Helminths of Ten Species of Geckos (Squamata: Gekkonidae) from Papua New Guinea, with Comparisons between Immigrant and Endemic Geckos1 Stephen R. Goldberg,2,5 Charles R. Bursey,3 and Fred Kraus 4 Abstract: Two hundred three individuals representing 10 species of gekkonid lizards from Papua New Guinea collected from 2002 to 2005 were examined for helminths: Cyrtodactylus epiroticus ðn ¼ 2Þ, C. klugei ðn ¼ 2Þ, C. loriae ðn ¼ 7Þ, C. novaeguineae ðn ¼ 3Þ, C. sermowaiensis ðn ¼ 30Þ, Gehyra mutilata ðn ¼ 22Þ, G. oceanica ðn ¼ 27Þ, Gekko vittatus ðn ¼ 41Þ, Hemidactylus frenatus ðn ¼ 29Þ, and Lepidodactylus lugubris ðn ¼ 40Þ. One species of Digenea, one species of Cestoda, 18 species of Nematoda, as well as three taxa of nematode larvae (in cysts) were found. Thirty-one new host records and six new locality (¼ country) records are reported. Prevalence in endemic geckos was significantly higher than in nonendemic geckos. The family Gekkonidae has an almost worldwide distribution and is well represented in the Papuan Region, where 47 species have been reported (Papuan Herpetofauna 2008). The purpose of this paper is to establish an initial helminth list for the Papuan endemics Cyrtodactylus epiroticus, Cyrtodactylus klugei, Cyrtodactylus loriae, Cyrtodactylus novaeguineae, and Cyrtodactylus sermowaiensis and provide additional helminth occurrences for Gehyra mutilata, Gehyra oceanica, Gekko vittatus, Hemidactylus frenatus, and Lepidodactylus lugubris. Range maps for these host geckos in Papua New Guinea are available at http://www.bishopmuseum.org/ 1 This research was supported by National Science Foundation grant DEB 0103794. This is contribution 2009-004 from the Pacific Biological Survey at the Bishop Museum. Manuscript accepted 19 April 2009. 2 Department of Biology, Whittier College, Whittier, California 90608. 3 Department of Biology, Pennsylvania State University, Shenango Campus, Sharon, Pennsylvania 16146 (e-mail: cxb13@psu.edu). 4 Bishop Museum, Department of Natural Sciences, 1525 Bernice Street, Honolulu, Hawaii 96817 (e-mail: fkraus@hawaii.edu). 5 Corresponding author (e-mail: sgoldberg@whittier .edu). Pacific Science (2010), vol. 64, no. 1:131–139 doi: 10.2984/64.1.131 : 2010 by University of Hawai‘i Press All rights reserved research/pbs/papuanherps/. The last five species have previously been reported to harbor helminths (Kreis 1938, Bursey and Goldberg 1996, Goldberg and Bursey 1997, 2000a, 2001, Goldberg et al. 1998, 2000). Comparisons are made between helminths present in immigrant versus endemic geckos. materials and methods Two hundred three individuals representing 10 species of gekkonid lizards from Papua New Guinea collected from 2002 to 2005 (Appendix 1) were examined for helminths: Cyrtodactylus epiroticus ðn ¼ 2Þ, C. klugei ðn ¼ 2Þ, C. loriae ðn ¼ 7Þ, C. novaeguineae ðn ¼ 3Þ, C. sermowaiensis ðn ¼ 30Þ, Gehyra mutilata ðn ¼ 22Þ, G. oceanica ðn ¼ 27Þ, Gekko vittatus ðn ¼ 41Þ, Hemidactylus frenatus ðn ¼ 29Þ, and Lepidodactylus lugubris ðn ¼ 40Þ. Immediately after capture geckos were killed by an overdose of sodium pentobarbital, fixed in neutral buffered 10% formalin, stored in 70% ethanol, and accessioned in the herpetology collection of the Bishop Museum ( bpbm), Honolulu, Hawai‘i. Subsequently, the body cavity was opened by a longitudinal abdominal incision and the gastrointestinal tract was removed by cutting across the esophagus and the rectum, and then shipped in a vial of alcohol to Whittier College, Whittier, California, where a detailed dissection of each tract was carried out using a dissecting microscope. 131 132 PACIFIC SCIENCE . January 2010 Helminths from individual hosts were removed to vials of 70% ethanol and later placed under a coverslip in a drop of lactophenol and allowed to clear. Nematodes were identified from these preparations. Digeneans and cestodes were washed in water, regressively stained in hematoxylin, and mounted in balsam for identification under a compound microscope. Although the helminths examined in this study were originally fixed in situ with formalin, each species exhibits unique characters that easily allow proper identification; no previously undescribed helminth species was found. Voucher specimens were deposited in the United States National Parasite Collection (usnpc), Beltsville, Maryland, and the Bishop Museum ( bpbm), Honolulu, Hawai‘i (Appendix 2). Parasite terminology is in accordance with Bush et al. (1997). Chi-square analysis, Morisita’s index, and percentage similarity were used to compare infections between endemic versus nonendemic geckos (Brower et al. 1998, Magurran 2004). results A total of 635 helminths was collected from 69 (34%) of 205 geckos. Of these, 413 (65%) were mature individuals representing one species of Digenea (Allopharynx macallisteri), one species of Cestoda (Oochoristica javaensis), and 15 species of Nematoda (Aplectana macintoshi, Maxvachonia adamsoni, Cosmocerca zugi, Maxvachonia chabaudi, Meteterakis crombiei, Moaciria chondropythonis, Moaciria moraveci, Oswaldocruzia bakeri, Parapharyngodon maplestoni, Pharyngodon novaeguineae, Pharyngodon oceanicus, Pseudorictularia dispsarilis, Skrjabinelazia machidai, Spauligodon gehyrae, Spauligodon hemidactylus). Pseudorictularia dispsarilis occurred in the stomach; Allopharynx macallisteri, Oochoristica javaensis, and Skrjabinelazia machidai were found in the small intestine. All other helminths were found in the large intestines. In total 222 (35%) were immature forms (Abbreviata sp., Ascarops sp., and Acuariidae gen. sp.) represented by larvae in cysts in the stomach wall. Although 20 species of helminths occurred in the sample, no host species harbored more than eight species (mean number of helminth species per host species was 4.1 G 2.2; range 1–8), and no individual host harbored more than four helminth species (mean number of helminth species per infected gecko was 1.5 G 0.8 SD; range 1–4). Of the 69 infected geckos, 46 (67%) harbored one helminth species, 15 (22%) harbored two species, 6 (9%) harbored three species, and two (3%) harbored four species. Number of individuals for each species and prevalence (number of infected hosts/number of hosts examined
100) as well as 29 new host and six new locality (¼ country) records are recorded in Table 1. discussion The digenean Allopharynx macallisteri was originally described from Lepidodactylus lugubris collected on Guam, Mariana Islands (Dailey et al. 1998). It has also been reported from L. lugubris collected on Rota, Mariana Islands (Goldberg et al. 1998), and Cyrtodactylus triparticus (then C. louisiadensis) from Papua New Guinea (Bursey et al. 2005a). The cestode Oochoristica javaensis was originally described from specimens taken from the small intestines of the geckos Hemidactylus platyurus (then Cosymbotus platyurus), Gehyra mutilata, and Hemidactylus frenatus collected in Java (Kennedy et al. 1982) and has also been reported from Gehyra oceanica and Lepidodactylus paurolepis in the Pacific realm (see Goldberg and Bursey [2002] for locality records) as well as from Sphenomorphus jobiensis from Papua New Guinea (Bursey et al. 2005b). Six (33%) of the 18 nematodes found in this study are currently known only from Papua New Guinea (i.e., Cosmocerca zugi, Maxvachonia adamsoni, Meteterakis crombiei, Moaciria chondropythonis, M. moraveci, and Pharyngodon novaeguineae). Host lists for these species are given in Table 2. One of the 18 species, Pseudorictularia disparilis, is currently known only from the Australo-Papuan Region. It was originally described as Rictularia disparilis from specimens taken from a skink, Pseudemoia entrecasteauxi (formerly Lygosoma entrecasteauxi), collected in Australia by IrwinSmith (1922) but reassigned to its current TABLE 1 Helminth Taxa N % N % N % N % N % N % N % Digenea Allopharynx macallisteri Dailey, Goldberg & — — — — — Bursey, 1998 Cestoda Oochoristica javaensis Kennedy, Killick & — — — — — Beverley-Burton, 1982 Nematoda Aplectana macintoshi (Stewart, 1914) — *16 100 — *91 55 — Cosmocerca zugi Goldberg, Bursey & — — — *1 3 — Kraus, 2005 Maxvachonia adamsoni Moravec & Sey, — *1 50 — — — 1990 yMaxvachonia chabaudi Mawson, 1972 — — — *6 10 — Meteterakis crombiei Bursey, Goldberg & — *3 100 — *28 10 — Kraus, 2005 Moaciria chondropythonis Gibbons, 1979 — — *1 14 — — Moaciria moraveci Goldberg, Bursey & — — — *2 6 — Kraus, 2007 Oswaldocruzia bakeri Moravec & Sey, — — — — — 1986 Parapharyngodon maplestoni Chatterji, 1933 — *2 50 — *9 13 — Pharyngodon novaeguineae Bursey, Goldberg & *125 50 — — — *26 33 Kraus, 2008 yPharyngodon oceanicus Bursey & Goldberg, — — — — — 1999 Lepidodactylus lugubris Hemidactylus frenatus Gekko vittatus Gehyra oceanica Gehyra mutilata Cyrtodactylus novaeguineae Cyrtodactylus sermowaiensis Cyrtodactylus loriae Cyrtodactylus klugei Cyrtodactylus epiroticus Number of Individuals (N), Prevalence (%), New Host Record (*), and New Locality (¼ Country) Record (y) for Helminths in 10 Species of Geckos from Papua New Guinea N % N % N % — — *8 2 — 11 2 — — *7 7 26 14 — — — — — — — — — — — — — — — *1 2 — — 4 — — — — — — — — — — — — — — — — — — — — — — — — — — — — — *1 — — — — — 3 4 *2 4 7 2 134 PACIFIC SCIENCE . January 2010 Helminth Taxa Pseudorictularia disparilis (Irwin-Smith, 1922) ySkrjabinelazia machidai Hasegawa, 1984 ySpauligodon gehyrae Bursey & Goldberg, 1996 ySpauligodon hemidactylus Bursey & Goldberg, 1996 Abbreviata sp. yAscarops sp. Acuariid gen. sp. N % % N % % N % N % N % N % Lepidodactylus lugubris Hemidactylus frenatus Gekko vittatus Gehyra oceanica Gehyra mutilata Cyrtodactylus novaeguineae Cyrtodactylus sermowaiensis N N % N % — — — — — — — — — — — — — — 4 18 — — — — — — — — — — — — — — — — — — *60 26 *93 6 — *2 50 N Cyrtodactylus loriae Cyrtodactylus klugei Cyrtodactylus epiroticus TABLE 1 (continued) *4 100 *13 100 — — — — status by Dollfus and Desportes (1945). The host list for P. dipsarilis includes the frogs Litoria inermis, L. nigrofrenatus, Rana daemeli, and R. supragrisea; the lizards Emoia caeruleocauda, E. kordoana, E. physicae, Pogona mitchelli, P. microlepidota, and Tiliqua occipitalis; and the mammal Dasyurus hallucatus (Owen and Moorhouse 1980, Jones 1986, Bursey et al. 2008a, Goldberg et al. 2008). Eight of the 18 species of nematodes as well as the three genera represented by larvae in cysts have much broader distribution patterns. Hosts lists for Aplectana macintoshi, Oswaldocruzia bakeri, and Abbreviata sp. (larvae) are given in Bursey et al. (2008a); for Maxvachonia chabaudi, Parapharyngodon maplestoni, Spauligodon gehyrae, Skrjabinelazia machidai, and Ascarops sp. in Goldberg and Bursey (2002); for Pharyngodon oceanicus and Spauligodon hemidactylus in Goldberg et al. (2005); and for Acuariidae gen. sp. in Goldberg et al. (2008). Additions to these host lists are given in Table 1. 5 4 — — — 5 *1 4 — 29 34 — *4 33 — *3 11 *25 10 *2 3 — — — — — — *2 5 *3 4 — *12 3 — — — 1 Eight (40%) of the 20 helminth species found in this study (i.e., Allopharynx macallisteri, Oochoristica javaensis, Meteterakis crombiei, Pseudorictularia disparilis, Skrjabinelazia machidai, Abbreviata sp., Ascarops sp., and acuariid larvae) require an intermediate host (Macy 1965, Conn 1985, Anderson 2000). These species were represented by 277 (44% of total) individuals and were most likely acquired by ingestion of an insect intermediate host. It should be noted that 222 (80%) of the 277 individuals requiring an intermediate host were larvae in cysts; the geckos most likely serve as paratenic hosts. The remaining 14 species, represented by 358 (56% of total) individuals, are thought to have direct life cycles (Anderson 2000). The data presented here indicate that helminth parasites (specifically those capable of reaching maturity) of Papua New Guinea geckos are largely generalists, the majority of which (357 individuals versus 55) have direct life cycles. TABLE 2 Host Lists for Six Helminth Species Known Only from Papua New Guinea Nematode Cosmocerca zugi Type Host Additional Hosts Gecko, Cyrtodactylus louisiadensis Gecko, Cyrtodactylus sermowaiensis Skink, Sphenomorphus jobiensis Maxvachonia adamsoni Hylid frog, Litoria infrafrenata Gecko, Cyrtodactylus klugei Gecko, Cyrtodactylus louisiadensis Skink, Emoia jakati Skink, Emoia longicauda Skink, Emoia obscura Skink, Emoia pallidiceps Gecko, Lepidodactylus lugubris Skink, Sphenomorphus jobiensis Meteterakis crombiei Skink, Sphenomorphus jobiensis Ranid frog, Platymantis boulengeri Ranid frog, Platymantis schmidti Gecko, Cyrtodactylus klugei Gecko, Cyrtodactylus sermowaiensis Skink, Emoia battersbyi Skink, Emoia obscura Skink, Emoia pallidiceps Skink, Emoia physicae Skink, Emoia popei Skink, Sphenomorphus simus Moaciria chondropythonis Python, Morelia viridis Moaciria moraveci Microhylid frog, Hylophorbus cf. rufescens Pharyngodon novaeguineae Gecko, Cyrtodactylus louisiadensis Gecko, Cyrtodactylus loriae Gecko, Cyrtodactylus sermowaiensis Gecko, Cyrtodactylus epiroticus Gecko, Cyrtodactylus novaeguineae Reference Bursey et al. (2005a) This paper Bursey et al. (2005b) Moravec and Sey (1990) This paper Bursey et al. (2005a) Goldberg et al. (2008) Goldberg et al. (2008) Goldberg et al. (2008) Goldberg et al. (2008) This paper Bursey et al. (2005b) Bursey et al. (2005b) Goldberg et al. (2009) Goldberg et al. (2009) This paper This paper Goldberg et al. (2008) Goldberg et al. (2008) Goldberg et al. (2008) Goldberg et al. (2008) Goldberg et al. (2008) Bursey et al. (2007a) Jones (1983) This paper Bursey et al. (2007b) This paper Bursey et al. (2008b) This paper This paper 136 Five species of geckos examined in this study (Cyrtodactylus epiroticus, C. klugei, C. loriae, C. novaeguineae, and C. sermowaiensis), currently known only from New Guinea and the immediately adjacent islands, are represented by 45 individuals and account for 22% of the host sample. The other five species (Gehyra mutilata, G. oceanica, Gekko vittatus, Hemidactylus frenatus, and Lepidodactylus lugubris) are widely distributed in Oceania (Bauer and Henle 1994). Six species of helminths found in this study (Cosmocerca zugi, Maxvachonia adamsoni, Meteterakis crombiei, Moaciria chondropythonis, M. moraveci, and Pharyngodon novaeguineae), also known only from Papua New Guinea, are represented by 188 individuals and account for 30% of the helminth sample. The endemic gecko species as a group harbored 12 helminth species (Table 1). The more widely distributed gecko species as a group also harbored 12 helminth species; however, only four helminth species were common to both groups (Sorensen similarity index ¼ 0.33). The endemic geckos as a group harbored 485 helminth individuals; 30 (67%) of 45 were infected, a mean helminth load of 16.9 per infected gecko. The wide-ranging geckos as a group harbored 150 helminth individuals; 40 (25%) of 160 were infected, a mean helminth load of 3.8 per infected gecko. The helminth loads of endemic versus nonendemic geckos are significantly higher ( w 2 ¼ 35:98, df ¼ 1, P < :001). Twenty taxa of helminths were found in this study. Of the helminth species known only from Papua New Guinea, only one species, Maxvachonia adamsoni, was found in the group of geckos of wider distribution. Thus, the wide-ranging group of geckos harbored 12 helminth species but just one individual of a helminth species known only from Papua New Guinea and 149 helminth individuals of wider distribution. Six of the 14 helminth species of wider distribution (Aplectana macintoshi, Maxvachonia chabaudi, Parapharyngodon maplestoni, Pseudorictularia disparilis, Abbreviata sp., and Ascarops sp.) were found in the endemic gecko group. Thus, the endemic gecko group harbored 12 helminth species, 185 helminth individuals of species known PACIFIC SCIENCE . January 2010 only from Papua New Guinea and 300 helminth individuals of species with wider distribution. Similarity indices, Morisita’s index and percentage similarity, were calculated for the helminth communities harbored by endemic geckos (community A) and by the more widely ranging geckos (community B): Morisita’s index ¼ 0.184; percentage similarity ¼ 19.3%. Morisita’s index of community similarity is based upon Simpson’s index of dominance (DA and DB ) and is defined as ð2SXi Yi Þ/ðDA þ DB ÞðSXi SYi Þ, where Xi ¼ number of individuals in taxon i in community A, Yi ¼ number of individuals in taxon i in community B. Percentage similarity ¼ S(lower of the two percentages for each taxon). Each index ranges from 0 (no similarity) to 1 (identical) (Brower et al. 1998). Thus, there is little similarity in the composition of the helminth community in these two groups of geckos. Questions arise: Is helminth community change driven by the helminths of immigrant hosts; are immigrant hosts less susceptible to native helminths; are native hosts more quickly infected by immigrant helminths? It is well known that hosts can escape much of their native parasite community via human-assisted transport (Mitchell and Power 2003, Torchin et al. 2003, Torchin and Mitchell 2004), and Gehyra mutilata, G. oceanica, Hemidactylus frenatus, and Lepidodactylus lugubris have all been widely moved via human activities (Kraus 2009). It is unknown how recently these species arrived in the Papuan Region, but their wide and actively expanding distributions elsewhere suggest that it may have been relatively recently. Goldberg and Bursey (2000b) have also shown that lizards such as Anolis sagrei are capable of transporting helminth species from their point of origin (Caribbean) to areas they colonize (Hawai‘i); but corresponding information on infections in immigrant lizards is not yet available to make conclusions about rates of helminth community change brought about by invading hosts. Nevertheless, our data are consistent with a hypothesis of infection of endemic species by helminths transported by recently immigrating hosts. Gecko Helminths from Papua New Guinea . Goldberg et al. acknowledgments We thank E. Teodoro, T. Doleck, and S. Goldsberry for assistance with dissections. Innumerable people assisted with the fieldwork, and we thank the inhabitants and landowners of the areas listed in Appendix 1. Particular thanks go to Jim Anamiato, Sedeka Andrew, Ilaiah Bigilale, Lydia Doni, Noel Doni, Fred Francisco, Telen Garra, Charlie Graham, Bulisa Iova, Conrad Kembwa, Dyson Libai, Fred Malesa, David Mitchell, Florence Paisparea, Pesto Roberts, Bendo Salepuna, John Sengo, Stanley Simalken, John Slapeinsky, Ben Yawi, and Isodore Yidika for providing logistical assistance. We thank the Papua New Guinea National Museum and Art Gallery for providing in-country collaborative assistance and the Department of Environment and Conservation, National Research Institute, and Central, East New Britain, Milne Bay, Morobe, and West Sepik provincial governments for permission to conduct this research. Literature Cited Anderson, R. C. 2000. Nematode parasites of vertebrates: Their development and transmission. 2nd ed. CABI Publishing, Wallingford, United Kingdom. Bauer, A. M., and K. Henle. 1994. Familia Gekkonidae (Reptilia, Sauria). Part I. Australia and Oceania. Das Tierreich. 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Acta Parasitol. 52:142–145. ———. 2007b. A new species of Moaciria (Nematoda, Heterakidae) and other helminths in the red Mawatta frog, Hylophorbus cf. rufescens (Anura, Microhylidae) from Papua New Guinea. Acta Parasitol. 52:233–237. ———. 2008a. A new species of Proteocephalus (Cestoda: Proteocephalidae), description of the male of Desmogathiema papuensis (Nematoda; Quimperiidae), and other endoparasites in Sylvirana supragrisea (Anura: Ranidae) from Papua New Guinea. Comp. Parasitol. 75:33–48. ———. 2008b. A new species of Pharyngodon (Nematoda, Pharyngodonidae) and other helminths in Cyrtodactylus louisiadensis (Sauria, Gekkonidae) from Papua New Guinea. Acta Parasitol. 53:41–45. Bush, A. O., K. D. Lafferty, J. M. Lotz, and A. W. Shostak. 1997. Parasitology meets ecology on its own terms: Margolis et al. revisited. J. Parasitol. 83:575–583. Conn, D. B. 1985. Life cycle and postembryonic development of Oochoristica anolis (Cyclophyllidea: Linstowiidae). J. Parasitol. 71:10–16. 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Vol. 4, Springer Series in Invasion Ecology. Springer Science, New York. Kreis, H. A. 1938. Beiträge zur kenntnis paratischer nematoden. VIII. Neue parasitische nematoden aus dem Naturhistorischen Museum Basel. Zentralbl. Bakteriol. Parasitenkd. Infektionskr. 142:329–352. Macy, R. W. 1965. On the life cycle of the trematode Prosthogonimus cuneatus (Rudolphi, 1809) (Plagiorchiidae) in Egypt. Trans. Am. Microsc. Soc. 84:577–580. Magurran, A. E. 2004. Measuring biological diversity. Blackwell Publishing, Malden, Massachusetts. Mitchell, C. E., and A. G. Power. 2003. Release of invasive plants from fungal and viral pathogens. Nature (Lond.) 421:625– 627. Moravec, F., and O. Sey. 1990. Some nematode parasites of frogs from Papua New Guinea and Australia. Acta Soc. Zool. Bohem. 54:268–286. Owen, L., and D. E. Moorhouse. 1980. The description of the male Pseudorictularia disparilis (Irwin-Smith, 1922) (Nematoda, Physalopteridae). Bull. Mus. Nat. Hist. Nat., Paris, 4th ser. 2:1013–1018. Papuan Herpetofauna. 2008. http:// www.bishopmuseum.org/research/pbs/ papuanherps/lizards.html (accessed 18 November 2008). Torchin, M. E., K. D. Lafferty, A. P. Dobson, V. J. McKenzie, and A. M. Kuris. 2003. Introduced species and their missing parasites. Nature (Lond.) 421:828–630. Torchin, M. E., and C. F. Mitchell. 2004. Gecko Helminths from Papua New Guinea . Goldberg et al. Parasites, pathogens, and invasions by plants and animals. Front. Ecol. Environ. 2:183–190. Appendix 1 Gecko Specimens from Papua New Guinea Examined from the Herpetology Collection of the Bishop Museum ( bpbm), Honolulu, Hawai‘i Cyrtodactylus epiroticus Kraus, 2008 ðn ¼ 2Þ. Morobe Province: Mt. Shungol ( bpbm 18653, 18654). Cyrtodactylus klugei Kraus, 2008 ðn ¼ 2Þ. Milne Bay Province: Sudest Island ( bpbm 19739, 19740). Cyrtodactylus loriae (Boulenger, 1897) ðn ¼ 7Þ. Milne Bay Province: Siyomu Village, ( bpbm 17255), Bunisi Village ( bpbm 17256, 17257); Central Province: Fane ( bpbm 18649, 18650); Morobe Province: NW slope Mt. Shungol ( bpbm 18651, 18652). Cyrtodactylus novaeguineae (Schlegel, 1837) ðn ¼ 3Þ. West Sepik Province: Torricelli Mts., 2.9 to 3.2 km SSE Mt. Sapau summit ( bpbm 23314–23316). Cyrtodactylus sermowaiensis (De Rooij, 1915) ðn ¼ 30Þ. West Sepik Province: Parkop ( bpbm 23317–23320), 3.2 km SSE Mt. Sapau summit, Torricelli Mts. ( bpbm 23321–23337, 23339–23347). Gehyra mutilata (Wiegmann, 1834) ðn ¼ 22Þ. Milne Bay Province: Pini Range, Duabo ( bpbm 15435–15438), Wagabu Island ( bpbm 15831), Normanby Island ( bpbm 15832), Woodlark Island, Guasopa ( bpbm 17708–17711), Sudest Island, Araeda ( bpbm 19745– 19751), Rossel Island, Damunu ( bpbm 19753, 19754), Rossel Island, Cheme ( bpbm 19755), Nimowa Island ( bpbm 19756, 19758). Gehyra oceanica (Lesson, 1830) ðn ¼ 27Þ. East New Britain Province: Marmar ( bpbm 21995), Ulege ( bpbm 21996); Milne Bay Province: Wagabu Island ( bpbm 15833), Fergusson Island, Basima ( bpbm 15834), Misima Island ( bpbm 17218–17222), Woodlark Island, Guasopa ( bpbm 17712–17714), Sudest Island ( bpbm 19759–19763), Rossel Island, Damunu ( bpbm 19764– 19769), Nimowa Island ( bpbm 19970), Normanby Island ( bpbm 19771); West Sepik Province: Parkop ( bpbm 23348), Sibilanga ( bpbm 23349). Gekko vittatus (Houttuyn, 1782) ðn ¼ 41Þ. East New Britain Province: Mamar ( bpbm 21997–22001); Milne Bay Province: Misima Island ( bpbm 16851–16853), Woodlark Island, Guasopa ( bpbm 17700–17706), Sudest Island, Araeda ( bpbm 19773–19777), Rossel Island ( bpbm 19778–19784, 19786, 19787); Morobe Province: Mt. Shungol, Apele ( bpbm 18656–18661); West Sepik Province: Parkop ( bpbm 23350–23355). Hemidactylus frenatus Schlegel, 1836 ðn ¼ 29Þ. Central Province: Moroka ( bpbm 22006); East New Britain Province: Mamar ( bpbm 22002–22005); Milne Bay Province: Duabo, Pini Range ( bpbm 15441), Misima Island ( bpbm 16830–16833, 19788, 19789), Nimowa Island ( bpbm 19790–19793), Guasopa, 139 Woodlark Island ( bpbm 17715–17722); Morobe Province: Lae ( bpbm 18662), NW slope Mt. Shungol, Morobe ( bpbm 18663). Lepidodactylus lugubris (Duméril and Bibron, 1836) ðn ¼ 41Þ. East New Britain Province: Mamar ( bpbm 22007–22009); Milne Bay Province: Wagabu Island ( bpbm 15835–15839), Gau Hi Lama Island, Killerton Islands ( bpbm 15840), Misima Island ( bpbm 17728, 19795–19798), Sudest Island ( bpbm 19799, 19780), Rossel Island ( bpbm 19801, 19802, 19804, 19805, 19807, 19808, 19810–19812, 19814–19820), Nimowa Island ( bpbm 19822), Woodlark Island ( bpbm 17727– 17731); Morobe Province: Mt. Shungol, Apele ( bpbm 18665–18668). Appendix 2 United States National Parasite Collection (usnpc) and Bernice P. Bishop Museum ( bpbm) Accession Numbers for Helminths from Gecko Specimens Collected in Papua New Guinea Examined from the Herpetology Collection of the Bishop Museum Cyrtodactylus epiroticus: Pharyngodon novaeguineae (usnpc 101460), Pseudorictularia disparilis (usnpc 101461), Abbreviata sp. (usnpc 101462). Cyrtodactylus klugei: Aplectana macintoshi (usnpc 101463), Maxvachonia adamsoni (usnpc 101464), Meteterakis crombiei (usnpc 101465), Parapharyngodon maplestoni (usnpc 101466), Abbreviata sp. (usnpc 101467). Cyrtodactylus loriae: Moaciria chondropythonis (usnpc 101468). Cyrtodactylus novaeguineae: Parapharyngodon novaeguineae (lost in processing). Cyrtodactylus sermowaiensis: Aplectata macintoshi (usnpc 101469; bpbm H391), Cosmocerca zugi (usnpc 101470), Maxvachonia chabaudi (usnpc 101471), Meteterakis crombiei (usnpc 101472), Moaciria moraveci (usnpc 101473), Parapharyngodon maplestoni (usnpc 101474), Abbreviata sp. (usnpc 101475; bpbm H392), Ascarops sp. (usnpc 101476; bpbm H393). Gehyra mutilata: Skrjabinelazia machidai (usnpc 101477; bpbm H394), Spauligodon hemidactylus (usnpc 101478), acuariid larvae (usnpc 101479). Gehyra oceanica: Oswaldocruzia bakeri (usnpc 101480), Pharyngodon oceanicus (usnpc 101481), Spauligodon gehyrae (usnpc 101482), Spauligodon hemidactylus (usnpc 101483), Abbreviata sp. (usnpc 101484), acuariid larvae (usnpc 101485). Gekko vittatus: Allopharynx macallisteri (usnpc 101486; bpbm F320), Oochoristica javaensis (usnpc 101487), Maxvachonia chabaudi (usnpc 101488; bpbm H395), Parapharyngodon maplestoni (usnpc 101489), Abbreviata sp. larva (usnpc 101490; bpbm H396). Hemidactylus frenatus: Oochoristica javaensis (usnpc 101491; bpbm F321), Spauligodon hemidactylus (usnpc 101492), Abbreviata sp. (usnpc 101493), acuariid larvae (usnpc 101494). Lepidodactylus lugubris: Allopharynx macallisteri (usnpc 101495; bpbm F322), Maxvachonia adamsoni (usnpc 101496).