Comp. Parasitol. 78(2), 2011, pp. 359–366

Helminths from 9 Species of (: ) from Sri Lanka

1,6 2 3 4 STEPHEN R. GOLDBERG, CHARLES R. BURSEY, AARON M. BAUER, ANSELM DE SILVA, AND 5 CHRISTOPHER C. AUSTIN 1 Department of Biology, Whittier College, Whittier, California 90608, U.S.A. (e-mail: [email protected]), 2 Department of Biology, Pennsylvania State University, Shenango Campus, Sharon, Pennsylvania 16146, U.S.A. (e-mail: [email protected]), 3 Department of Biology, Villanova University, Villanova, Pennsylvania 19085, U.S.A. (e-mail: [email protected]), 4 Amphibia and Research Organization of Sri Lanka, 15/1 Dolosbage Road, Gampola, Sri Lanka (e-mail: [email protected]), and 5 Department of Biological Sciences and Museum of Natural Science, Louisiana State University, Baton Rouge, Louisiana 70803, U.S.A. (e-mail: [email protected])

ABSTRACT: Nine species of geckos from Sri Lanka were examined for helminths: Calodactylodes illingworthorum, Cnemaspis kandiana, Cnemaspis scalpensis, Cnemaspis tropidogaster, Gehyra mutilata, parvimaculatus, Hemidactylus depressus, Hemidactylus frenatus, and Hemidactylus leschenaultii. Species found include 1 species of Digenea, Mesocoelium monas; 2 species of Cestoda, Cylindrotaenia philauti and Oochoristica paurensis; 8 species of Nematoda, Parapharyngodon maplestoni, Pharyngodon gekko, Pharyngodon oceanicus, Physaloperoides dactyluris, Skrjabinelazia hemidactyli, Spauligodon hemidactylus, Thamugadia hemidactyla, and Acuariidae gen sp. (larva in cysts); and 1 species of Acanthocephala, Acanthocephalus serendibensis. Mean number of helminth species per infected was 1.3 6 0.5 (range 1–3). Nineteen new hosts and 7 new locality records are reported. KEY WORDS: helminths, Digenea, Cestoda, Nematoda, Acanthocephala, geckos, Sri Lanka.

There have been few reports of helminths in Sri 1877), and the rough-bellied gecko Cnemaspis tropi- Lankan geckos: von Linstow (1906) described a dogaster (Boulenger, 1885). Manamendra-Arachchi et nematode, Oxyuris megaloon (currently Parapharyn- al. (2007) recently described many new species, godon species inquirenda), from Hemidactyulus restricted the application of the names C. kandiana leschenaultii; Crusz and Ching (1975) reported the and C. scalpensis to specimens from limited geograph- acanthocephalan Acanthocephalus serendibensis in ic areas within the island, and considered the Cnemaspis kandiana; Breckenridge and Nathanael provenance of C. tropidogaster sensu stricto to be (1983) studied the morphology of acanthocephalans uncertain. Both the geographic distribution and range obtained from Cnemaspis kandiana; Crusz (1984) of morphological variability of most of the newly found a digenean, Paradistomum sp., in Hemidactylus described taxa remain incompletely known, and it is parvimaculatus; and de Silva et al. (2004) reported probable that our samples of Cnemaspis are composite unidentified strongyloid nematodes in Calodactylodes under their interpretation. Only our Central Province C. illingworthorum and Hemidactylus maculatus. kandiana specimens are unambiguously assignable to Herein, we report helminths in 9 species of geckos the named species as narrowly construed. The from Sri Lanka. Six of these species are Sri Lankan remaining specimens cannot confidently be assigned endemics: Sri Lankan golden gecko, Calodactylodes to species within the framework of Manamendra- illingworthorum Deraniyagala, 1953, a large lizard Arachchi et al. (2007), but they are consistent with the found in rocky outcrops of the dry zone of Sri Lanka; names as used in the literature prior to 2007 (e.g., Das Kandyan gecko, Hemidactylus depressus Gray, 1842, a and de Silva, 2005). The other species have wider rupicolous gecko that occurs on the plains; spotted distribution patterns: stump-tailed gecko, Gehyra house gecko, Hemidactylus parvimaculatus Deraniya- mutilata Wiegmann, 1834, ranging from India east to gala, 1953, a widely distributed species common in Myanmar, Indo-China, the Malay Peninsula, and disturbed habitats; and three morphospecies of the Indonesia to Oceania; common house gecko, Hemi- genus Cnemaspis, which we refer to as the Kandy day dactylus frenatus (Dume´ril and Bibron, 1836), which is gecko, Cnemaspis kandiana (Kelaart, 1852), Fergu- widespread in Sri Lanka and southern and Southeast son’s day gecko, Cnemaspis scalpensis (Ferguson, Asia, and has been introduced into almost every part of the tropics and subtropics; and Leschenault’s leaf-toed gecko, Hemidactylus leschenaultii Dume´ril and Bi- 6 Corresponding author. bron, 1836, which occurs in the dry zones of Sri Lanka

359 360 COMPARATIVE PARASITOLOGY, 78(2), JULY 2011

Table 1. Number (N), prevalence (%), and mean intensity of infection (mean [M] ± standard deviation [SD]) for helminths in 9 species of Sri Lankan gekkonid lizards.

Calodactylodes Cnemaspis illingworthorum Cnemaspis kandiana Cnemaspis scalpensis tropidogaster N % M 6 SD N % M 6 SD N % M 6 SD N % M 6 SD

Digenea Mesocoelium monas –383–– Cestoda Cylindrotaenia philauti –– – 2916 0 Oochoristica paurensis –– –– Nematoda Parapharyngodon maplestoni –– –– Pharyngodon gekko 4 33 4 – – 1 5 1 Pharyngodon oceanicus –– –– Physalopteroides dactyluris 3 33 3 – 3 25 3 2 5 2 Skrjabinelazia hemidactyli 1 33 1 9 46 1.5 6 1.2 – 3 14 1 6 0 Spauligodon hemidactylus –– –– Thamugadia hemidactyla –– – 151 Acuariidae gen sp. (larva in cysts) – – – 10 9 5 6 4 Acanthocephala Acanthocephalus serendibensis –– – 151 but is widespread in southern Asia from eastern RESULTS Pakistan to peninsular India (Das and de Silva, 2005). We found 1 species of Digenea (Mesocoelium monas [Rudolphi, 1819] Freitas, 1957), 2 species of Cestoda MATERIALS AND METHODS (Cylindrotaenia philauti Crusz and Sanmugasunderam, Seventy-six individuals representing the following 9 species 1971 and Oochoristica paurensis Malhotra and Capoor, of geckos from Sri Lanka collected in 2002 were examined for 1984), 8 species of Nematoda (Parapharyngodon helminths: Calodactylodes illingworthorum (n 5 3, mean snout-vent length (SVL) 5 76.3 mm 6 18.5 SD, range 5 55– maplestoni Chatterji, 1933; Pharyngodon gekko [Chak- 88 mm); Cnemaspis kandiana (n 5 13, mean SVL 5 29.7 mm ravarty and Bhaduri, 1948] Petter and Quentin, 1976; 6 2.6 SD, range 5 24–33 mm); Cnemaspis scalpensis (n 5 4, Pharyngodon oceanicus Bursey & Goldberg, 1999; mean SVL 5 33.8 mm 6 1.7 SD, range 5 32–36 mm); Physaloperoides dactyluris [Karve, 1938] Chabaud and Cnemaspis tropidogaster (n 5 22, mean SVL 5 30.9 mm 6 3.7 SD, range 5 23–38 mm); Gehyra mutilata (n 5 4, mean Brygoo, 1960; Skrjabinelazia hemidactyli Shamim and SVL 5 48.0 mm 6 7.5 SD, range 5 38–56 mm); Hemidactylus Deshmukh, 1982; Spauligodon hemidactylus Bursey parvimaculatus (n 5 12, mean SVL 5 43.5 mm 6 7.3 SD, and Goldberg, 1996; Thamugadia hemidactyla Desh- range 5 33–54 mm); Hemidactylus depressus (n 5 7, mean mukh and Ali, 1966; Acuariidae gen sp. [larva in SVL 5 68.0 mm 6 4.8 SD, range 5 59–73 mm); Hemidactylus frenatus (n 5 8, mean SVL 5 49.2 mm 6 3.9 SD, range 5 45– cysts]), and 1 species of Acanthocephala (Acanthoce- 54 mm); and Hemidactylus leschenaultii (n 5 3, mean SVL 5 phalus serendibensis Crusz and Mills, 1970). Number 81.0 6 1.4 SD, range 5 80–82 mm). of helminths, prevalence (%), and mean intensity 6 SD Geckos were collected by hand, euthanized within 12 hr of are presented in Table 1. Parasite terminology is in capture, preserved in 10% formalin, and stored in 70% ethanol. The body cavity was opened by a longitudinal incision, and the accordance with Bush et al. (1997). digestive tract was removed and opened. The esophagus, In total, 192 helminths were found: 47 (24.5%) stomach, small intestine, and large intestine were examined for digeneans; 3 (1.5%) cestodes; 141 (73.5%) nematodes, helminths under a dissecting microscope. Nematodes and and 1 (0.5%) acanthocephalan. Mean number of acanthocephalans were placed on a glass slide in a drop of glycerol, a coverslip was added, and identification was made helminths per infected host was 6.2 6 13.7. Of the from these temporary wetmounts. Digeneans and cestodes helminths found, 10 (5%) were larval forms not were regressively stained in hematoxylin, mounted in balsam, capable of completing their life cycle in geckos. and examined. Locality data are given in Appendix 1. Geckos Twelve species of helminths were found; however, were deposited in the herpetology collection of the National no individual host harbored more than 3 species. Thirty Museum of Sri Lanka, Colombo, Sri Lanka. Helminths were deposited in the United States National Parasite Museum, one (40%) of 77 geckos were infected. Of these, 26 Beltsville, Maryland (Appendix 2). (84%) each harbored 1 helminth species; 38 (13%) GOLDBERG ET AL.—HELMINTHS FROM GECKOS IN SRI LANKA 361

Table 1. Extended.

Hemidactylus Hemidactylus Hemidactylus Gehyra mutilata parvimaculatus depressus Hemidactylus frenatus leschenaultii N % M 6 SD N % M 6 SD N % M 6 SD N % M 6 SD N % M 6 SD

– – – 44 13 44 –

––––– –181–––

– 1 8 1 – – 11 100 3.0 6 2.7 ––––– – – 2 14 2 – – – 3 17 1.5 6 0.7 – – 7 66 3.5 6 3.5 2 25 2 – – – – – – 76 28 38 6 41 4 13 4 – ––––– –––––

–––––

harbored 2 helminth species each; and 1 (3%) harbored Oochoristica pauriensis was described from Hemi- 3 species (mean number of helminth species per dactylus brookii and Hemidactylus flaviviridis, both infected gecko was 1.3 6 0.5; range 1–3). Mean from India (Malhotra and Capoor, 1984). To our number of helminth species per host species was 2.6 6 knowledge, it has not been reported since its 1.8 (range 1–7). discovery. Sri Lanka is a new locality record for O. pauriensis, and H. parvimaculatus (recently resur- DISCUSSION rected from the synonymy of H. brookii; Bauer et al., 2010) is a new host record. Mesocoelium monas is cosmopolitan in distribu- Parapharyngodon maplestoni was originally de- tion and requires a mollusk intermediate host scribed from the intestine of an agamid lizard, (Kennedy et al., 1987). It is known from fishes, Calotes versicolor, collected in Burma by Chatterji amphibians, and ; records are summarized in (1933). Additional hosts include the agamid Bronch- Bursey et al. (2007). Mesocoelium monas has ocela cristatella; the anguid Pseudopus apodus; the previously been found in H. frenatus from Indonesia gekkonids Cyrtodactylus louisiadensis, Hemidactylus (Killick and Beverley-Burton, 1982; Kennedy et al., flaviviridis, and H. frenatus; and the scincids Emoia 1987). Cnemapsis kandiana represents a new host longicauda, Emoia nigra, Emoia obscura, Emoia record for M. monas. Other hosts from Sri Lanka pallidiceps, Lipinia noctua, Prasinohaema virens, include Polypedates eques, Polypedates maculatus, Sphenomorphus emigrans, and Sphenomorphus jo- Ichthyophis glutinosus, Ichthyophis pseudangularis, biensis (Bursey et al., 2005a, b; Goldberg et al., 2008, and Nessia burtonii (Fernando, 1933; Crusz and 2010). Of these, Baylis (1936) lists C. versicolor and H. Santiapillai, 1982; Crusz and Daundasekera, 1988). flaviviridis as hosts from Sri Lanka for P. maplestoni; Cylindrotaenia philauti was originally described however, Crusz and Daundasekera (1988) considered from the rhacophorid frog Philautus (currently Sri Lanka a doubtful locale for P. maplestoni. Pseudophilautus) variabilis, collected in Sri Lanka Hemidactylus parvimaculatus and Hemidactylus (Crusz and Sanmugasunderam, 1971). It apparently leschenaultii represent new host records for P. has not been reported since its discovery. Our maplestoni. Baylis’s (1936) record of H. flaviviridis specimens were not well preserved, but they do fit probably actually pertains to the somewhat similar H. the original description. Cnemaspis tropidogaster leschenaultii, because H. flaviviridis does not occur in represents a new host record for C. philauti. Sri Lanka (Somaweera and Somaweera, 2009). 6 OPRTV AAIOOY 82,JL 2011 JULY 78(2), PARASITOLOGY, COMPARATIVE 362

Table 2. Reported helminths from 11 species of geckos from Sri Lanka.

Gecko Calodactyloides Cnemaspis Cnemaspis Cnemaspis Gehyra Hemidactylus Hemidactylus Hemidactylus Hemidactylus Hemidactylus Hemidactylus Helminth illingworthorum kandiana scalpenis tropidogaster mutilata parvimaculatus depressus frenatus flaviviridis leschenaultii maculatus

Mesocoelium monas – 1* – – – – – 1 – – – Paradisomum sp. – – – – – 2 – – – – – Cylindrotaenia philauti –––1––––––– Oochoristica paurensis –––––1––––– Parapharyngodon maplestoni –––––1––31– Parapharyngodon megaloon –––––––––4– Pharyngodon gekko 1––1––––––– Pharyngodon oceanicus ––––––1–––– Physaloperoides dactyluris 1–11–1–––1– Skrjabinelazia hemidactyli 1––11–––––– Spauligodon hemidactylus ––––––11––– Thamugadia hemidactyla –––1––––––– Acuariidae gen. sp. – – – 1 – – – – – – – Strongylidae gen. sp. 5 – – – – – – – – – 5 Acanthocephalus serendibensis –6–1–– – ––––

* 1. This study; 2. Crusz (1984); 3. Baylis (1936); 4. von Linstow (1906); 5. de Silva et al. (2004); 6. Crusz and Ching (1975). GOLDBERG ET AL.—HELMINTHS FROM GECKOS IN SRI LANKA 363

Pharyngodon gekko, originally Neopharyngodon Acuariid nematodes inhabit the gizzard of terres- gekko, was described from specimens taken from the trial birds; insects are intermediate hosts (Anderson, large intestine of Gekko gecko collected in Calcutta, 2000). Lizards presumably are paratenic (transport) India, by Chakravarty and Bhaduri (1948) and hosts, and development does not go beyond the larval reassigned by Petter and Quentin (1976). It appar- stage. This is the first report of acuariid larvae in Sri ently has not been reported since its discovery. Lankan geckos. Calodactylodes illingworthorum and Cnemaspis aff. Acanthocephalus serendibensis was originally tropidogaster represent new host records for P. described from specimens taken from the agamid gekko; Sri Lanka is a new locality record. Pharyngodon Ceratophora stoddarti, collected in Sri Lanka by oceanicus was described from the gecko Gehyra Crusz and Mills (1970), and was redescribed from oceanica from the Cook and Society Islands by Bursey specimens taken from the same host, as well as the and Goldberg (1999). There is an additional report of P. gekkonid Cnemaspis aff. kandiana, both host species oceanicus in G. oceanica collected in Moorea (Gold- collected in Sri Lanka (Crusz and Ching, 1975). berg et al., 2000). Hemidactylus depressus represents a Cnemaspis aff. tropidogaster represents a new host new host record for Pharyngodon oceanicus; Sri Lanka record for A. serendibensis. is a new locality record. Bush et al. (1997) presented a hierarchy of parasite Physalopteroides dactylurus, originally Thubu- community terms, including infracommunity (hel- naea dactyluris, was described from specimens taken minths in a single host), component community from Hemidactylus flaviviridis and Calotes versicolor (helminths of a host species are presented in Table 1), by Karve (1938) and reassigned to Physalopteroides and supracommunity (helminths in sympatric hosts). by Chabaud and Brygoo (1960). Additional hosts For this discussion, we will consider the supracom- include H. frenatus and a viperid snake, Echis munity to be Sri Lankan geckos. Helminths have carinatus, from India and a skink, Eumeces (currently been classified as core and secondary species Eurylepis) taeniolatus, from Turkmenistan (Baker, according to their prevalence (P): Species with 1987). Calodactylodes illingworthorum, Cnemaspis prevalences .30% are deemed core species (Roca, aff. scalpensis, C. aff. tropidogaster, H. parvimacu- 1993); species with 10–30% prevalence are consid- latus, and H. leschenaultii represent new host records ered secondary species (Hanski, 1982; Roca, 1993). for P. dactylurus; Sri Lanka is a new locality record. Within the supracommunity, no helminth species was Skrjabinelazia hemidactyli was described from found to represent a core species; Skrjabinelazia specimens taken from Hemidactylus brookii collected hemidactyli represents a secondary species (P 5 14). in Marathwada, Maharashtra, India, by Shamim and However, for the various component communities, Deshmukh (1982). It, to our knowledge, has not been the following species represent core species: Phar- reported since its discovery. Calodactylodes illing- yngodon gekko, Physalopteroides dactyluris, and worthorum, C. tropidogaster, and H. leschenaultii Skrjabinelazia hemidactyli in C. illingworthorum; represent new host records for S. hemidactyli; Sri Skrjabinelazia hemidactyli in C. kandiana; and P. Lanka is a new locality record. maplestoni in H. leschenaultii (see Table 1 for P). Spauligodon hemidactylus was described from H. The following species represent secondary species frenatus from 11 islands in Oceania by Bursey and within component communities: Physalopteroides Goldberg (1996). It has been reported in the dactyluris in C. scalpensis; Skrjabinelazia hemidac- gekkonids Cosymbotus (currently Hemidactylus) tyli in G. mutilata; Spauligodon hemidactylus in H. platyurus, Gehyra mutilata, Hemidactylus garnotii, depressus; and Physalopteroides dactyluris in H. Hemiphyllodactylus typus, and Lepidodactylus lugu- leschenaultii (see Table 1 for P). bris (Goldberg and Bursey, 2001, 2002). Hemidacty- At this point in time, one generalization can be lus depressus represents a new host record for S. made: Sri Lankan geckos are infected by generalist hemidactylus; Sri Lanka is a new locality record. helminth species, that is, species capable of infecting Thamagudia hemidactylia was described from two or more host species (Table 2). One question that specimens taken from subcutaneous tissues of Hemi- remains to be answered is: Why do differential rates of dactylus giganteus collected in India by Deshmukh infection occur within these hosts? Is it related to the and Ali (1966). It apparently has not been reported small sample of hosts examined or are there ecological since its description. Cnemaspis tropidogaster repre- or physiological factors at work? A second question is: sents a new host record for T. hemidactylia; Sri Lanka Will the remaining species of geckos, when examined, is a new locality record. harbor the same species of helminths so far reported? 364 COMPARATIVE PARASITOLOGY, 78(2), JULY 2011

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Goldberg, S. R., C. R. Bursey, and H. Cheam. 2000. C. Austin. All specimens have been deposited in the Gastrointestinal helminths of four lizard species from National Museum of Sri Lanka, Colombo (registra- Moorea, French Polynesia. Comparative Parasitology 67:118–121. tion numbers pending). Goldberg, S. R., C. R. Bursey, and F. Kraus. 2008. Calodactylodes illingworthorum (n 5 3): AMB Gastrointestinal helminths of eleven species of Emoia (Squamata: Scincidae) from Papua New Guinea. 7414, 7421, Uva Province, Monaragala District, Journal of Natural History 42:1923–1935. Serawa, Pitakumbura (07u159500N; 81u219250E); Goldberg, S. R., C. R. Bursey, and F. Kraus. 2010. AMB 7434 Eastern Province, Ampara District, Aran- Metazoan endoparasites of 14 species of skinks telawa east of Maha Oya (07 319590N; 81 269230E). (Squamata: Scincidae) from Papua New Guinea. u u Journal of Natural History 44:447–467. Cnemaspis kandiana (n 5 13): AMB 7422, 7423, Hanski, I. 1982. Dynamics of regional distribution: the core Uva Province, Monaragala District (06 569000N; and satellite species hypothesis. Oikos 38:210–221. u Karve, J. N. 1938. Some nematode parasites of lizards. Pages 81u179170E); AMB 7431, Uva Province, Monaragala 251–258 in Livro Jubilar do Professor Lauro Travassos, District, Rathatakanda Butlele (06u469000N; editado para Commemorar o 25 Anniversario de suas 81u159430E); AMB 7448, 7451, North Central Actividades Scientificas (1913–1938). Typographia do Province, Anuradhapura District, Mihintale Instituto Oswaldo Cruz, Rio de Janeiro, Brazil. Kennedy, M. J., L. M. Killick, and M. Beverley-Burton. (08u219080N; 80u309480E); AMB 7471, 7472, Cen- 1987. Life cycle studies on two Digenea, Paradistomum tral Province, Kandy District, Kandy (07u159360N; geckonum (Dicrocoeliidae) and Mesocoelium sociale 80u379110E); AMB, 7476–7479, 7484, Central (Mesocoeliidae), in geckonid lizards from Indonesia. Province, Matale District, Matale (07u319480N; Canadian Journal of Zoology 65:2491–2497. Killick, L. M., and M. Beverley-Burton. 1982. Observa- 80u379390E); AMB 7486, Central Province, Kandy tions on digeneans from lizards (Sauria) in Indonesia District, Gampola (07u099059N; 80u339050E). (Paradistomum geckonum, Mesocoelium sociale, and Postorchigenes ovatus) with a revision of Paradisto- Cnemaspis scalpensis (n 5 4): AMB 7446, 7447, mum Kossack, 1910 (Dicrocoeliidae). Canadian Jour- North Central Province, Anuradhapura District, nal of Zoology 60:2093–2106. Ritigala (08u079060N; 80u399510E); AMB 7417, Malhotra, S. K., and V. N. Capoor. 1984. On a new 7418, Uva Province, Monaragala District, Serawa, reptilian cestode Oochoristica pauriensis n. sp. from Hemidactylus brooki (Gray) and Hemidactylus flavivir- Pitakumbura (07u159500N; 81u219250E). idis (Ru¨ppell) from Garhwal Hills, U.P., India. Korean Cnemaspis tropidogaster (n 5 24): AMB 7470, Journal of Parasitology 22:99–101. Manamendra-Arachchi, K., S. Batuwita, and R. Central Province, Kandy District, Kandy Pethiyagoda. 2007. A taxonomic revision of the Sri (07u159360N; 80u379110E); AMB 7488–7490, 7492, Lankan day-geckos (Reptilia: Gekkonidae; Cnemaspis) 7493, Central Province, Kandy District, Helboda with description of new species from Sri Lanka and (07u059370N; 80u399260E); AMB 7494, Central southern India. Zeylanica 7:9–122. Petter, A. J., and J. C. Quentin. 1976. Keys to the genera Province, Nuwara Eliya District, Kondagala, Laboo- of the Oxyuroidea. Pages 1–30 in R. C. Anderson, A. kellie (07u029120, 80u429290E); AMB 7496, 7497, G. Chabaud, and S. Willmott, eds. CIH Keys to the 7499–7501, Central Province, Nuwara Eliya District, Nematode Parasites of Vertebrates. Commonwealth Upcot (06u479070N; 80u379370E); AMB 7502, Cen- Agricultural Bureaux, Farnham Royal, Slough, U.K. 30 tral Province, Nuwara Eliya District, Fairlawn Tea pp. Roca, V. 1993. Helmintofauna dels reptiles. Monografies de la Estate (06u469100N; 80u379180E); AMB 7510–7513, Societat d’Histo`ria Natural de les Balears 2:273–292. Sabargamuwa Province, Ratnapura District, Rak- Shamim, S., and P. G. D. Deshmukh. 1982. Skrjabinelazia wana-Deniyaya Road (06u279040N; 80u379370E); hemidactylus n. sp. from a reptilian host Hemidactylus AMB 7526, Southern Province, Galle District, brooki. Rivista di Parassitologia 43:469–472. Somaweera, R., and N. Somaweera. 2009. Lizards of Sri Udugama (06u129530N; 80u209230E); CCA 2460– Lanka; a Colour Guide with Field Keys. Edition 2462, Southern Province, Galle District, Kottawa Chimaira. Frankfurt am Main, Germany. 303 pp. (06u059520N; 80u189400E); CCA 2463 Southern von Linstow, O. 1906. Helminths from the collection of the Province, Galle District, 5 km south of Kottawa Colombo Museum. Spolia Zeylanica 3:163–188. (06u049480N; 80u179360E); AMB 7522, Southern Province, Galle District, Bataganwilla (06u029200N; APPENDIX 1 80u13917E); AMB 7525, Southern Province, Galle District, Kitulampitiya (06 049000N; 80 13900E). Location by district and province for 9 species of u u geckos from Sri Lanka found to harbor helminth Gehyra mutilata (n 5 4): AMB 7419, Uva parasites. Collection acronyms: AdS 5 Anselm de Province, Monaragala District, Rathagala (07u189020N; Silva, AMB 5 Aaron M. Bauer, CCA 5 Christopher 81u23911uE); AMB 7506, Central Province, Nuwara 366 COMPARATIVE PARASITOLOGY, 78(2), JULY 2011

Eliya District, Upcot (06u479070N; 80u379370E); AMB Hemidactylus leschenaultii (n 5 3): AdS 12, North 7515, Southern Province, Galle District, Nimalawa Central Province, Polonnaruwa District, Yakkure (06u03908uN; 80u259550E); AMB 7527, Southern (07u499000N; 81u019000E); AMB 7443, North Cen- Province, Galle District, Haycock (06u189480N; tral Province, Polonnaruwa District, Polonnaruwa 80u189550E). (07u559590N; 81u009290E); AMB 7444, North Cen- Hemidactylus parvimaculatus (n 5 12): AdS 36, tral Province, Polonnaruwa District, Minneriya (08 029480N; 80 549350E). Eastern Province, Ampara District, Kartivu (07u219520, u u 81u519160E); AMB 7413, Uva Province, Monaragala District, Pidenipitiya (07u139270N; 81u029170E); AMB 7426, 7427 Uva Province, Monaragala District, Gonaga- APPENDIX 2 nara (06u369530N; 81u169130E); AMB 7432, Eastern Helminths from 9 species of geckos from Sri Province, Ampara District, Tampataya (07u359260N; Lanka were deposited in the United States National 81u259380E); AMB 7438, North Central Province, Parasite Museum (USNPC), Beltsville, Maryland. Polonnaruwa District, Dimbulagala (07u529249N; Calodactylodes illingworthorum: Pharyngodon 81u099090E); AMB 7454, North Central Province, gekko (USNPC 101127), Physalopteroides dactyluris Anuradhapura District, Poonewa (08u339510N; (USNPC 101128), Skrjabinelazia hemidactyli (USNPC 80u379390E); AMB 7466, Northwestern Province, Putta- 101129). lam District, Mamburiya (07u599380N; 79u449330E); AMB 7480, Central Province, Matale District, Matale Cnemaspis kandiana: Mesocoelium monas (07u319480N; 80u379390E); AMB 7516, Southern (USNPC 101130), Skrjabinelazia hemidactyli (USNPC Province, Galle District, Nimalawa (06u039080N; 101131). 80u259550E); AMB 7520, 7521, Southern Province, Galle District, Boosa (06u049190N; 80u099460E). Cnemaspis scalpensis: Physalopteroides dactyluris (USNPC 101132). Hemidactylus depressus (n 5 7): AdS 69, Uva Province, Monaragala District, Kuruwekotha Cnemaspis tropidogaster: Pharyngodon gekko (07u059550N; 81u139030E); AMB 7440, 7441, North (USNPC 101136), Physalopteroides dactyluris (USNPC Central Province, Polonnaruwa District, Dimbula- 101137), Skrjabinelazia hemidactyli (USNPC 101138), gawa (07u529240N; 81u099090W); AMB 7445 North Thamugadia hemidactyla (USNPC 101139), acuariid Central Province, Anuradhapura District, Ritigala larvae (USNPC 101140), Acanthocephalus serendiben- (08u079060N; 80u399510E); AMB 7481 Central sis (USNPC 101141). Province, Matale District, Matale (07u319480N; Gehyra mutilata: Skrjabinelazia hemidactyli 80u379390E); AMB 7524, Southern Province, Galle (USNPC 101142). District, Kitulampitiya (06u049000N; 80u139000E); AMB 7528 Southern Province, Galle District, Hemidactylus parvimaculatus: Oochoristica paur- Haycock (06u189480N; 80u189550E). iensis (USNPC 101143), Parapharyngodon maples- toni (USNPC 101144), Physalopteroides dactyluris ( 5 6) AdS 16-A, Central Hemidactylus frenatus n : (USNPC 101145). Province, Kandy District, Gompola (07u099050N; 80u339050E); AMB 7411, Uva Province, Monaragala Hemidactylus depressus: Pharyngodon oceanicus District, Pidenipitiya (07u139270N; 81u029170E); (USNPC 101146), Spauligodon hemidactylus (USNPC AMB 7420, Uva Province, Monaragala District, 101147). Rathagala (07 189020N; 81 239110E); AMB 7452, u u Hemidactylus frenatus: Mesocoelium monas North Central Province, Anuradhapura District, (USNPC 101148), Spauligodon hemidactylus (USNPC Mihintale (08 219080N; 80 309480E); AMB 7455, u u 101149). North Central Province, Anuradhapura District, Poonewa (08u339510N; 80u289500E); AMB 7518, Hemidactylus leschenaultia: Parapharyngodon Southern Province, Galle District, Kandewatte maplestoni (USNPC 101150), Physalopteroides dac- (06u029240N; 80u129390E). tyluris (USNPC 101145).