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Nematode Diversity of Phytotelmata of Nepenthes Spp. in Mount Hamiguitan Range Wildlife Sanctuary, Philippines

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Nematode Diversity of Phytotelmata of Nepenthes Spp. in Mount Hamiguitan Range Wildlife Sanctuary, Philippines Ghent University Faculty of Sciences Department of Biology Academic Year 2011-2013 Nematode diversity of phytotelmata of Nepenthes spp. in Mount Hamiguitan Range Wildlife Sanctuary, Philippines JOESEPH SELLADO QUISADO Promoter: Prof. Dr. WIM BERT & Thesis submitted to obtain the degree of Dr. IRMA TANDINGAN DE LEY Master of Science in Nematology Nematode diversity of phytotelmata of Nepenthes spp. in Mount Hamiguitan Range Wildlife Sanctuary, Philippines JOESEPH SELLADO QUISADO Nematology Section, Department of Biology, Faculty of Sciences, Ghent University; K.L. Ledeganckstraat 35, 9000 Ghent, Belgium [email protected]/[email protected] Summary – Nematodes from phytotelmata of Nepenthes hamiguitanensis and N. peltata in Mt. Hamiguitan, Philippines included three new species of the genera: Molgolaimus Ditlevsen 1921, Dominicactinolaimus Jairajpuri and Ahmad 1992, Tripylella (Bütschli, 1873) Brzeski & Winiszewska-Ślipińska, 1993; two known: Tylocephalus auriculatus (Bütschli, 1873) Anderson, 1966, Pelodera strongyloides (Schneider, 1860) Schneider 1866; and three uncertain species of the genera: Paractinolaimus Meyl 1957, Plectus Bastian 1865, and Anaplectus De Coninck & Schuurmans Stekhoven 1933. Measurements and illustrations are provided. Molgolaimus sp. nov. is characterized by the absence of pre-cloacal supplements, shape of the spicule with lamina widened distally, conical tail with swollen tip and without digitate prolongation, and sexual dimorphism in the shape of the cardia (elongated in male and more round in females). Moreover, a comprehensive key for the genus Molgolaimus is presented. Dominicactinolaimus sp. nov. is characterized by short body length, long tail (c = 6.6-7.6) and 6-7 pre-cloacal supplements. The generic position of Dominicatinolaimus is reaffirmed and the synonymy with Trachypleurosum or Trachactinolaimus is rejected. Tripyllela sp. nov. is morphologically close to T. iucunda but differs in the female reproductive system having reduced posterior branch. The morphology and morphometry of Tylocephalus auriculatus and Pelodera strongyloides specimens agree with the original descriptions of Anderson, 1966 and Schneider, 1866 respectively. Phylogenetic analysis of small subunit rDNA for T. auriculatus and D2-D3 expansion segment of LSU rDNA for P. strongyloides supported sister relationship with respective species sequences available in GenBank. Furthermore, Molgolaimus and Actinonema were observed from the samples which support the initial discovery of marine nematodes in the Nepenthes phytotelmata in 2008. However, detailed taxonomical identification of Actinonema is not provided due to loss of sample during the process. In addition, the presence of freshwater nematode throws a new light on a better understanding of the complex scheme of Nepenthes carnivory and enzyme production of pitcher plants. Keywords - new species, marine, freshwater, integrative taxonomy, dichotomous key, ecology 1 The Philippines is a tropical country located between 116° 40’ and 126° 34’ E longitude and 4° 40’ and 21° 10’ N. It is composed of 7107 islands and currently heads the world list of megadiversity countries, with a study concluding that its waters contain the world’s highest diversity of marine life (Heaney & Regalado Jr, 1998). About more than 510 species of land vertebrates are unique to the islands and estimates of endemism for vascular plants range from 45 % to 60%, an exceptionally higher level of endemism for a country of its size (Heaney & Mittermeier, 1997). In addition, Myers et al. (2000) includes the Philippines on the list of biodiversity hotspot for conservation priorities as it possesses exceptional concentration of endemic species facing also an exceptional loss of habitat. Nematodes are unfortunately overlooked in biodiversity programs and often not even mentioned as a terrestrial invertebrate (Hill et al., 2005). Philippine nematology in the past four decades focused on applied aspects (management) of plant parasitic nematodes and free- living nematodes are oftentimes neglected for taxonomical identification and characterization. Despite its reported biodiversity richness, nematodes remained unexplored in the Philippines. In the current study we will evaluate the diversity of nematodes inhabiting Nepenthes phytotelmata. The term “phytotelmata” comes from a Greek words phyton which means “plants” and telm which means “pond” (Maguire Jr, 1971). It is used to describe bodies of water impounded by plant parts and tree holes (Kitching, 1971). Phytotelmata comes in various forms Kitching and Pimm (1985), and Kitching (2000) grouped these into 5 major classes of habitats which include water-filled tree holes, plant axils, bromeliad tanks, bamboo internodes and pitchers. The latter includes the genus Nepenthes, a member of the family Nepenthaceae and the largest genus of pitcher plants with approximately 90 recognized species at present. Amoroso & Aspiras (2011) recorded 7 species of Nepenthes in Mt. Hamiguitan. Nepenthes are carnivorous plants, capable of modifying its leaves into a shape of cup or pitcher (Clarke & Lee, 2004). This serves as prey-trapping mechanism which features a deep cavity filled with liquid known as a pitfall trap (Król et al., 2012). These phytotelms of pitcher plants serve as a special kind of habitat for insect larvae, and various microorganisms including nematodes (Maguire Jr, 1971). Only a small number of studies have been conducted on nematode inhabitants of water held by Nepenthes spp. The most recent and detailed finding was the discovery of panagrolaimid nematode, Baujardia mirabilis (Bert et al., 2003). It was assumed that this nematode is naturally inhabiting the phytotelmata of Nepenthes mirabilis since it was found alive in several pitchers (Bert et al., 2003). The most comprehensive study done by Menzel 2 (1922) was on N. gymnamphora, who found several nematodes belonging to genera Diplogaster, Dorylaimus, Plectus and Rhabditis. He also described and considered that only Panagrellus nepenthicola is the real inhabitant of pitcher plants. A study of community structure of inhabitants of Nepenthes alata in West Sumatra recorded 25 nematode individuals but the species were unknown (Sota et al., 2006). A survey was initiated last 2010 headed by Dr. Irma Tandingan De Ley in Mt. Hamiguitan. Samples were taken from soil, litter, moss, epiphytes and from phytotelmata of Nepenthes. Unexpectedly, marine nematodes were found in the Nepenthes samples but this was never given enough attention as they thought those were just a contamination from other samples in their laboratory. This follow-up survey was initiated to provide a contribution towards the documentation of the nematodes present in Mount Hamiguitan Range Wildlife Sanctuary with an emphasis on pitcher plant phytotelmata. To verify whether nematodes associated with pitcher plants in Mount Hamiguitan are similar with the previous studies from other location; i.e. to validate the discovery of the marine nematodes from phytotelmata samples. To provide comprehensive morphological and phylogenetic information of the nematodes found in the phytotelmata. To hypothesize possible explanation how these nematodes survive if they inhabit the phytotelmata of Nepenthes which is known to produce enzymes and other compounds used to digest prey. Materials and methods CHARACTERISTIC OF THE AREA AND SAMPLING SITE Mount Hamiguitan Range is one of the wildlife sanctuaries in the Philippines, located 6°40’01” to 6°46’60” N and 126°09’02” to 126°31’01” E in the province of Davao Oriental. The mountain is bordered by bodies of water, Pacific Ocean to the east and Davao Gulf to the west (Figure 1). The 6,834-hectare total surface area of Hamiguitan is characterized by five 5 vegetation types each of these forest type harbors endemic and rare species of flora and fauna, including different species of Nepenthes (Amoroso & Aspiras 2011). 3 Figure 1. Sampling site. Mount Hamiguitan, Davao Oriental, Island of Mindanao, Philippines SAMPLE COLLECTION AND EXTRACTION Samples were obtained from three different species of Nepenthes (Nepenthes hamiguitanensis, N. peltata and N. micramphora). Collection was done by a random sampling of pitcher plants along the trail. A total of 13 sampling sites with respective coordinates, altitude (m) were chosen (Table 1). Each sampling site might represent one composite sample of all pitchers in the specific location while others represent only one pitcher. Three samples were recorded to have nematodes (Table 2). Liquid inside the pitchers of Nepenthes spp. were decanted into a 28 µm mesh sieves. The filtrates were washed using DESS (Yoder et al., 2006) then collected into a 100 ml falcon tubes and transported to Ghent University. Nematodes were isolated from filtrate using the Ludox floatation technique (Somerfield and Warwick, 1996). Specimens were kept and fixed back in DESS. Table 1. Site description of Nepenthes species samples and pH value. Sample Species of Nepenthes Coordinates Elevation (m asl1) pH 1 N. peltata 06°43.615'N 126°09.167'E 639 4.8 2 N. hamiguitanensis 06°43.481'N 126°09.133'E 639 3.6 3 N. hamiguitanensis* 06°43.481'N 126°09.133'E 639 3.9 4 N. hamiguitanensis 06°43,453'N 126°09.527'E 821 5.6 5 N. peltata 06°43,453'N 126°09.527'E 821 6.2 6 N. peltata* 06°43.014'N 126°11.004'E 1188 4.4 7 N. peltata 06°43.014'N 126°11.004'E 1188 5.8 8 N. micramphora 06°00.344'N 126°10.975'E 1606 2.7 9 N. micramphora 06°00.344'N 126°10.975'E 1606 3.5 10 N. peltata* 06°00.344'N 126°10.975'E 1606 3.1
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