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Helminths of Ten Species of (: ) from Papua New Guinea, with Comparisons between Immigrant and Endemic Geckos1

Stephen R. Goldberg,2,5 Charles R. Bursey,3 and Fred Kraus4

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, mutilata ðn ¼ 22Þ, G. oceanica ðn ¼ 27Þ, Gekko vittatus ðn ¼ 41Þ, 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 non- endemic geckos.

The family Gekkonidae has an almost research/pbs/papuanherps/. The last five spe- worldwide distribution and is well repre- cies have previously been reported to harbor sented in the Papuan Region, where 47 helminths (Kreis 1938, Bursey and Goldberg species have been reported (Papuan Herpeto- 1996, Goldberg and Bursey 1997, 2000a, fauna 2008). The purpose of this paper is 2001, Goldberg et al. 1998, 2000). Compari- to establish an initial helminth list for the sons are made between helminths present in Papuan endemics Cyrtodactylus epiroticus, immigrant versus endemic geckos. Cyrtodactylus klugei, Cyrtodactylus loriae, Cyrto- dactylus novaeguineae, and Cyrtodactylus sermo- materials and methods waiensis and provide additional helminth occurrences for Gehyra mutilata, Gehyra oce- Two hundred three individuals representing anica, Gekko vittatus, Hemidactylus frenatus, 10 species of gekkonid lizards from Papua and Lepidodactylus lugubris. Range maps for New Guinea collected from 2002 to 2005 these host geckos in Papua New Guinea are (Appendix 1) were examined for helminths: available at http://www.bishopmuseum.org/ Cyrtodactylus epiroticus ðn ¼ 2Þ, C. klugei ðn ¼ 2Þ, C. loriae ðn ¼ 7Þ, C. novaeguineae ðn ¼ 3Þ, 1 This research was supported by National Science C. sermowaiensis ðn ¼ 30Þ, Gehyra mutilata Foundation grant DEB 0103794. This is contribution ðn ¼ 22Þ, G. oceanica ðn ¼ 27Þ, Gekko vittatus 2009-004 from the Pacific Biological Survey at the ðn ¼ 41Þ, Hemidactylus frenatus ðn ¼ 29Þ, and Bishop Museum. Manuscript accepted 19 April 2009. Lepidodactylus lugubris ðn ¼ 40Þ. Immediately 2 Department of Biology, Whittier College, Whittier, California 90608. after capture geckos were killed by an over- 3 Department of Biology, Pennsylvania State Uni- dose of sodium pentobarbital, fixed in neu- versity, Shenango Campus, Sharon, Pennsylvania 16146 tral buffered 10% formalin, stored in 70% (e-mail: [email protected]). ethanol, and accessioned in the herpetology 4 Bishop Museum, Department of Natural Sciences, collection of the Bishop Museum (bpbm), 1525 Bernice Street, Honolulu, Hawaii 96817 (e-mail: [email protected]). Honolulu, Hawai‘i. Subsequently, the body 5 Corresponding author (e-mail: sgoldberg@whittier cavity was opened by a longitudinal abdomi- .edu). nal incision and the gastrointestinal tract was removed by cutting across the esophagus and the rectum, and then shipped in a vial of alco- Pacific Science (2010), vol. 64, no. 1:131–139 doi: 10.2984/64.1.131 hol to Whittier College, Whittier, California, : 2010 by University of Hawai‘i Press where a detailed dissection of each tract was All rights reserved carried out using a dissecting microscope.

131 132 PACIFIC SCIENCE . January 2010

Helminths from individual hosts were re- than eight species (mean number of helminth moved to vials of 70% ethanol and later species per host species was 4.1 G 2.2; range placed under a coverslip in a drop of lacto- 1–8), and no individual host harbored more phenol and allowed to clear. Nematodes were than four helminth species (mean number identified from these preparations. Digeneans of helminth species per infected was and cestodes were washed in water, regres- 1.5 G 0.8 SD; range 1–4). Of the 69 infected sively stained in hematoxylin, and mounted geckos, 46 (67%) harbored one helminth spe- in balsam for identification under a com- cies, 15 (22%) harbored two species, 6 (9%) pound microscope. Although the helminths harbored three species, and two (3%) har- examined in this study were originally fixed bored four species. Number of individuals in situ with formalin, each species exhibits for each species and prevalence (number of unique characters that easily allow proper infected hosts/number of hosts examined identification; no previously undescribed hel- 100) as well as 29 new host and six new local- minth species was found. Voucher specimens ity (¼ country) records are recorded in Table were deposited in the United States Na- 1. usnpc tional Parasite Collection ( ), Beltsville, discussion Maryland, and the Bishop Museum (bpbm), Honolulu, Hawai‘i (Appendix 2). Parasite ter- The digenean Allopharynx macallisteri was minology is in accordance with Bush et al. originally described from Lepidodactylus lugub- (1997). Chi-square analysis, Morisita’s index, ris collected on Guam, Mariana Islands (Dai- and percentage similarity were used to com- ley et al. 1998). It has also been reported from pare infections between endemic versus L. lugubris collected on Rota, Mariana Is- nonendemic geckos (Brower et al. 1998, Ma- lands (Goldberg et al. 1998), and Cyrtodactylus gurran 2004). triparticus (then C. louisiadensis) from Papua New Guinea (Bursey et al. 2005a). results The cestode Oochoristica javaensis was orig- inally described from specimens taken from A total of 635 helminths was collected from the small intestines of the geckos Hemidac- 69 (34%) of 205 geckos. Of these, 413 (65%) tylus platyurus (then Cosymbotus platyurus), were mature individuals representing one Gehyra mutilata, and Hemidactylus frenatus species of Digenea (Allopharynx macallisteri), collected in Java (Kennedy et al. 1982) and one species of Cestoda (Oochoristica javaensis), has also been reported from Gehyra oceanica and 15 species of Nematoda (Aplectana macin- and Lepidodactylus paurolepis in the Pacific toshi, Maxvachonia adamsoni, Cosmocerca zugi, realm (see Goldberg and Bursey [2002] for Maxvachonia chabaudi, Meteterakis crombiei, locality records) as well as from Sphenomor- Moaciria chondropythonis, Moaciria moraveci, phus jobiensis from Papua New Guinea (Bur- Oswaldocruzia bakeri, Parapharyngodon maples- sey et al. 2005b). toni, Pharyngodon novaeguineae, Pharyngodon Six (33%) of the 18 nematodes found in oceanicus, Pseudorictularia dispsarilis, Skrjabine- this study are currently known only from Pa- lazia machidai, Spauligodon gehyrae, Spauligo- pua New Guinea (i.e., Cosmocerca zugi, Maxva- don hemidactylus). Pseudorictularia dispsarilis chonia adamsoni, Meteterakis crombiei, Moaciria occurred in the stomach; Allopharynx macallis- chondropythonis, M. moraveci, and Pharyngodon teri, Oochoristica javaensis, and Skrjabinelazia novaeguineae). Host lists for these species are machidai were found in the small intestine. given in Table 2. One of the 18 species, Pseu- All other helminths were found in the large dorictularia disparilis, is currently known only intestines. In total 222 (35%) were immature from the Australo-Papuan Region. It was forms (Abbreviata sp., Ascarops sp., and Acuari- originally described as Rictularia disparilis idae gen. sp.) represented by larvae in cysts in from specimens taken from a skink, Pseude- the stomach wall. moia entrecasteauxi (formerly Lygosoma entre- Although 20 species of helminths occurred casteauxi), collected in Australia by Irwin- in the sample, no host species harbored more Smith (1922) but reassigned to its current TABLE 1 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 Hemidactylus frenatus Lepidodactylus lugubris Cyrtodactylus epiroticus Cyrtodactylus klugei Cyrtodactylus loriae Cyrtodactylus sermowaiensis Cyrtodactylus novaeguineae Gehyra mutilata Gehyra oceanica Gekko vittatus

Helminth Taxa N % N % N % N % N % N % N % N % N % N %

Digenea Allopharynx macallisteri Dailey, Goldberg & — — —— ———*82—112 Bursey, 1998 Cestoda Oochoristica javaensis Kennedy, Killick & — — — — — — — *7 7 26 14 — Beverley-Burton, 1982 Nematoda Aplectana macintoshi (Stewart, 1914) — *16 100 — *91 55 — — — — — — Cosmocerca zugi Goldberg, Bursey & — — —*13—————— Kraus, 2005 Maxvachonia adamsoni Moravec & Sey, —*150———————*12 1990 yMaxvachonia chabaudi Mawson, 1972 — — — *6 10 — — — 4 7 — — Meteterakis crombiei Bursey, Goldberg & — *3 100 — *28 10 — — — — — — Kraus, 2005 Moaciria chondropythonis Gibbons, 1979 — — *1 14 — — — — — — — Moaciria moraveci Goldberg, Bursey & — — —*26—————— Kraus, 2007 Oswaldocruzia bakeri Moravec & Sey, — — —— ——*14——— 1986 Parapharyngodon maplestoni Chatterji, 1933 — *2 50 — *9 13 — — — *2 2 — — Pharyngodon novaeguineae Bursey, Goldberg & *125 50 — — — *26 33 — — — — — Kraus, 2008 yPharyngodon oceanicus Bursey & Goldberg, — — —— ——34——— 1999 134 PACIFIC SCIENCE . January 2010

TABLE 1 (continued) Hemidactylus frenatus Lepidodactylus lugubris Cyrtodactylus epiroticus Cyrtodactylus klugei Cyrtodactylus loriae Cyrtodactylus sermowaiensis Cyrtodactylus novaeguineae Gehyra mutilata Gehyra oceanica Gekko vittatus

Helminth Taxa N % N % N % N % N % N % N % N % N % N %

Pseudorictularia disparilis (Irwin-Smith, 1922) *250——— ———— —— ySkrjabinelazia machidai Hasegawa, 1984 — — — — — 4 18 — — — — ySpauligodon gehyrae Bursey & Goldberg, — — —— ——54——— 1996 ySpauligodon hemidactylus Bursey & Goldberg, — — — — —15*14—2934— 1996 Abbreviata sp. *4 100 *13 100 — *60 26 *4 33 — *3 11 *25 10 *2 3 — yAscarops sp. — — — *93 6 — — — — — — Acuariid gen. sp. — — — — — *2 5 *3 4 — *12 3 — status by Dollfus and Desportes (1945). The Eight (40%) of the 20 helminth species host list for P. dipsarilis includes the frogs found in this study (i.e., Allopharynx macallis- Litoria inermis, L. nigrofrenatus, Rana daemeli, teri, Oochoristica javaensis, Meteterakis crombiei, and R. supragrisea; the lizards Emoia caeruleo- Pseudorictularia disparilis, Skrjabinelazia machi- cauda, E. kordoana, E. physicae, Pogona mitchelli, dai, Abbreviata sp., Ascarops sp., and acuariid P. microlepidota, and Tiliqua occipitalis; and the larvae) require an intermediate host (Macy mammal Dasyurus hallucatus (Owen and 1965, Conn 1985, Anderson 2000). These Moorhouse 1980, Jones 1986, Bursey et al. species were represented by 277 (44% of to- 2008a, Goldberg et al. 2008). Eight of the 18 tal) individuals and were most likely acquired species of nematodes as well as the three gen- by ingestion of an insect intermediate host. era represented by larvae in cysts have much It should be noted that 222 (80%) of the 277 broader distribution patterns. Hosts lists for individuals requiring an intermediate host Aplectana macintoshi, Oswaldocruzia bakeri, and were larvae in cysts; the geckos most likely Abbreviata sp. (larvae) are given in Bursey serve as paratenic hosts. The remaining 14 et al. (2008a); for Maxvachonia chabaudi, Para- species, represented by 358 (56% of total) pharyngodon maplestoni, Spauligodon gehyrae, individuals, are thought to have direct life Skrjabinelazia machidai, and Ascarops sp. in cycles (Anderson 2000). The data presented Goldberg and Bursey (2002); for Pharyngodon here indicate that helminth parasites (specifi- oceanicus and Spauligodon hemidactylus in Gold- cally those capable of reaching maturity) of berg et al. (2005); and for Acuariidae gen. sp. Papua New Guinea geckos are largely gener- in Goldberg et al. (2008). Additions to these alists, the majority of which (357 individuals host lists are given in Table 1. versus 55) have direct life cycles. TABLE 2 Host Lists for Six Helminth Species Known Only from Papua New Guinea

Nematode Type Host Additional Hosts Reference

Cosmocerca zugi Gecko, Cyrtodactylus louisiadensis Bursey et al. (2005a) Gecko, Cyrtodactylus sermowaiensis This paper Skink, Sphenomorphus jobiensis Bursey et al. (2005b) Maxvachonia adamsoni Hylid frog, Litoria infrafrenata Moravec and Sey (1990) Gecko, Cyrtodactylus klugei This paper Gecko, Cyrtodactylus louisiadensis Bursey et al. (2005a) Skink, Emoia jakati Goldberg et al. (2008) Skink, Emoia longicauda Goldberg et al. (2008) Skink, Emoia obscura Goldberg et al. (2008) Skink, Emoia pallidiceps Goldberg et al. (2008) Gecko, Lepidodactylus lugubris This paper Skink, Sphenomorphus jobiensis Bursey et al. (2005b) Meteterakis crombiei Skink, Sphenomorphus jobiensis Bursey et al. (2005b) Ranid frog, Platymantis boulengeri Goldberg et al. (2009) Ranid frog, Platymantis schmidti Goldberg et al. (2009) Gecko, Cyrtodactylus klugei This paper Gecko, Cyrtodactylus sermowaiensis This paper Skink, Emoia battersbyi Goldberg et al. (2008) Skink, Emoia obscura Goldberg et al. (2008) Skink, Emoia pallidiceps Goldberg et al. (2008) Skink, Emoia physicae Goldberg et al. (2008) Skink, Emoia popei Goldberg et al. (2008) Skink, Sphenomorphus simus Bursey et al. (2007a) Moaciria chondropythonis Python, Morelia viridis Jones (1983) Gecko, Cyrtodactylus loriae This paper Moaciria moraveci Microhylid frog, Hylophorbus cf. rufescens Bursey et al. (2007b) Gecko, Cyrtodactylus sermowaiensis This paper Pharyngodon novaeguineae Gecko, Cyrtodactylus louisiadensis Bursey et al. (2008b) Gecko, Cyrtodactylus epiroticus This paper Gecko, Cyrtodactylus novaeguineae This paper 136 PACIFIC SCIENCE . January 2010

Five species of geckos examined in this only from Papua New Guinea and 300 hel- study (Cyrtodactylus epiroticus, C. klugei, C. minth individuals of species with wider distri- loriae, C. novaeguineae, and C. sermowaiensis), bution. Similarity indices, Morisita’s index currently known only from New Guinea and and percentage similarity, were calculated for the immediately adjacent islands, are repre- the helminth communities harbored by en- sented by 45 individuals and account for 22% demic geckos (community A) and by the of the host sample. The other five species more widely ranging geckos (community (Gehyra mutilata, G. oceanica, Gekko vittatus, B): Morisita’s index ¼ 0.184; percentage Hemidactylus frenatus, and Lepidodactylus lu- similarity ¼ 19.3%. Morisita’s index of com- gubris) are widely distributed in Oceania munity similarity is based upon Simpson’s (Bauer and Henle 1994). Six species of hel- index of dominance (DA and DB) and is de- minths found in this study (Cosmocerca zugi, fined as ð2SXiYiÞ/ðDA þ DBÞðSXiSYiÞ, where Maxvachonia adamsoni, Meteterakis crombiei, Xi ¼ number of individuals in taxon i in com- Moaciria chondropythonis, M. moraveci, and munity A, Yi ¼ number of individuals in Pharyngodon novaeguineae), also known only taxon i in community B. Percentage similarity from Papua New Guinea, are represented by ¼ S(lower of the two percentages for each 188 individuals and account for 30% of the taxon). Each index ranges from 0 (no simi- helminth sample. The endemic gecko species larity) to 1 (identical) (Brower et al. 1998). as a group harbored 12 helminth species (Ta- Thus, there is little similarity in the composi- ble 1). The more widely distributed gecko tion of the helminth community in these two species as a group also harbored 12 helminth groups of geckos. Questions arise: Is hel- species; however, only four helminth species minth community change driven by the hel- were common to both groups (Sorensen sim- minths of immigrant hosts; are immigrant ilarity index ¼ 0.33). The endemic geckos as hosts less susceptible to native helminths; are a group harbored 485 helminth individuals; native hosts more quickly infected by immi- 30 (67%) of 45 were infected, a mean hel- grant helminths? It is well known that hosts minth load of 16.9 per infected gecko. The can escape much of their native parasite com- wide-ranging geckos as a group harbored munity via human-assisted transport (Mitch- 150 helminth individuals; 40 (25%) of 160 ell and Power 2003, Torchin et al. 2003, were infected, a mean helminth load of 3.8 Torchin and Mitchell 2004), and Gehyra mu- per infected gecko. The helminth loads of tilata, G. oceanica, Hemidactylus frenatus, and endemic versus nonendemic geckos are signi- Lepidodactylus lugubris have all been widely ficantly higher (w2 ¼ 35:98, df ¼ 1, P <:001). moved via human activities (Kraus 2009). It Twenty taxa of helminths were found in is unknown how recently these species ar- this study. Of the helminth species known rived in the Papuan Region, but their wide only from Papua New Guinea, only one spe- and actively expanding distributions else- cies, Maxvachonia adamsoni, was found in the where suggest that it may have been relatively group of geckos of wider distribution. Thus, recently. Goldberg and Bursey (2000b) have the wide-ranging group of geckos harbored also shown that lizards such as Anolis sagrei 12 helminth species but just one individual are capable of transporting helminth species of a helminth species known only from Papua from their point of origin (Caribbean) to New Guinea and 149 helminth individuals of areas they colonize (Hawai‘i); but corre- wider distribution. Six of the 14 helminth sponding information on infections in immi- species of wider distribution (Aplectana macin- grant lizards is not yet available to make toshi, Maxvachonia chabaudi, Parapharyngodon conclusions about rates of helminth commu- maplestoni, Pseudorictularia disparilis, Abbre- nity change brought about by invading hosts. viata sp., and Ascarops sp.) were found in the Nevertheless, our data are consistent with a endemic gecko group. Thus, the endemic hypothesis of infection of endemic species gecko group harbored 12 helminth species, by helminths transported by recently immi- 185 helminth individuals of species known grating hosts. Gecko Helminths from Papua New Guinea . Goldberg et al. 137

acknowledgments konidae) from Papua New Guinea. J. Para- sitol. 91:882–889. We thank E. Teodoro, T. Doleck, and S. ———. 2005b. Endoparasites in Spheno- Goldsberry for assistance with dissections. In- morphus jobiensis (Sauria: Scincidae) from numerable people assisted with the fieldwork, Papua New Guinea with description of and we thank the inhabitants and landowners three new species. J. Parasitol. 91:1385– of the areas listed in Appendix 1. Particular 1394. thanks go to Jim Anamiato, Sedeka Andrew, ———. 2007a. A new species of Falcaustra Ilaiah Bigilale, Lydia Doni, Noel Doni, Fred (Nematoda, Kathlaniidae) and other nem- Francisco, Telen Garra, Charlie Graham, Bu- atodes from Sphenomorphus simus (Squa- lisa Iova, Conrad Kembwa, Dyson Libai, mata, Scincidae) from Papua New Guinea. Fred Malesa, David Mitchell, Florence Pais- Acta Parasitol. 52:142–145. parea, Pesto Roberts, Bendo Salepuna, John ———. 2007b. A new species of Moaciria Sengo, Stanley Simalken, John Slapeinsky, (Nematoda, Heterakidae) and other hel- Ben Yawi, and Isodore Yidika for providing minths in the red Mawatta frog, Hylo- logistical assistance. We thank the Papua phorbus cf. rufescens (Anura, Microhylidae) New Guinea National Museum and Art Gal- from Papua New Guinea. Acta Parasitol. lery for providing in-country collaborative as- 52:233–237. sistance and the Department of Environment ———. 2008a. A new species of Proteoce- and Conservation, National Research Insti- phalus (Cestoda: Proteocephalidae), de- tute, and Central, East New Britain, Milne scription of the male of Desmogathiema Bay, Morobe, and West Sepik provincial gov- papuensis (Nematoda; Quimperiidae), and ernments for permission to conduct this re- other endoparasites in Sylvirana supragri- search. sea (Anura: Ranidae) from Papua New Guinea. Comp. Parasitol. 75:33–48. ——— Literature Cited . 2008b. A new species of Pharyngodon (Nematoda, Pharyngodonidae) and other Anderson, R. C. 2000. Nematode parasites of helminths in Cyrtodactylus louisiadensis vertebrates: Their development and trans- (Sauria, Gekkonidae) from Papua New mission. 2nd ed. CABI Publishing, Wall- Guinea. Acta Parasitol. 53:41–45. ingford, United Kingdom. Bush, A. O., K. D. Lafferty, J. M. Lotz, and Bauer, A. M., and K. Henle. 1994. Familia A. W. Shostak. 1997. Parasitology meets Gekkonidae (Reptilia, Sauria). Part I. Aus- ecology on its own terms: Margolis et al. tralia and Oceania. Das Tierreich. Walter revisited. J. Parasitol. 83:575–583. de Gruyter, Berlin 109:1–306. Conn, D. B. 1985. Life cycle and postem- Brower, J. E., J. H. Zar, and C. N. von Ende. bryonic development of Oochoristica anolis 1998. Field and laboratory methods for (Cyclophyllidea: Linstowiidae). J. Parasi- general ecology. 4th ed. McGraw-Hill, tol. 71:10–16. Boston, Massachusetts. Dailey, M. D., S. R. Goldberg, and C. R. Bursey, C. R., and S. R. Goldberg. 1996. Bursey. 1998. Allopharynx macallisteri sp. Spauligodon gehyrae n. sp. (Nematoda: n. (Trematoda: Plagiorchiidae) from the Pharyngodonidae) from Gehyra oceanica mourning gecko, Lepidodactylus lugubris, (Sauria: Gekkonidae) from Guam, Mariana from Guam, Mariana Islands, Micronesia, Islands, Micronesia. J. Parasitol. 82:962– with a key to the species of the genus Allo- 964. pharynx. J. Helminthol. Soc. Wash. 65:16– Bursey, C. R., S. R. Goldberg, and F. Kraus. 20. 2005a. New genus, new species of Cestoda Dollfus, R. P., and C. Desportes. 1945. Sur (Anoplocephalidae), new species of Nema- le genre Rictularia Froelich 1802 (Nema- toda (Cosmocercidae) and other helminths todes, Spiruroidea). Ann. Parasitol. Hum. in Cyrtodactylus louisiadensis (Sauria: Gek- Comp. 20:6–34. 138 PACIFIC SCIENCE . January 2010

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Parasites, pathogens, and invasions by Woodlark Island (bpbm 17715–17722); Morobe plants and . Front. Ecol. Environ. Province: Lae (bpbm 18662), NW slope Mt. Shungol, bpbm 2:183–190. Morobe ( 18663). Lepidodactylus lugubris (Dume´ril 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, Appendix 1 19795–19798), Sudest Island (bpbm 19799, 19780), Gecko Specimens from Papua New Guinea Examined Rossel Island (bpbm 19801, 19802, 19804, 19805, from the Herpetology Collection of the Bishop Museum 19807, 19808, 19810–19812, 19814–19820), Nimowa (bpbm), Honolulu, Hawai‘i Island (bpbm 19822), Woodlark Island (bpbm 17727– 17731); Morobe Province: Mt. Shungol, Apele (bpbm ð ¼ Þ Cyrtodactylus epiroticus Kraus, 2008 n 2 . Morobe 18665–18668). Province: Mt. Shungol (bpbm 18653, 18654). Cyrtodactylus klugei Kraus, 2008 ðn ¼ 2Þ. Milne Bay Prov- ince: Sudest Island (bpbm 19739, 19740). Appendix 2 Cyrtodactylus loriae (Boulenger, 1897) ðn ¼ 7Þ. Milne Bay usnpc bpbm United States National Parasite Collection ( ) and Province: Siyomu Village, ( 17255), Bunisi Vil- Bernice P. Bishop Museum (bpbm) Accession Numbers lage (bpbm 17256, 17257); Central Province: Fane bpbm for Helminths from Gecko Specimens Collected in ( 18649, 18650); Morobe Province: NW slope Papua New Guinea Examined from the Herpetology Mt. Shungol (bpbm 18651, 18652). Collection of the Bishop Museum Cyrtodactylus novaeguineae (Schlegel, 1837) ðn ¼ 3Þ. West Sepik Province: Torricelli Mts., 2.9 to 3.2 km SSE Cyrtodactylus epiroticus: Pharyngodon novaeguineae (usnpc Mt. Sapau summit (bpbm 23314–23316). 101460), Pseudorictularia disparilis (usnpc 101461), Ab- Cyrtodactylus sermowaiensis (De Rooij, 1915) ðn ¼ 30Þ. breviata sp. (usnpc 101462). West Sepik Province: Parkop (bpbm 23317–23320), Cyrtodactylus klugei: Aplectana macintoshi (usnpc 101463), 3.2 km SSE Mt. Sapau summit, Torricelli Mts. (bpbm Maxvachonia adamsoni (usnpc 101464), Meteterakis 23321–23337, 23339–23347). crombiei (usnpc 101465), Parapharyngodon maplestoni Gehyra mutilata (Wiegmann, 1834) ðn ¼ 22Þ. Milne Bay (usnpc 101466), Abbreviata sp. (usnpc 101467). Province: Pini Range, Duabo (bpbm 15435–15438), Cyrtodactylus loriae: Moaciria chondropythonis (usnpc Wagabu Island (bpbm 15831), Normanby Island 101468). (bpbm 15832), Woodlark Island, Guasopa (bpbm Cyrtodactylus novaeguineae: Parapharyngodon novaeguineae 17708–17711), Sudest Island, Araeda (bpbm 19745– (lost in processing). 19751), Rossel Island, Damunu (bpbm 19753, 19754), Cyrtodactylus sermowaiensis: Aplectata macintoshi (usnpc Rossel Island, Cheme (bpbm 19755), Nimowa Island 101469; bpbm H391), Cosmocerca zugi (usnpc 101470), (bpbm 19756, 19758). Maxvachonia chabaudi (usnpc 101471), Meteterakis Gehyra oceanica (Lesson, 1830) ðn ¼ 27Þ. East New Brit- crombiei (usnpc 101472), Moaciria moraveci (usnpc ain Province: Marmar (bpbm 21995), Ulege (bpbm 101473), Parapharyngodon maplestoni (usnpc 101474), 21996); Milne Bay Province: Wagabu Island (bpbm Abbreviata sp. (usnpc 101475; bpbm H392), Ascarops 15833), Fergusson Island, Basima (bpbm 15834), Mi- sp. (usnpc 101476; bpbm H393). sima Island (bpbm 17218–17222), Woodlark Island, Gehyra mutilata: Skrjabinelazia machidai (usnpc 101477; Guasopa (bpbm 17712–17714), Sudest Island (bpbm bpbm H394), Spauligodon hemidactylus (usnpc 19759–19763), Rossel Island, Damunu (bpbm 19764– 101478), acuariid larvae (usnpc 101479). 19769), Nimowa Island (bpbm 19970), Normanby Gehyra oceanica: Oswaldocruzia bakeri (usnpc 101480), Island (bpbm 19771); West Sepik Province: Parkop Pharyngodon oceanicus (usnpc 101481), Spauligodon ge- (bpbm 23348), Sibilanga (bpbm 23349). hyrae (usnpc 101482), Spauligodon hemidactylus (usnpc Gekko vittatus (Houttuyn, 1782) ðn ¼ 41Þ. East New Brit- 101483), Abbreviata sp. (usnpc 101484), acuariid lar- ain Province: Mamar (bpbm 21997–22001); Milne vae (usnpc 101485). Bay Province: Misima Island (bpbm 16851–16853), Gekko vittatus: Allopharynx macallisteri (usnpc 101486; Woodlark Island, Guasopa (bpbm 17700–17706), bpbm F320), Oochoristica javaensis (usnpc 101487), Sudest Island, Araeda (bpbm 19773–19777), Rossel Maxvachonia chabaudi (usnpc 101488; bpbm H395), Island (bpbm 19778–19784, 19786, 19787); Morobe Parapharyngodon maplestoni (usnpc 101489), Abbre- Province: Mt. Shungol, Apele (bpbm 18656–18661); viata sp. larva (usnpc 101490; bpbm H396). West Sepik Province: Parkop (bpbm 23350–23355). Hemidactylus frenatus: Oochoristica javaensis (usnpc 101491; Hemidactylus frenatus Schlegel, 1836 ðn ¼ 29Þ. Central bpbm F321), Spauligodon hemidactylus (usnpc 101492), Province: Moroka (bpbm 22006); East New Brit- Abbreviata sp. (usnpc 101493), acuariid larvae (usnpc ain Province: Mamar (bpbm 22002–22005); Milne 101494). Bay Province: Duabo, Pini Range (bpbm 15441), Lepidodactylus lugubris: Allopharynx macallisteri (usnpc Misima Island (bpbm 16830–16833, 19788, 19789), 101495; bpbm F322), Maxvachonia adamsoni (usnpc Nimowa Island (bpbm 19790–19793), Guasopa, 101496).