excisum has only been reported may in the long-term cause changes in the from Lake Laguna de Bay (Mamaril Sr. 2001; Mamaril community which will later have an impact on the Sr. & Fernando 1978). It is characterized by its large head zooplankton community as well. with a well-developed dorsal part and a moderate to large eye. The ventral margin of the carapace valves has a flap This is also the first time that scanning electron that joins the margin almost at a right-angle. Samples of micrographs have been used in the Philippines to augment D. excisum collected from the lake had mean lengths of light microscopy in validating micro-characters present 0.65 mm. Similar to the other two Diaphanosoma spp. in in selected species. SEM has been widely used by Lake Taal, it occurred in low densities in 2008 with highest other scientists to study zooplankton micro-characters, recorded densities at < 30 ind. / l (April). especially minute details or complex three-dimensional structures (Dussart & Defaye 2001) such as appendages, setae and setule patterns, spine and spinule ornamentation Diaphanosoma tropicum Korovchinsky, 1998 etc. Many publications have already began to include This species has only been recently separated from D. scanning electron micrographs with traditional line modigliani. The revision was done based on samples drawings in the description of novel zooplankton species collected from Sri Lanka, Malaysia, Thailand, China and (Dussart & Defaye 2001; Suarez-Morales & Elias- the Philippines. D. modigliani is believed to be endemic Gutierrez 2001; Walter et al. 2006). In this study, the use to Indonesian lakes Toba and Tempe (Korinek 2002; of SEM was useful in validating the presence or absence Korovchinsky 1998). The Philippine specimens were of certain structures or minute details such as spinule collected from Talisay, Batangas which is situated on patterns in some appendages that were otherwise more the northern shores of Lake Taal. Further examination of difficult to observe using conventional light microscopy. samples collected from our survey validated the existence Some examples are the spinules at the terminal portion of of D. tropicum in the lake. It is characterized by its large the P5 endopod of A. dorsalis (Fig. 3D), the gonophore head with a protruding dorsal part and moderate to large and spinule ornamentation in the last thoracic segment of eyes. It may be differentiated from D. modigliani by its female P. brehmi (Fig. 4A & B). The utilization of SEM strongly curved spine on the end of the upper antennal in systematic studies of crustacean zooplankton taxa branch, long ventral valve inflexion, large and fewer has been very useful in verifying certain structures that denticles in the ventro-posterior valve margin, among were otherwise more difficult to examine in detail using others (Korovchinsky 1998). It is the largest of the three conventional microscopy. Although there are still a few Diaphanosoma spp. in the lake at 0.67 mm mean length. SEM facilities in the country, the availability of cheaper D. tropicum were observed to occur in similar densities table-top SEMs that require minimal specimen preparation in the north and south basins of the lake and was least nowadays will probably lead to its wider utilization in abundant of all the cladoceran species present. 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 DISCUSSION of zooplankton biodiversity. A total of nine species (six cladocera, three copepoda) In spite of the cosmopolitan nature of many of the species of crustacean zooplankton were recorded from Lake encountered, our study yielded two interesting species for Taal (Fig. 1). Seven genera from seven families were Lake Taal which may be due to the increased sampling represented by the following number of species: effort put in this study compared to the past surveys and Cyclopidae (1); Diaptomidae (1); Pseudodiaptomidae (1); the introduction of non-indigenous species into the lake. Bosminidae (1); Daphniidae (1); Moinidae (1); Sididae The presence of Pseudodiaptomus brehmi in Lake Taal (3). The species composition of zooplankton in Lake may be considered as one of the few occasions that a Taal has been described as typically tropical, with low Pseudodiaptomid has been recorded in a freshwater diversity for limnetic zooplankton, the absence of large environment in the Philippines (Kiefer 1939a; Petersen cladocerans such as Daphnia and the existence of small- & Carlos 1984). This is made more significant by the fact bodied species (Mamaril Sr. 2001). Predation pressure that they have not been found in more recent collections from zooplanktivores such as Sardinella tawilis may have from the two other lakes where they have previously been led to the present size structure of limnetic zooplankton recorded (Laguna de Bay and Lake Naujan). An earlier in the lake (Fernando 2002; Papa et al. 2008). The paper by the senior author has listed P. brehmi as a new declining populations of this zooplanktivore in Lake Taal record for Lake Taal (Papa & Zafaralla 2011). The other may however trigger a shift in zooplankton community

249 Philippine Journal of Science Papa et al.: Taxonomic Account of Limnetic Vol. 141 No. 2, December 2012 Zooplankton in Lake Taal, Philippines interesting species found in the lake is Arctodiaptomus alterations on the littoral zones of lakes brought about dorsalis (Marsh 1907), a Neotropical invasive species. The by human-mediated changes, such as the impact of presence of A. dorsalis confirms the role of aquaculture aquaculture on limnetic zooplankton composition. Other in the introduction of non-indigenous zooplankton, as smaller freshwater ecosystems must also be sampled given the spread of A. dorsalis has been linked to the presence how deteriorated some lake ecosystems are at present as of tilapia cage culture (Papa et al. 2012). This may these places may reveal higher diversity compared to the also be the reason behind the introduction and spread limnetic zones of tropical lakes. of the Neotropical 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), ACKNOWLEDGMENTS modelled after the success of aquaculture in nearby Lake We would like to thank R. Eckmann and J. Hentschel Laguna de Bay. A. dorsalis was first recorded in the (University of Konstanz, Germany) for the use of the Philippines from Laguna de Bay in 1991 (Tuyor & Baay SEM; T. Chad Walter, H. Dumont, and M. Holynska for 2001) and since Laguna de Bay serves as the source of verifying our identification of P. brehmi, A. dorsalis and T. tilapia fingerlings that are used to stock other Philippine crassus, respectively; and Aissa Domingo for the scientific lakes, it could have easily dispersed A. dorsalis to the illustrations. The SEMs were taken during the DAAD other lakes as well. Calanoid copepods were first recorded Research Fellowship of the first author. The samplings in the lake in August 1989 (unpublished data of Mamaril were funded by the Tonolli Fund Postgraduate Fellowship Sr. 2001). As under-regulated aquaculture practices and of the International Society of Limnology, the Philippine the introduction of alien species continue in many lakes Commission on Higher Education, and the UST Research in the archipelago, more alterations to the native limnetic Center for the Natural and Applied Sciences. 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 REFERENCES Taal as littoral areas were not sampled, it more or less ALEKSEEV VR. 2002. Copepoda. In: Fernando CH, gives a complete account for the limnetic zone based on ed. A Guide to Tropical Freshwater Zooplankton the 18 sampling trips in six sampling sites that covered -Identification, Ecology and Impact on Fisheries. both open water and aquaculture areas in a span of two Leiden, the Netherlands: Backhuys Publishers. p. years (2008-2010). The difference in sampling strategies 123-128 that has to be employed for littoral zooplankton also entails a separate analysis for this group (Matsumara- AMARASINGHE PB, ARIYARANTNE MG T. Tundisi & Tundisi 2005). It is expected that the species CHITAPALAPONG, VIJVERBERG J 2008. richness of zooplankton in Lake Taal will increase after Production, Biomass and Productivity of Copepods the inclusion of littoral species, and further new records and Cladocerans in Tropical Asian Water Bodies and may be established given that it has been 26 years since the Carrying Capacity for Zooplanktivorous Fish. a comprehensive listing of littoral species (Mamaril Sr. In: SCHIEMER F, SIMON D, AMARASINGHE 1986) and 11 years since a review of zooplankton species US, MOREAU J, eds. Aquatic Ecosystems and diversity in Philippine lakes were published (Mamaril Development: Comparative Asian Perspectives. Sr. 2001). The littoral zooplankton community is highly Biology of Inland Waters Series. Leiden, The diverse, and support species that are not usually found in Netherlands: Backhuys Publishers. p. 173-194. the pelagic zone (Fernando 2002; Peters & Lodge 2010). 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