Are Geckos Paratenic Hosts for Caribbean Island Acanthocephalans?

Are Geckos Paratenic Hosts for Caribbean Island Acanthocephalans? Evidence from Gonatodes antillensis and a Global Review of Squamate Reptiles Acting as Transport Hosts Alex Dornburg 1, April D. Lamb 1,2 , Dan Warren 3, Gregory J. Watkins-Colwell 4, Gregory A. Lewbart 5, and James Flowers 5

1 Corresponding author: North Carolina Museum of Natural Sciences, Raleigh, NC 27601 USA —email: [email protected] 2 Department of Applied Ecology, North Carolina State University, Raleigh, NC 27695 USA 3 Senckenberg Biodiversity and Climate Research Institute, Frankfurt am Main 60325 Germany 4 Division of Vertebrate Zoology, Peabody Museum of Natural History, Yale University, New Haven, CT 06520 USA 5 College of Veterinary Medicine, North Carolina State University, Raleigh, NC 27607 USA

ABSTRACT It is well known that reptiles can act as paratenic hosts for parasites that use mammals as their definitive hosts. However, studies of potential paratenic hosts in the Caribbean have been tempo - rally restricted to only diurnal species of lizards, thereby neglecting a dominant component of the nocturnal reptilian community: geckos. Many gecko species are human commensals with activ - ity periods that overlap temporally with those of domestic cats, making them prime candidates as potential transport hosts for cat parasites. However, no studies have reported geckos as paratenic hosts for felid parasites on any Caribbean island. Here we report the first records of subcutaneous oligacanthorhynchid cystacanths on the Venezuelan Coastal Clawed Gecko ( Gonatodes antillen - sis ) based on specimens collected in Curaçao and Bonaire. The cysts were identified as belonging to the genus Oncicola , likely those of Oncicola venezuelensis. This study reports these geckos as a new host record for oligacanthorhynchid cystacanths, as well as Curaçao and Bonaire as new geographic locales for these acanthocephalan parasites. We additionally provide a review of saurian cystacanths, comparing the restricted taxonomic focus of transport hosts in Caribbean is - lands to the distribution of paratenic squamate hosts both in the Neotropics and globally. We find evidence that the ability of squamate reptiles to act as transport hosts is a pervasive feature across their Tree of Life, suggesting that these animals may serve as important vectors for transporting parasites between intermediate and definitive hosts.

KEYWORDS Acanthocephala, intestinal helminths, cats, parasite, trophic transmission, mammals, birds, host–parasite interactions, vector species

Introduction associating these species and the parasites they are transporting are uncommon across the Caribbean. The transport of parasites by non-definitive trans - This lack of data creates a knowledge gap that chal - port, or paratenic, hosts creates an opportunity for lenges our ability to accurately forecast the spread parasitic organisms to utilize and potentially estab - of internal parasites to both domestic animals and lish novel transmission pathways (Marcogliese wildlife. In particular, despite the high potential of 2007; Parker et al. 2015; Cable et al. 2017). reptiles acting as paratenic hosts to the acantho - Although it is well established that nonmam - cephalan parasites of cats and dogs, these poten - malian vertebrates often act as paratenic hosts for tial transmission vectors have received little a variety of mammalian parasites (Bolt et al. 1993; attention. To date, studies of lizards as paratenic Anderson 2000; Strube et al. 2013), baseline data hosts in the Caribbean have been restricted to

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TABLE 1. Institution and catalog numbers of Gonatodes antillensis specimens examined in this study. Abbrevia - tions: CAS, California Academy of Sciences; LACM, Natural History Museum of Los Angeles County; MCZ, Museum of Comparative Zoology; NCMNS, North Carolina Museum of Natural Sciences; RMNH, Naturalis Bio - diversity Center (former Rijksmuseum voor Natuurlijke Historie); UMMZ, University of Michigan Museum of Zoology; USNM, National Museum of Natural History, Smithsonian Institution; YPM HERR, Division of Ver - tebrate Zoology Herpetology Reptile Collection, Peabody Museum of Natural History, Yale University.

Institution code Specimen numbers

CAS 113319–113345 LACM 126062–126078 MCZ R-141593, R-141600, R-149268, R-149349, R-149351, R-149367, R-149369–R-149371, R-27548, R-60579, R-69577, R-69580, R-69581, R-69583, R-69584, R-69586, R-69587, R-69592, R-69596, R-69597, R-81514, R-82968, R-82969, R-82971, R-82973, R-82974, R-82976, R-82978, R-82979 NCMNS 89384–89387, 89390, 89392–89412, 89419–89443, 89445–89448, 89452, 89491 RMNH 13421, 14742A–B, 16682A–B, 18000, 18001A–C, 18002, 18004, 18008 UMMZ 57317–57345, 127795, 127796, 142675, 151501, 179309–179314 USNM 79231, 94980 YPM YPM HERR 017579–017584, YPM HERR 018188–018199, YPM HERR 018438–018449, YPM HERR 018451–018460, YPM HERR 018603–018606, YPM HERR 018640–018644

diurnal species such as various species of Anolis or combination of field-collected and museum spec - Ameiva (Dobson et al. 1992; Goldberg et al. 1998a; imens. To place our findings into a broader con - Nickol et al. 2006). This temporal bias neglects an text, we provide a review of all known Caribbean important nocturnal component of Caribbean island squamates as well a global survey of squa - lizard communities: geckos. mates that have been found to act as paratenic With more than 1,600 species, geckos repre - hosts for acanthocephalans. sent one of the most successful radiations of lizards and are common commensal organisms in Methods urban and suburban landscapes worldwide (Gamble et al. 2012). Although several studies As part of an ongoing study of the ecology and have found geckos to harbor cystacanths of acan - evolutionary biology of geckos on the southern thocephalans in Southeast Asia (Saehoong and Caribbean island of Curaçao (Dornburg et al. Wongsawad 1997; Mahagedara and Rajakaruna 2011, 2016; Lamb et al. 2017), we collected a total 2015), Africa (Oluwafemi et al. 2017), Australia of 140 Gonatodes antillensis between 2011 and (Barton 2015), South America (Anjos et al. 2005), 2017 from 10 locales across Curaçao, which were or Central America (Goldberg and Bursey 2004a; deposited in the Peabody Museum of Natural Bursey et al. 2007), no studies have investigated History, Yale University, New Haven, Connecti - whether Caribbean island species act as paratenic cut, USA, and the North Carolina Museum of hosts. Given both the spatial and temporal over - Natural Sciences, Raleigh, North Carolina, USA lap of many Caribbean species with domestic and (Table 1). All specimens were visually inspected feral cats, geckos make likely paratenic host can - on collection, and those containing subcuta - didates. Ground-dwelling species may be partic - neous cysts were documented ( Figure 1). To ularly vulnerable to felid predation, making an determine the historic prevalence of similar sub - assessment of their parasite transmission potential cutaneous cysts, we were aided by collection staff of high utility for veterinary and wildlife medicine. who collectively visually inspected the specimen Herein we examine whether the ground-dwelling holdings of the Division of Vertebrate Zoology Venezuelan Coastal Clawed Gecko ( Gonatodes Herpetology Reptile Collection, Yale Peabody antillensis ; Lidth de Jeude 1887) acts as paratenic Museum of Natural History; the Museum of host for acanthocephalan parasites by using a Comparative Zoology, Harvard University,

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FIGURE 1. Oncicola cystacanths in Gonatodes antillensis . A. Individual G. antillensis with visible subcutaneous cystacanths highlighted. B. Dorsal view of G. antillensis. C. Slide of Oncicola cystacanth extracted from a cyst.

Cambridge, Massachusetts, USA; National Medicine at North Carolina State University, Museum of Natural History, Smithsonian Insti - Raleigh, North Carolina, USA, for identification. tution, Washington, DC, USA; the Natural His - Larval acanthocephalans (cystacanths) were tory Museum of Los Angeles County, Los extracted from cysts with the aid of a dissecting Angeles, California, USA; the Naturalis Biodi - microscope. Cystacanths were stained with versity Center (former Rijksmuseum voor Natu - Semichon’s carmine, dehydrated in an ethanol urlijke Historie), Leiden, Netherlands; the series, cleared in xylene and mounted in balsam California Academy of Sciences, San Francisco, for identification. Identifications were based on California, USA; and the University of Michigan taxonomic keys and descriptions (Schmidt 1972, Museum of Zoology, Ann Arbor, Michigan, 1985; Nickol and Dunagan 1989; Smales 1997; USA, for specimens with similar subcutaneous Nickol et al. 2006; Santos et al. 2017). Compara - cysts ( Table 1). Subcutaneous cysts found in any tive voucher specimens (HWML 48252, 48254, samples were excised and sent to the parasitol - 48256, 48257, 48258, 49710) were borrowed ogy laboratory at the College of Veterinary from the Harold W. Manter Laboratory for

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Parasitology, University of Nebraska State Museum, canths indicate inclusion in the family Oligacan - Lincoln, Nebraska, USA (HWML). thorhynchidae and the genus Oncicola . The long To place our results into context, we compiled length of the lemnisci (Nickol et al. 2006) and the a list of all known paratenic saurian hosts for oli - subcutaneous site of infection in Caribbean gacanthorhynchid and centrorhynchid cysta - saurian hosts are suggestive of Oncicola venezue - canths across all Caribbean islands. These two lensis (Marteau 1977). Unfortunately, the pro - groups represent two major radiations of acantho - boscis of our cystacanth specimens remained cephalans infecting terrestrial vertebrates that prey inverted, preventing a definitive species diagno - on squamates (mammals and birds; Petrochenko sis. All cystacanth specimens were deposited 1956), and would therefore be likely to co-opt in the Division of Invertebrate Zoology, Yale squamates as transport hosts. Searches were con - Peabody Museum of Natural History (YPM IZ ducted using Google Scholar and ISI Web of 102933–102938). Knowledge using the following key terms: lizard, Results of our literature and database survey cystacanth, Caribbean, transport host, paratenic, yielded 15 publications and 4 museum records saurian, gecko, Anolis , Iguanidae, snake, helminth, containing information on paratenic saurian hosts squamate, reptile, Acanthocephala, Oligantho - on Caribbean islands ( Table 2). Anolis was the hynchidae and Centrorhynchus . Additionally, most frequently represented genus, with 21 references within each found document were species reported as paratenic hosts. Of these, only checked for additional publications not captured three species were reported with oligacan - by our key terms. Although additional key search thorhynchid cystacanths, whereas all 21 Anolis terms are certainly possible, no additional terms species as well as a species of Ameiva and the we attempted yielded new reports. We further snake Leimadophis reginae were reported with searched the Harold W. Manter Laboratory of Par - centrorhynchid cystacanths ( Table 2). No subcu - asitology and the United States National Parasite taneous centrorhynchid infections were reported. Collection, National Museum of Natural History, In contrast, both our findings for Gonatodes antil - Smithsonian Institution, for records not reported lensis and prior work in Anolis cristatellus and in the primary literature. This search was then Anolis stratulus reported subcutaneous infections expanded to compare Caribbean records to all (Table 2). records of these parasites in squamate reptiles The results of our search of paratenic hosts globally. globally yielded 202 records that spanned 35 countries and 184 species (Appendix). The United Results States was the most represented country with 36 records, followed by Costa Rica with 33 (Appen - Across our survey of the 140 Venezuelan Coastal dix). We found records of 119 species possessing Clawed Geckos collected from Curaçao, only oligacanthorhynchid and 75 species possessing specimens (6 of 22) from one locale, Director’s Centrorhynchus infections (Appendix). Infections Bay, possessed subcutaneous cysts ( Figure 2). in the body cavity (coelom) were most common Cysts were primarily located below the anterior in both oligacanthorhynchid and centrorhynchid surface of the neck and laterally along the main infections, respectively representing 60% and body, with several specimens also exhibiting cysts 50% of the reported records. Ophidians were the near the posterior insertion of the humerus with most reported group of squamates (56%), fol - the scapula. Visual inspections of museum speci - lowed by iguanians (19%) and lacertoids (13%). mens for geckos that possessed subcutaneous Only a single record of a centrorhynchid cyst was cysts ( Table 1) yielded only one specimen from found for another Gonatodes , with Hemidacty - the University of Michigan Museum of Zoology, lus the most frequently represented (54%) gecko Ann Arbor, Michigan, USA (UMMZ 179309). genus (Appendix). That gecko had been collected in 1984 from Bonaire, Netherlands Antilles, and we found it to Discussion contain three previously undocumented subcuta - neous cysts. Morphological characteristics (i.e., Understanding the transmission pathways that size, shape, proboscis hooks) of extracted cysta - parasites can utilize to move between hosts at

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FIGURE 2. Frequency and spatial distribution of Oncicola cystacanths in Curaçao detected from our surveys and museum specimens. A. Frequency of cystacanths per site. B. Location of sites on the island. Blue indicates absence of cystacanths and yellow indicates presence of cystacanths. Abbreviation: CARMABI, Caribbean Research and Management of Biodiversity.

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different life stages is a fundamental component of both veterinary and human medicine. Our 2 2 study represents the first documentation of Onci - 9 9 9 9 1 1

cola in Gonatodes antillensis and the first record a a l l a a of a Caribbean island gecko acting as a paratenic c c a a P P host for acanthocephalans. While definitive hosts

d d n n of Oncicola species include felids and birds, Onci - e a a

c n n n cola venezuelensis is known only from felids, o o e s s r b b e specifically domestic cats and ocelots ( Leopardus f o o e D D R

4 pardalis ) (Marteau 1977; Patton et al. 1986; Fuller ; ; 9 2 2 9 9 9 7 7 and Nickol 2011; Santos et al. 2017). Rep (1975) 1 9 9 8 8

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L n n e likely been on the islands for at least four decades. y y o o b s s M d d d b b l n n Our finding of an additional Oncicola cystacanth W o o o u u H D B D G B from a specimen collected in 1984 from the island of Bonaire provides additional evidence that the Venezuelan Coastal Clawed Gecko has been act - ing as a paratenic host for nearly as long. How - ever, this raises the question of the role G. antillensis plays in the life cycle of Oncicola . Oncicola venezuelensis utilizes termites of the genus Nasutitermes as intermediate hosts, with s a d transmission facilitated through the encounter of n m o a a l g h s

a infected fecal matter (Fuller and Nickol 2011). Par - a I b B o

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i . m m n n r t a a transmission (Nickol et al. 2006). Unfortunately, T A J S A J the natural history of Gonatodes antillensis has n

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e e e e 2017), and resolution of the feeding ecology of G. d t t e o o o t n n i I C I B C antillensis is broadly classified as comprising small S invertebrates (Van Buurt 2005). Although diets of G. antillensis have not been studied explicitly, ter - e a i

n mites of the genus Nasutitermes are considered i n i g n z e e t

r native to Curaçao (Van Buurt and Debrot 2012) i r i

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e and species of Gonatodes for which diets are t l g p h a H a c a o w v s s known have all been documented to eat termites

d s s s s a i i i i l l l l

m (Quesnel 1957; Vitt et al. 2000; Miranda and o o o o i n n n n e L A A A A Andrade 2003). This suggests that the feeding ecol - ogy of G. antillensis may predispose this species as .

D a transmission vector for O. venezuelensis . Further E U d

i studies of the feeding ecology of this gecko are N I h T c

N needed to test this hypothesis. n h t O y n C h

Given that no native felid species occur in the r a 2 c o r E a

t Leeward Antilles, domestic cats likely represent t L s n B e y A the only definitive host of Oncicola venezuelensis C T C

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on these islands. This suggests a life cycle of O. ther suggesting that this mode of transmission venezuelensis beginning in a termite that has may be a pervasive squamate-wide phenomenon. encountered infected fecal matter. Infected ter - Of all squamate groups, snakes were the most fre - mites are then ingested by Gonatodes antillensis, quently represented. However, this should not be which acts as a paratenic host, and the parasite is taken to imply that snakes are particularly prone ultimately transmitted to domestic cats to restart to acting as paratenic hosts, as a survey based on the cycle. This intermediate termite to paratenic the combined pool of natural history notes and lizard to definitive cat host life cycle has been pro - museum records cannot account for sampling posed for Oncicola on other Caribbean islands biases (Dornburg et al. 2017). Further work is (Fuller and Nickol 2011), adding a veterinary needed to discern geographic, taxonomic and life health concern to the global conservation concern history-based biases in sampling efforts and com - of predation by feral cats driving reptile extirpa - plete a more robust understanding of patterns in tion (Gibbon et al. 2000; Medina et al. 2011). paratenic parasitism in squamates globally. How - Although cat predation on G. antillensis has not ever, comparing the restricted taxonomic focus of been reported, many studies have described feral transport hosts in Caribbean islands to the distri - cat predation on geckos (Bonnaud et al. 2011; Kutt bution of paratenic hosts both in the Neotropics 2011), including other species of Gonatodes and globally (Appendix) strongly suggests a sig - (Alonso et al. 2009). As such, the likelihood of at nificant knowledge gap in the region. least some felid predation events is certainly not To date, our knowledge of squamates acting negligible. Additionally, we observed a high fre - as paratenic hosts on Caribbean islands has been quency of feral cats at the Director’s Bay site where limited to records from Anolis and a single lacer - all cystacanth-containing G. antillensis specimens toid and ophidian ( Table 2). Our study expands were collected. Although cat frequencies and the reservoir of potential Caribbean transport habitat characteristics were not quantified, this hosts to include geckos of the genus Gonatodes . was by far the most ecologically degraded site of Cystacanths of Centrorhynchus have previously all sampling locations, with large amounts of trash been found in Gonatodes albogularis in Panama; and debris scattered throughout. Large quantities however, our record is the first instance of Onci - of garbage are known to attract cats by providing cola . Further, G. albogularis is diurnal whereas food resources, thereby increasing the risk of G. antillensis is nocturnal (Van Buurt 2005), pathogen infections (Plaza and Lambertucci thereby overlapping in diel activity with the 2017). The condition of this site could explain the other geckos found in our survey (Appendix). Of high frequency of infection as the harsh arid con - the other nocturnal geckos encountered in our ditions of the island might prevent aggregations survey, geckos of the genus Hemidactylus are of of feral cats in nonurbanized settings. Further, particular note as potential transmission vectors. Nasutitermes termites have repeatedly been found Hemidactylus geckos have invaded tropical to be among the most common termites in highly ecosystems worldwide and have been docu - disturbed or degraded habitats worldwide (de mented to prey on other geckos (Bolger and Case Souza and Brown 1994; Eggleton et al. 1995; Ban - 1992; Dornburg et al. 2011, 2016), including G. deira et al. 2003), suggesting that this site could antillensis in Curaçao (Dornburg et al. 2011), constitute a perfect storm for acanthocephalan making this an additional prime candidate for transmission potential. investigation. More broadly, our survey of paratenic squamate hosts undoubtedly underes - Paratenic Parasitism of Squamates timates the extent to which squamates act as and the Helminth Life Cycle paratenic hosts. The results of our literature and database survey The ability of acanthocephalan species to suggest squamates to represent an underappreci - infect nondefinitive hosts while remaining trans - ated component of the ecology and life cycle of mit table offers a solution to the ecological prob - acanthocephalans (Appendix). We found records lem of transitioning between different feeding from 184 species of squamates acting as potential guilds at different trophic levels between life transmission vectors in 35 countries (Appendix). stages. Many acanthocephalans such as Oncicola These species span the squamate Tree of Life, fur - begin their life cycle in arthropod intermediate

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hosts, yet require birds or mammals feeding at Acknowledgments higher trophic levels for their adult stage (Schmidt 1985). The results of our survey and instances of We thank Mark Vermeij and the staff of paratenic parasitism by other parasites with sim - CARMABI for facilitating collecting and permit - ilar complex life histories (e.g., Marcogleise 2002; ting, the Westpunt syndicate for help with field - Choisy et al. 2003; Cirtwill et al. 2017) reiterates a work, Bryan Stuart for help with specimen question: When do parasitic lineages evolve to co- curation and imaging of Gonatodes antillensis opt pathways of energy flow across community specimens and Gregory Schneider at the Univer - food webs to reach their target definitive hosts? It sity of Michigan, College of Literature, Science, should not be surprising that squamates are and the Arts, Museum of Zoology, for his aid with broadly used as transport hosts, as these often rep - specimen loans. We additionally thank Lauren resent direct links between lower trophic level Scheinberg at the California Academy of Sciences, consumers (e.g., arthropods) and higher level ver - Neftali Camacho at the Los Angeles County tebrate predators (Watkins-Colwell et al. 2006). Museum and Rayna Bell of the Smithsonian Insti - Given that we found documented instances of tution for checking specimens of G. antillensis for squamate paratenic parasitism that span the globe, external cysts. Also, Gabor Racz and Scott Gar - we suggest that investigating the ecology and evo - dener of the Harold W. Manter Laboratory of Par - lution of paratenic parasitism in squamates rep - asitology for the loan of Oncicola venezuelensis resents a potentially rich area of research in the specimens. evolution of parasite transmission pathways of high importance to both wildlife and human Received 16 May 2018; revised and accepted 26 medicine. September 2018.

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Literature Cited in the Western Diamondback Rattlesnake, Crotalus atrox (Serpentes: Viperidae). Journal of Parasitology 83:751–752. ACHOLONU , A.D. 1976. Helminth fauna of saurians from Puerto —1997b. Oligacanthorhynchid cystacanths (Acanthocephala) Rico with observations on the life cycle of Lueheia inscripta in a long-nosed snake, Rhinocheilus lecontei lecontei (Colu - (Westrumb, 1821) and description of Allopharynx puertori - bridae) and a Mojave rattlesnake, Crotalus scutulatus scutu - censis sp. n. Proceedings of the Helminthological Society of latus (Viperidae) from Maricopa County, Arizona. Washington 43:106–116. Southwestern Naturalist 42(2):232–236. ALONSO , M.L., J.M. C OTRINA , D.A. P ARDO , E. F ONT AND M. —1998a. Natural occurrence of Macracanthorhynchus ingens MOLINA -B ORJA . 2009. Sex differences in antipredator tail- (Acanthocephala: Oligacanthorhynchidae) in a northern waving displays of the diurnal yellow-headed gecko Gona - black racer, Coluber constrictor constrictor (Serpentes: Colu - todes albogularis from tropical forests of Colombia. Journal bridae). Texas Journal of Science 50:85–89. of Ethology 28:305–311. —1998b. Helminths of the Prairie Rattlesnake, Crotalus viridis ANDERSON , R.C. 2000. Nematode Parasites of Vertebrates: Their viridis (Serpentes: Viperidae), from western South Dakota. Development and Transmission. Wallingford, UK: CABI Journal of the Helminthological Society of Washington Publishing. 650 pp. 65:105–107. ANDRUSHKO , A.M. AND G.S. M ARKOV . 1956. The helminth fauna BOLGER , D.T. AND T.J. C ASE . 1992. Intra- and interspecific inter - of lizards from the deserts of Central Asia and its ecological ference behaviour among sexual and asexual geckos. Ani - character. Vestnik Leningradskogo Gosudarstvennogo Uni - mal Behaviour 44:21–30. versiteta, Seriya Biologii 21:61–71. BOLT , G., J. M ONRAD , F. F RANDSEN , P. H ENRIKSEN AND H.H. ANJOS , L.A., C. B EZERRA , D.C. P ASSOS , D. Z ANCHI AND C.A.B. DIETZ . 1993. The common frog ( Rana temporaria ) as a GALDINO . 2011. Helminth fauna of two gecko lizards, Hemi - potential paratenic and intermediate host for Angiostrongy - dactylus agrius and Lygodactylus klugei (Gekkonidade), lus vasorum . Parasitology Research 79:428–430. from Caatinga biome, northeastern Brazil. Neotropical BONNAUD , E., F.M. M EDINA , E. V IDAL , M. N OGALES , B. T ERSHY , Helminthology 5:285–290. E. Z AVALETA , C.J. D ONLAN , B. K EITT , M. L E CORRE AND S.V. ANJOS , L.A., C.F.D. R OCHA , D. V RCIBRADIC AND J.J. V ICENTE . HORWATH . 2011. The diet of feral cats on islands: A review 2005. Helminths of the exotic lizard Hemidactylus mabouia and a call for more studies. Biological Invasions 13:581–603. from a rock outcrop area in southeastern Brazil. Journal of BRITO , S.V., G. C ORSO , A.M. A LMEIDA ., F.S. F ERREIRA , W.O. Helminthology 79:307–313. ALMEIDA , L.A. A NJOS , D.O. M ESQUITA AND A. V ASCONCEL - ÁVILA , R.W. AND R.J. S ILVA . 2010. Checklist of helminths from LOS . 2014. Phylogeny and micro-habitats utilized by lizards lizards and amphisbaenians (Reptilia, Squamata) of South determine the composition of their endoparasites in the America. Journal of Venomous Animals and Toxins includ - semiarid Caatinga of Northeast Brazil. Parasitology ing Tropical Diseases 16:543–572. Research 113:3963–3972. BANDEIRA A.G., A. V ASCONCELLOS , M.P. S ILVA AND R. C ON - BROOKS , G.R. 1972. Intestinal parasites of the lizard Lygosoma STANTINO . 2003. Effects of habitat disturbance on the termite laterale . Quarterly Journal of the Florida Academy of Sci - fauna in a highland humid forest in the Caatinga domain, ences 35:8–14. Brazil. Sociobiology 42:117–128. BUNDY , D.A.P., P. V OGEL AND E.A. H ARRIS . 1987. Helminth par - BARTON , D.P. 2015. Helminth parasites of the introduced Asian asites of Jamaican anoles (Reptilian: Iguanidae): A compar - House Gecko (‘ Hemidactylus frenatus ’) (Gekkonidae), in the ison of the helminth fauna of 6 Anolis species. Journal of Northern Territory, Australia. Northern Territory Natural - Helminthology 61:77–83. ist 26:44–55. BURSEY , C.R. AND S.R. G OLDBERG . 2003. Acanthocephalus BARUS , V. AND F. T ENORA . 1976. New data on parasitic nema - saurius n. sp. (Acanthocephala: Echinorhynchidae) and todes and acanthocephalans recovered from Amphibia and other helminths from the lizard Norops limifrons (Sauria: Reptilia from Afghanistan. Acta Universitatis Agriculturae, Polychrotidae) from Costa Rica. Journal of Parasitology Brno, Facultas Agronomica 24:339–350. 89:573–576. BENES , E.S. 1985. Helminth parasitism in some central Arizona —2006. Helminths in Mesaspis monticola (Squamata: lizards. Southwestern Naturalist 27:467–473. Anguidae) from Costa Rica, with the description of a new BENNETT , A.F. AND G.C. G ORMAN . 1979. Population density species of Entomelas (Nematoda: Rhabdiasidae) and a new and energetics of lizards on a tropical island. Oecologia species of Skrjabinodon (Nematoda: Pharyngodonidae). 42:339–358. Parasite 13:183–191. BIRLIK , S., H.S. Y ILDIRIMHAN , Ç. I LGAZ AND Y. K UMLUTAş . —2007. A new species of Parapharyngodon (Nematoda: 2018. Helminth fauna of Spiny Tailed Lizard, Darevskia Pharyngodonidae) and other helminths in Typhlosaurus lin - rudis (Bedriaga, 1886) (Sauria: Lacertidae) from Turkey. eatus (Squamata: Scincidae), from southern Africa. Onder - Helminthologia 55:45–51. stepoort Journal of Veterinary Research 74:143–147. BIRLIK , S., H.S. Y ILDIRIMHAN , N. S üMER , Ç. I LGAZ , Ö. G üÇLü BURSEY , C.R., S.R. G OLDBERG AND C.L. M ILLER . 2004. Two new AND S.H. D URMUş . 2015. The helminth fauna of Apathya species of Falcaustra and comments on helminths of Norops cappadocica (Werner, 1902) (Anatolian Lizard) (Squa - tropidolepis (Sauria: Polychrotidae) from Costa Rica. Journal mata: Lacertidae) from Turkey. Helminthologia 52:310– of Parasitology 90:598–603. 315. BURSEY , C.R., S.R. G OLDBERG AND S.R. T ELFORD JR. 2005. Pla - BOLETTE , D.P. 1997a. First record of Pachysentis canicola (Acan - giorchis taiwanensis (Digenea: Plagiorchiidae), Kurilonema thocephala: Oligacanthorhynchida) and the occurrence of markovi (Nematoda: Rhabdiasidae) and other helminths in Mesocestoides sp. Tetrathyridia (Cestoidea: Cyclophyllidea) Eumeces latiscutatus (Scincidae) and Takydromus tachydro -

Downloaded From: https://bioone.org/journals/Bulletin-of-the-Peabody-Museum-of-Natural-History on 11 Jul 2019 Terms of Use: https://bioone.org/terms-of-use Access provided by University of Mississippi Geckos Paratenic Hosts for Caribbean Island Acanthocephalans • Dornburg et al. 75

moides (Lacertidae) from Japan. Comparative Parasitology DORNBURG , A., C. L IPPI , S. F EDERMAN , J.A. M OORE , D.L. W AR - 72:234–240. REN , T.L. I GLESIAS , M.C. B RANDLEY , G.J. W ATKINS -C OLWELL , —2007. Gastrointestinal helminths of 14 species of lizards from A.D. L AMB AND A. J ONES . 2016. Disentangling the influence Panama with descriptions of five new species. Comparative of urbanization and invasion on endemic geckos in tropical Parasitology 74:108–140. biodiversity hot spots: A case study of Phyllodactylus martini BURSEY , C.R., S.R. G OLDBERG , S.R. T ELFORD JR. AND L.J. V ITT . (Squamata: Phyllodactylidae) along an urban gradient in 2012. Metazoan endoparasites of 13 species of Central Curaçao. Bulletin of the Peabody Museum of Natural His - American anoles (Sauria: Polychrotidae: Anolis ) with a tory 57:147–164. review of the helminth communities of Caribbean, Mexi - DORNBURG , A., D.L. W ARREN , T. I GLESIAS AND M.C. B RANDLEY . can, North American, and South American anoles. Com - 2011. Natural history observations of the ichthyological and parative Parasitology 79:75–132. herpetological fauna on the island of Curaçao (Netherlands). BURSEY , C.R., S.R. G OLDBERG AND L.J. V ITT . 2006. New species Bulletin of the Peabody Museum of Natural History of Oswaldocruzia (Nematoda: Molineidae) in Ameiva festiva 52:181–186. (Squamata: Teiidae) from Nicaragua. Journal of Parasitol - DüSEN , S., Y. K UMLUTAş , Ç. I LGAZ , H. Y AKA AND F. K ARADAYI . ogy 92:350–352. 2013. Helminth parasites of the three racerunner lizards: CABLE , J., I. B ARBER , B. B OAG , A.R. E LLISON , E.R. M ORGAN , K. Eremias pleskei Nikolsky, 1905 (Pleske’s Racerunner–Tran - MURRAY , E.L. P ASCOE , S.M. S AIT , A.J. W ILSON AND M. B OOTH . scaucasian Racerunner), Eremias strauchi Kessler, 1878 2017. Global change, parasite transmission and disease con - (Strauch’s Racerunner) and Eremias suphani Basoglu & trol: Lessons from ecology. Philosophical Transactions of the Hellmich, 1968 (Suphan Racerunner) collected from eastern Royal Society B: Biological Sciences 372:20160088. part of Turkey. Helminthologia 50:108–111. CABRERA -G UZMÁN , E. AND L. G ARRIDO -O LVERA . 2014. EGGLETON , P., D.E. B IGNELL , W.A. S ANDS , B. W AITE , T.G. Helminth parasites of the lesser scaly anole, Anolis uniformis WOOD AND J.H. L AWTON . 1995. The species richness of ter - (Squamata: Dactyloidae), from Los Tuxtlas, Southern Mex - mites (Isoptera) under differing levels of forest disturbance ico: Evidence of diet and habitat use. South American Jour - in the Mbalmayo Forest Reserve, southern Cameroon. Jour - nal of Herpetology 9:183–189. nal of Tropical Ecology 11:85–98. CHOISY , M., S.P. B ROWN , K.D. L AFFERTY AND F. T HOMAS . 2003. ELKINS , C.A. AND B.B. N ICKOL . 1983. The epizootiology of Evolution of trophic transmission in parasites: Why add Macracanthorhynchus ingens in Louisiana. Journal of Para - intermediate hosts? American Naturalist 162:172–181. sitology 69:951–956. CIRTWILL A.R., C. L AGRUE , R. P OULIN AND D.B. S TOUFFER . EVERARD , C.O.R. 1975. Endoparasites of some amphibia, rep - 2017. Host taxonomy constrains the properties of trophic tiles and small mammals from Trinidad. Living World: Jour - transmission routes for parasites in lake food webs. Ecology nal of the Trinidad and Tobago Field Naturalists’ Club 98:2401–2412. 1975:72–79. COLLINS , R.F. 1968. The helminths of Natrix spp. and Agk - FARZALIEV , A.M. AND V.I. P ETROCHENKO . 1980. New data on istrodon piscivorus piscivorus (Reptilia: Ophidia) in eastern the life-cycle of the acanthocephalan Macracanthorhynchus North Carolina [master’s thesis]. Winston-Salem, NC: Wake catulinus Kostylew, 1927 (Acanthocephala), a parasite of car - Forest University. nivores. Trudy Vsesoyuznogo Instituta Gel’mintologii —1969. The helminths of Natrix spp. and Agkistrodon pis - 25:140–144. (in Russian) civorus piscivorus (Reptilia: Ophidia) in eastern North Car - FOSTER , G.W., P.E. M OLER , J.M. K INSELLA , S.P. T ERRELL AND D.J. olina. Journal of the Elisha Mitchell Scientific Society FORRESTER . 2000. Parasites of eastern indigo snakes ( Dry - 85:141–144. marchon corais couperi ) from Florida, USA. Comparative CRISCIONE , C.D. AND W.F. F ONT . 2001. The guest playing host: Parasitology 67:124–128. Colonization of the introduced Mediterranean gecko, Hemi - FULLER , C.A. AND B.B. N ICKOL . 2011. A description of mature dactylus turcicus , by helminth parasites in southeastern Oncicola venezuelensis (Acanthocephala: Oligacan - Louisiana. Journal of Parasitology 87:1273–1278. thorhynchidae) from a feral house cat in the U. S. Virgin DAS , E.N. 1950. On some juvenile forms of Acanthocephala of Islands. Journal of Parasitology 97:1099–1100. the genus Centrorhynchas from India. Indian Journal of FULLER , C.A., P. R OCK AND T. P HILIPS . 2003. Behavior, color Helminthology 2:49–56. changes, and predation risk induced by acanthocephalan DAVIS , E., J.C. B EANE AND J.R. F LOWERS . 2016. Helminth para - parasitism in the Caribbean termite Nasutitermes acajutlae . sites of pit vipers from North Carolina. Southeastern Natu - Caribbean Journal of Science 39:128–135. ralist 15:729–741. GAMBLE , T., E. G REENBAUM , T.R. J ACKMAN , A.P. R USSELL AND DOBSON , A.P. AND S.W. P ACALA . 1992. The parasites of Anolis A.M. B AUER . 2012. Repeated origin and loss of adhesive lizards in the northern Lesser Antilles. II. The structure of toepads in geckos. PLoS One 7:e39429. the parasite community. Oecologia 91:118–125. GIBBON , J.W., D.E. S COTT , T.J. R YAN , K.A. B UHLMANN , DOBSON , A.P., S.V. P ACALA , J.D. R OUGHGARDEN , E.R. C ARPER T.D. T UBERVILLE , B.S. M ETTS , J.L. G REENE , T. M ILLS , Y. L EI - AND E.A. H ARRIS . 1992. The parasites of Anolis lizards in the DEN , S. P OPPY ET AL . 2000. The Global Decline of Rep - northern Lesser Antilles. Oecologia 91:110–117. tiles, Déjà Vu Amphibians: Reptile species are declining DORNBURG , A., E. F ORRESTEL , J. M OORE , T. I GLESIAS , A. J ONES , on a global scale. Six significant threats to reptile popu - L. R AO AND D. W ARREN . 2017. An assessment of sampling lations are habitat loss and degradation, introduced biases across studies of diel activity patterns in marine ray- invasive species, environmental pollution, disease, unsus - finned fishes (Actinopterygii). Bulletin of Marine Science tainable use, and global climate change. BioScience 50: 93:611–639. 653–666.

Downloaded From: https://bioone.org/journals/Bulletin-of-the-Peabody-Museum-of-Natural-History on 11 Jul 2019 Terms of Use: https://bioone.org/terms-of-use Access provided by University of Mississippi 76 Bulletin of the Peabody Museum of Natural History 60(1) • April 2019

GOLDBERG , S.R. AND C.R. B URSEY . 1996. Gastrointestinal fornia Sur, Mexico. Bulletin of the Southern California helminths of the anole Anolis oculatus (Polychridae) from Academy of Sciences 102(3):143–147. Dominica, Lesser Antilles. Journal of the Helminthological GOLDBERG , S.R., C.R. B URSEY AND R.L. B EZY . 1996. Gastroin - Society of Washington 63:125–128. testinal helminths of Yarrow’s spiny lizard, Scleroporus jar - —1999. Crotalus lepidus (Rock Rattlesnake), Crotalus molossus rovii (Phrynosomatidae) in Mexico. American Midland (Blacktail Rattlesnake), Crotalus pricei (Twin-spotted Rat - Naturalist 135:299–309. tlesnake), Crotalus tigris (Tiger Rattlesnake). Endoparasites. GOLDBERG , S.R., C.R. B URSEY AND H. C HEAM . 1996. Gastroin - Herpetological Review 30:44–45. testinal helminths of six anole species, Anolis armouri , A. —2000a. Phrynosoma ditmarsi (Rock Horned Lizard). barahonae , A. bahorucoensis , A. brevirostris , A. chlorocyanus Endoparasites. Herpetological Review 31:242. and A. coelestinus (Polychrotidae) from Hispaniola. —2000b. Crotalus mitchellii (Speckled Rattlesnake) and Cro - Caribbean Journal of Science 32:112–115. talus willardi (Ridgenose Rattlesnake). Endoparasites. Her - —1997a. Helminths of Anolis acutus (Sauria: Polychrotidae) petological Review 31:104. from St. Croix, U.S. Virgin Islands. Journal of Parasitology —2000c. Micruroides euryxanthus (Western Coral Snake). 83:530–531. Endoparasites. Herpetological Review 31:105–106. —1997b. Helminths of 12 species of Anolis lizards (Polychroti - —2001. Hypsiglena torquata (Night Snake) and Oxybelis aeneus dae) from the Lesser Antilles, West Indies. Journal of the (Brown Vine Snake). Endoparasites. Herpetological Review Helminthological Society of Washington 64:248–257. 32:263. —1997c. Helminths from the Sonoran spotted whiptail, Cnemi - —2004a. Lepidoblepharis xanthostigma (Orange-tailed Gecko). dophorus sonorae , and the western whiptail, Cnemidopho - Endoparasites. Herpetological Review 35:268. rus tigris (Sauria: Teiidae), from southern Arizona with —2004b. Coelomic metazoan endoparasites of 15 colubrid and comments on Abbreviata terrapenis (Nematoda: Physa - two elapid snake species from Costa Rica. Caribbean Jour - lopteridae). Great Basin Naturalist 57:273–277. nal of Science 40:62–69. —1998a. Helminths of the lizard Anolis cristatellus (Polychroti - —2004c. Coelomic endoparasites in four species of colubrid dae) from the British Virgin Islands, West Indies. Journal of snakes, Drymobius margaritiferus , Masticophis mentovar - the Helminthological Society of Washington 65:259–262. ius , Salvadora mexicana and Trimorphodon tau from Mex - —1998b. Helminths of six species of Anolis lizards (Polychroti - ico. Bulletin of the Maryland Herpetological Society dae) from Hispaniola, West Indies. Journal of Parasitology 40:179–183. 84:1291–1295. —2004d. Conopsis lineata (Lined Tolucan Ground Snake) and —1999. Composition of the helminth community of a mon - Leptodeira maculata (Southwestern Cat-eyed Snake). tane population of the side-blotched lizard, Uta stansburi - Endoparasites. Herpetological Review 35:59. ana (Phrynosomatidae) from Los Angeles County, —2004e. Diadophis punctatus (Ring-necked Snake). Endopar - California. American Midland Naturalist 141:204–208. asites. Herpetological Review 35:64. GOLDBERG , S.R., C.R. B URSEY AND H.J. H OLSHUH . 1998. Preva - —2004f. Oligacanthorhynchid acanthocephalan cystacanths in lence and distribution of cystacanths of an oligacan - the Yaqui blackhead snake, Tantilla yaquia (Serpentes: Col - thorhynchid acanthocephalan from the longnose snake, ubridae) from Arizona. Journal of the Arizona-Nevada Rhinocheilus lecontei (Colubridae), in southwestern North Academy of Science 37:83–84. America. Journal of the Helminthological Society of Wash - —2005. Coelomic helminths in the Speckled Racer, Drymobius ington 65:262–265. margaritiferus (Serpentes, Colubridae) from Central Amer - GOLDBERG , S.R., C.R. B URSEY AND F. K RAUS . 2009. Endoparasites ica. Texas Journal of Science 57:103–106. in 12 species of Sphenomorphus (Squamata: Scincidae) from —2007. Coelomic helminths of five colubrid snake species (Ser - Papua New Guinea. Comparative Parasitology 76:58–83. pentes, Colubridae) from Costa Rica. Phyllomedusa: Journal GOLDBERG , S.R., C.R. B URSEY AND C.T. M CALLISTER . 1995. Gas - of Herpetology 6:69–72. trointestinal helminths of nine species of Sceloporus lizards —2009a. Ameiva undulata (Rainbow Ameiva). Endoparasites. (Phrynosomatidae) from Texas. Journal of the Helmintho - Herpetological Review 40:82. logical Society of Washington 62:188–196. —2009b. Imantodes cenchoa (Blunt-headed Tree Snake), Iman - GOLDBERG , S.R., C.R. B URSEY AND C.W. P AINTER . 2002. todes gemmistratus (Central American Tree Snake), Iman - Helminths of the western diamondback rattlesnake, Crotalus todes inornatus (Western Tree Snake). Endoparasites. atrox , from southeast New Mexico rattlesnake roundups. Herpetological Review 40:230. Southwestern Naturalist 47:307–310. —2011. Ameiva leptophrys (Delicate Ameiva). Endoparasites. GOLDBERG , S.R., C.R. B URSEY AND R. T AWIL . 1994. Helminth Herpetological Review 42:271. parasites of the bark anole, Anolis distichus and the brown —2012a. Endohelminths of the Socorro Island Tree Lizard, anole, Anolis sagrei (Polychridae) from Florida and the Urosaurus auriculatus (Squamata: Phrynosomatidae), from Bahamas. Caribbean Journal of Science 30:275–277. Colima, Mexico. Comparative Parasitology 79:269–274. GOLDBERG , S.R., C.R. B URSEY AND S.R. T ELFORD JR. 2003. Meta - —2012b. Ameiva bifrontata (Cope’s Ameiva). Endoparasites. zoan endoparasites of 11 species of lizards from Pakistan. Herpetological Review 43:477. Comparative Parasitology 70:46–54. —2012c. Plestiodon multivirgatus (Many-lined Skink). Endopar - HECKMANN , R.A., A. K HAN AND O.M. A MIN . 2011. Juveniles of asites. Herpetological Review 43:4. genus Oligacanthorhynchus Travassos, 1915 (Acantho - GOLDBERG , S.R., C.R. B URSEY AND K.M. B EAMAN . 2003. Gas - cephala: Oligacanthorhynchidae Southwell & Macfie, 1925) trointestinal helminths of the black-tailed brush lizard, from snakes in Karachi, Pakistan. Pakistan Journal of Zool - Urosaurus nigricaudus (Phrynosomatidae), from Baja Cali - ogy 43:604–606.

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KRASNOSHCHEKOV , G. AND O. L ISITSYNA . 2009. On the migra - MCALLISTER , C.T., C.R. B URSEY AND M.B. C ONNIOR . 2015. tion of cystacanths Sphaerirostris picae (Acanthocephala, Helminth parasites (Trematoda, Cestoda, Nematoda, Acan - Centrorhynchidae) in paratenic host Lacerta agilis , thocephala) of herpetofauna from southeastern Oklahoma: histopathology. Vestnik Zoologii 43:433–440. New host and geographic records. Proceedings of the Okla - KUTT , A.S. 2011. The diet of the feral cat ( Felis catus ) in north- homa Academy of Science 95:125–134. eastern Australia. Acta Theriologica 56:157–169. —2017. New host and distributional records for parasites (Api - LAMAS , M.F. AND V.H. Z ARACHO . 2006. Tropidurus torquatus complexa, Trematoda, Nematoda, Acanthocephala, Aca - (Brown Lizard) . Endoparasites. Herpetological Review rina) of Texas herpetofauna. Comparative Parasitology 37:474–475. 84:42–50. LAMB , A.D., G.J. W ATKINS -C OLWELL , J.A. M OORE , D.L. W AR - MCALLISTER , C.T., C.R. B URSEY AND J.F. R OBERTS . 2004. REN , T.L. I GLESIAS , M.C. B RANDLEY AND A. D ORNBURG . 2017. Physocephalus sexalatus (Nematoda: Spirurida: Spirocer - Endolymphatic sac use and reproductive activity in the cidae) in three species of rattlesnakes, Crotalus atrox , Cro - Lesser Antilles endemic gecko Gonatodes antillensis talus lepidus , and Crotalus scutulatus , from southwestern (Gekkota: Sphaerodactylidae). Bulletin of the Peabody Texas. Journal of Herpetological Medicine and Surgery Museum of Natural History 58:17–29. 14:10–12. MAHAGEDARA , P.D. AND R.S. R AJAKARUNA . 2015. Parasites of MCALLISTER , C.T., J.E. C ORDES AND J.M. W ALKER . 1991. two co-occurring house gecko species, Hemidactylus frena - Helminth parasites of unisexual and bisexual whiptail lizards tus and Gehyra mutilata from Central Sri Lanka. Ving - (Teiidae) in North America. 6. The gray checkered whiptail nanam Journal of Science 11:32–38. (Cnemidophorus dixoni ). Texas Journal of Science 43: MARCOGLIESE , D.J. 2002. Food webs and the transmission of 309–314. parasites to marine fish. Parasitology 124:83–99. —1995. Helminth parasites of unisexual and bisexual whiptail —2007. Evolution of parasitic life in the ocean: Paratenic hosts lizards (Teiidae) in North America. IX. The plateau spotted enhance lateral incorporation. Trends in Parasitology whiptail ( Cnemidophorus gularis septemvittatus ). Texas Jour - 23:519–521. nal of Science 47:83–88. MARKOV , G.S., O.P. B OGDANOV , V.M. M AKEEV AND A.A. MCALLISTER , C.T., J. R ILEY , P.S. F REED AND D.A. F REED . 1993. KHUTORYANSKI . 1968. New data on the helminth fauna of Endoparasites of some Malagasy colubrids (Reptilia: Ser - Naja oxiana . Izdatel’stvo Akademii Nauk SSSR 1968:244– pentes), with descriptions of two new species of Raillietiella 248. (in Russian) (Pentastomida: Cephalobaenida). Transactions of the Amer - MARKOV , G.S., O.P. B OGDANOV AND L.A. P ERSIANOVA . 1970. ican Microscopical Society 112:35–42. [Helminth fauna of the carpet viper]. In: Yadovitye zhivot - MEDINA , F.M., E. B ONNAUD , E. V IDAL , B.R. T ERSHY , E.S. nye srednei Azii i ikh yady. (Materialy Sredneaziatskoi Kon - ZAVALETA , C. J OSH DONLAN , B.S. K EITT , M. C ORRE , S.V. ferentsii; 1–3 Oktyabrya 1968). Tashkent, USSR: Izdat HORWATH AND M. N OGALES . 2011. A global review of the "FAN". pp. 46–60. (in Russian) impacts of invasive cats on island endangered vertebrates. MARKOV , G.S., M.P. Z INYAKOVA AND A.S. L UTTA . 1967. New data Global Change Biology 17:3503–3510. in the parasitology of Central Asiatic Vipera lebetina . Sbornik MIRANDA , J.P. AND G.V. A NDRADE . 2003. Seasonality in diet, Nauchnykh Rabot Volgogradskogo Pedagogicheskogo Insti - perch use, and reproduction of the gecko Gonatodes humer - tuta AS Serafimovicha 1967:98–107. (in Russian) alis from Eastern Brazilian Amazon. Journal of Herpetology MARTEAU , M. 1977. [ Oncicola venezuelensis n. sp. (Archiacan - 37:433–438. thocephala-Oligacanthorhynchida), a parasite of the ocelot MURVANIDZE , L., T.S. L OMIDZE , K. N IKOLAISHVILI AND E. (Felis pardalis L.)]. Annales de Parasitologie Humaine et JANKARASHVILI . 2008. The annotated list of reptile helminths Comparée 52:25–33. (in French) of Georgia. Proceedings of the Institute of Zoology (Tbilisi) MAYéN -P EñA , E. AND G. S ALGADO -M ALDONADO . 1998. 23:54–61. Helminths of four lizards from Nayarit, México: Anolis neb - NICKOL , B.B. AND T.T. D UNAGAN . 1989. Reconsideration of the ulosus (Polychrotidae), Ctenosaura pectinata (Iguanidae), acanthocephalan genus Echinopardalis , with a description Phyllodactylus lanei (Gekkonidae), and Sceloporus nelsoni of adult E. atrata and a key to genera of the Oligacan - (Phrynosomatidae). Journal of the Helminthological Society thorhynchidae. Proceedings of the Helminthological Society of Washington 65:108–111. of Washington 56:8–13. MCALLISTER , C.T. 1990a. Helminth parasites of unisexual and NICKOL B.B., C.A. F ULLER AND P. R OCK . 2006. Cystacanths of bisexual whiptail lizards (Teiidae) in North America. IV. The Oncicola venezuelensis (Acanthocephala: Oligacan - Texas spotted whiptail ( Cnemidophorus gularis ). Texas Jour - thorhynchidae) in Caribbean termites and various paratenic nal of Science 42:381–388. hosts in the U.S. Virgin Islands. Journal of Parasitology —1990b. Helminth parasites of unisexual and bisexual whiptail 92:539–542. lizards (Teiidae) in North America. II. The New Mexico OLUWAFEMI , T., O. S OWEMIMO AND A. B AMIDELE . 2017. Para - whiptail ( Cnemidophorus neomexicanus ). Journal of Wildlife sitic infections of two species of lizard ( Hemidactylus frena - Diseases 26:403–406. tus and Mabuya quinquetaeniata ) from two cities in MCALLISTER , C.T. AND C.R. B URSEY . 2007. Some nematode and Southwest Nigeria. Annual Research & Review in Biology acanthocephalan parasites of the longnose leopard lizard, 18(2):1–13. Gambelia wislizenii (Lacertilia: Crotaphytidae), from Ari - PARKER , G.A., M.A. B ALL AND J.C. C HUBB . 2015. Evolution of zona, California, and Texas, with a summary of the complex life cycles in trophically transmitted helminths. I. helminths reported from this host. Comparative Parasitol - Host incorporation and trophic ascent. Journal of Evolution - ogy 74:179–184. ary Biology 28:267–291.

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PATTON , S., A. R ABINOWITZ , S. R ANDOLPH AND S.S. J OHNSON . SCHMIDT , G.D. 1972. Revision of the Class Archiacantho - 1986. A coprological survey of parasites of wild neotropical cephala Meyer, 1931 (Phylum Acanthocephala), with felidae. Journal of Parasitology 72:517–520. emphasis on Oligacanthorhynchidae Southwell et Macfie, PETROCHENKO , V.I. 1956. Acanthocephala of Domestic and 1925. Journal of Parasitology 58:290–297. Wild Animals, Volume I. Translated from Russian by Israel —1985. Development and life cycles. In: D.W.T. Crompton and Program for Scientific Translations, Jerusalem, 1971. Wash - B.B. Nickol, eds. Biology of the Acanthocephala. Cambridge: ington, DC: U.S. Department of Agriculture and the Cambridge University Press. pp. 273–305. National Science Foundation. 465 pp. SCHMIDT , G.D. AND R.E. K UNTZ . 1969. Centrorhynchus spilor - PIZZATTO , L. AND R.R. M ADI . 2002. Micrurus corallinus (Coral nae sp. n. (Acanthocephala), and other Centrorhynchidae Snake). Endoparasites. Herpetological Review 33:215. from the Far East. Journal of Parasitology 55:329–334. PIZZATTO , L. AND O.A. M ARQUES . 2006. Interpopulational vari - SHARPILO , V. 1962. To the study of helminthofauna of racers of ation in sexual dimorphism, reproductive output, and par - Transcaucasus. Zbornik prac zoologicheskogo muzeiu AN asitism of Liophis miliaris (Colubridae) in the Atlantic forest USSR 31:63–69. (in Ukrainian) of Brazil. Amphibia-Reptilia 27:37–46. —1976. The Parasitic Helminths of Reptiles of Fauna of USSR. PLAZA , P.I. AND S.A. L AMBERTUCCI . 2017. How are garbage Kiev: Naukova Dumka. 286 pp. (in Russian) dumps impacting vertebrate demography, health, and con - SHARPILO , V.P., V. B ISERKOV , A. K OSTADINOVA , J.M. B EHNKE AND servation? Global Ecology and Conservation 12:9–20. Y.I. K UZMIN . 2001. Helminths of the sand lizard, Lacerta PRUDHOE , S. AND M.T. H ARRIS . 1971. Helminths from agilis (Reptilia, Lacertidae), in the Palaearctic: Faunal diver - chameleons in southern Africa. Revue de Zoologie et de sity and spatial patterns of variation in the composition Botanique Africaines 84:334–343. and structure of component communities. Parasitology QUESNEL , V.C. 1957. Life history of the Streak Lizard, Gona - 23:389–400. todes vittatus . Living World: Journal of the Trinidad and SILVA , L.A. 2014. Helmintofauna associada a répteis prove - Tobago Field Naturalists’ Club 1957:5–14. nientes da Reserva Particular do Patrimônio Natural Foz do QUIRINO , T.F., A.J.M.G. F ERREIRA , M.C. S ILVA , R.J. S ILVA , D.H. Rio Aguapeí, Estado de São Paulo [dissertation]. Sao Paulo: MORAIS AND R.W. Á VILA . 2018. New records of helminths in Universidade Estadual Paulista–Campus de Botucatu. Avail - reptiles from five states of Brazil. Brazilian Journal of Biology able from: https://repositorio.unesp.br/bitstream/handle/ 78:750–754. 11449/110418/000786523.pdf RENGARAJU , V. AND E.N. D AS . 1981. On a new species of the SIMONSEN , P.E. AND R.K. S ARDA . 1985. Helminth and arthropod genus Oligacanthorhynchus Travassos, 1915 (Acantho - parasites of Hemidactylus mabouia from Tanzania. Journal cephala: Oligacanthorhynchidae) from India. Records of the of Herpetology 19:428–430. Zoological Survey of India 78:37–40. SMALES , L.R. 1997. A review of the helminth parasites of Aus - REP , B.H. 1975. Intestinal helminths in dogs and cats on the tralian rodents. Australian Journal of Zoology 45:505–521. Antillian Islands Aruba, Curaçao and Bonaire. Tropical and —2007. Acanthocephala in amphibians (Anura) and reptiles Geographical Medicine 27:317–323. (Squamata) from Brazil and Paraguay with description of a RICHARDSON , D.J. AND B.B. N ICKOL . 1995. The genus Cen - new species. Journal of Parasitology 93:392–398. trorhynchus (Acanthocephala) in North America with DE SOUZA , O.F.F. AND V.K. B ROWN . 1994. Effects of habitat frag - description of Centrorhynchus robustus n. sp., redescription mentation on Amazonian termite communities. Journal of of Centrorhynchus conspectus , and a key to species. Journal Tropical Ecology 10:197–206. of Parasitology 81:767–772. STRUBE , C., L. H EUER AND E. J ANECEK . 2013. Toxocara spp. ROCA , V. 1995. An approach to the knowledge of the helminth infections in paratenic hosts. Veterinary Parasitology infracommunities of Mediterranean insular lizards ( Podar - 193:375–389. cis spp.). Scientia Herpetologica 1995:285–292. TELFORD , S.R. 1997. The Ecology of a Symbiotic Community, ROCA , V., F. J ORGE AND M.Á. C ARRETERO . 2012. Synopsis of the Volume 2. The Component Symbiote Community of the helminth communities of the lacertid lizards from the Japanese Lizard Takydromus tachydromoides (Schlegel) (Lac - Balearic and Canary Islands. Basic and Applied Herpetol - ertidae). Malabar, FL: Krieger Publishing Company. 153 pp. ogy 26:107–116. TRAVASSOS , L. 1917. Contribuições para o conhecimento da ROCA , V., J.E. M ARTIN AND E. C ARBONELL . 1999. Helminths fauna helmintolojica brazileira. Memórias do Instituto parasitising endemic geckoes from Canary Islands. Miscel - Oswaldo Cruz 9:5–62. lània Zoològica 22:101–108. VAN BUURT , G. 2005. Field Guide to the Amphibians and Rep - RODRIGUES , H.O. 1970. Estudo da fauna helmintologica de tiles of Aruba, Curaçao and Bonaire. Frankfurt am Main, Hemidactylus mabouia (M. de J.) no Estado da Guanabara. Germany: Chimaira. 160 pp. Atas da Sociedade Biologica do Rio de Janeiro 12(Suppl.): VAN BUURT , G. AND A.O. D EBROT . 2012. Introduced Agricul - 15–23. tural Pests, Plant and Animals Diseases and Vectors in the SAEHOONG , P. AND C. W ONGSAWAD . 1997. Helminths in house Dutch Caribbean, with an "Alert Species" List. Wageningen, lizards (Reptilia: Gekkonidae). Southeast Asian Journal of Netherlands: Institute for Marine Resources & Ecosystem Tropical Medicine and Public Health 28:184–189. Studies. 35 pp. (IMARES Report C193/11.) SANTOS , E.G.N., M. C HAME , V.A. C HAGAS -M OUTINHO AND C.P. VELIKANOV , V.P . 1989. The role of amphibians and reptiles like SANTOS . 2017. Morphology and molecular analysis of Onci - intermediate and paratenic hosts of helminths in the condi - cola venezuelensis (Acanthocephala: Oligacanthorhynchi - tions of the Turkmenian SSR. Izvestiya Akademii Nauk dae) from the ocelot Leopardus pardalis in Brazil. Journal of Turkmenskoi SSR Seriya Biologicheskikh Nauk 6:43–49. (in Helminthology 91:605–612. Russian)

Downloaded From: https://bioone.org/journals/Bulletin-of-the-Peabody-Museum-of-Natural-History on 11 Jul 2019 Terms of Use: https://bioone.org/terms-of-use Access provided by University of Mississippi Geckos Paratenic Hosts for Caribbean Island Acanthocephalans • Dornburg et al. 79

VICENTE , J.J. 1978. Helmintos de Tropidurus (Lacertilia: (Scincidae) in Queimada Grande Island, Sao Paulo, Brazil. Iguanidae) da Coleço Helmintológica do Instituto Oswaldo Revista Brasileira de Zoologia 22:1185–1190. Cruz, I. Trematoda, Cestoda, Acanthocephala, Linguatul - VRCIBRADIC , D., C.F.D. R OCHA , C.R. B URSEY AND J.J. V ICENTE . ida. Atas da Sociedade de Biologia do Rio de Janeiro 2002. Helminth communities of two sympatric skinks 19:71–78. (in Portuguese) (Mabuya agilis and Mabuya macrorhyncha ) from two VITT , L.J., R.A. S OUZA , S.S. S ARTORIUS , T.C.S. A VILA -P IRES AND ‘restinga’ habitats in southeastern Brazil. Journal of M.C. E SPóSITO . 2000. Comparative ecology of sympatric Helminthology 76:355–361. Gonatodes (Squamata: Gekkonidae) in the western Amazon VRCIBRADIC , D., J.J. V ICENTE AND C.R. B URSEY . 2007. Helminths of Brazil. Copeia 2000:83–95. infecting the lizard Enyalius bilineatus (Iguanidae; Leiosauri - VOGEL , P. AND D.A.P. B UNDY . 1987. Helminth parasites of nae) from an Atlantic Rainforest area in Espírito Santo state, Jamaican anoles (Reptilia: Iguanidae): Variation in preva - southeastern Brazil. Amphibia-Reptilia 28:166–169. lence and intensity with host age and sex in a population of WARD , H.L. 1940. Notes on juvenile Acanthocephala. Journal of Anolis lineatopus . Parasitology 94:399–404. Parasitology 26:191–194. VRCIBRADIC , D., L.A. A NJOS , J.J. V ICENTE AND C.R. B URSEY . WATKINS -C OLWELL , G.J., J. DE LEON AND S.B. H OCHGRAFF . 2008. Helminth parasites of two sympatric lizards, Enyalius 2006. Leptotyphlops tenellus (Guyana Blindsnake). Preda - iheringii and E. perditus (Leiosauridae), from an Atlantic tion. Herpetological Review 37:353. Rainforest area of southeastern Brazil. Acta Parasitologica YILDIRIMHAN , H.S., N. S üMER AND N. Y ILMAZ . 2008. The 53:222–225. helminth fauna of Hemidactylus turcicus (Mediterranean VRCIBRADIC , D. AND C.F. R OCHA . 2005. Observations on the Gecko) (Linnaeus, 1758) from Hatay Province. Acta Para - natural history of the lizard Mabuya macrorhyncha Hoge sitologica Turcica 32:393–395.