-Associated from Dominica and the Bahamas With Notes on Hyperparasitic Erythraeid Author(s): Lance A. Durden, Charles R. Knapp, Lorenza Beati, and Stephanie Dold Source: Journal of Parasitology, 101(1):24-27. Published By: American Society of Parasitologists DOI: http://dx.doi.org/10.1645/14-602.1 URL: http://www.bioone.org/doi/full/10.1645/14-602.1

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BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit publishers, academic institutions, research libraries, and research funders in the common goal of maximizing access to critical research. J. Parasitol., 101(1), 2015, pp. 24–27 Ó American Society of Parasitologists 2015

REPTILE-ASSOCIATED TICKS FROM DOMINICA AND THE BAHAMAS WITH NOTES ON HYPERPARASITIC ERYTHRAEID MITES

Lance A. Durden, Charles R. Knapp*, Lorenza Beati†, and Stephanie Dold‡ Department of Biology, Georgia Southern University, Statesboro, Georgia 30458. Correspondence should be sent to: [email protected]

ABSTRACT: Ticks were collected or recorded from 522 individual on Dominica and from 658 reptiles from the Bahamas. Two species of ticks were collected on Dominica: Amblyomma antillorum and Amblyomma rotundatum. Similarly, 2 species were collected in the Bahamas: Amblyomma albopictum and Amblyomma torrei. On Dominica, A. antillorum was recorded from 517 Lesser Antillean iguanas (Iguana delicatissima), 2 boa constrictors (Boa nebulosa), 1 Antilles snake (Alsophis sibonius), and 1 Dominican ground lizard (Ameiva fuscata), whereas A. rotundatum was recorded from 1 Lesser Antillean ( mabouya). In the Bahamas, A. albopictum was recorded from 131 Andros iguanas (Cyclura cychlura cychlura), 271 Exuma Island iguanas (Cyclura cychlura figginsi), and 1 Andros curlytail lizard (Leiocephalus carinatus coryi), whereas A. torrei was recorded from 255 Exuma Island iguanas. In the Bahamas, A. albopictum parasitized iguanas on Andros Island and the central Exuma Islands, and A. torrei parasitized iguanas in the southern Exumas. An exception to this trend was that A. torrei was collected from iguanas on Pasture Cay in the central Exumas, an anomaly that is explained by the fact that iguanas (with attached ticks) on Pasture Cay were introduced by humans in the past from islands further south. External hyperparasitic larval erythraeid mites ( sp.) were recorded from A. torrei in the Bahamas.

Reptiles in the Caribbean region are parasitized by a unique we counted ticks from each individual and collected a random sample suite of amblyommine ticks, including Amblyomma albopictum from a random selection of iguanas including both host sexes and all age classes. This study was approved by the Zoological Society of San Diego Neumann, Amblyomma antillorum Kohls, Amblyomma cruciferum Institutional Care and Use Committee (project approval #267) Neumann, Amblyomma dissimile Koch, Amblyomma quadricavum and the John G. Shedd Aquarium Research Committee. (Schulze), Amblyomma rotundatum Koch, and Amblyomma torrei Ectoparasites were collected primarily from 1 iguana species in the Vigeuras (Robinson, 1926; Whittick, 1939; Clifford and Kohls, Bahamas, Cyclura cychlura, which is listed as ‘‘vulnerable’’ according to the International Union for Conservation of Nature (Knapp et al., 2004), 1962; Cerny,´ 1969a; Kohls, 1969a, 1969b; Keirans, 1985; Keirans although the subspecies Cyclura cychlura cychlura and Cyclura cychlura and Garris, 1986; Keirans and Klompen, 1996; Guglielmone et figginsi are listed as ‘‘endangered’’ and ‘‘critically endangered,’’ respec- al., 2003; Durden and Knapp, 2005). Of these species, A. tively. Ectoparasites from reptile hosts on Andros Island and on the albopictum, A. antillorum, A. cruciferum, and A. torrei are known Exuma Islands, Bahamas, were collected during March, May, and/or to be ectoparasitic specialists of iguanas (Robinson, 1926; June. Specific sampling localities on Andros Island were Alcorine Cay (24808.5320N, 77850.9460W), Mangrove Cay (24809.4290N, 77844.8340W), Keirans, 1985; Guglielmone et al., 2003). However, the geograph- Sandy Cay (24805.1440N, 77841.6350W), and Ratman Cay (23841.6620N, ical distribution of these species on the various Caribbean 77836.9400W). Specific localities in the Exuma Islands were Bitter Guana islands and their host associations are incompletely known. Cay (24808.7370N, 76825.0890W), Gaulin Cay (24807.3350N, 76824.3460W), 0 0 Additionally, to our knowledge, no external hyperparasites of any Noddy Cay (23848.047 N, 76808.412 W), North Adderly Cay (23847.5280N, 76807.3870W), Pasture Cay (24819.0830N, 76833.5810W), of these iguana-associated ticks have been reported previously. and White Bay Cay (23847.9190N, 76808.7340W). In this paper, we document the host associations for 2 of the Ectoparasites from reptile hosts on Dominica were collected in April, above-listed tick species, A. albopictum and A. torrei, from the May, June, July, August, and/or September from the west coast of the Bahamas, and for 1 congener, A. antillorum, from Dominica. We island primarily in 2 study locations—Batali Beach (15826.9380N, 0 0 0 also document A. rotundatum from a skink in Dominica and 61826.871 W) and Champagne Bay (15814.782 N, 61822.357 W). Ectopar- asites were collected primarily from 1 species, the Lesser Antillean Iguana report ectoparasitic erythraeid mites from A. torrei in the (Iguana delicatissima), which was common historically in the northern Bahamas. We also briefly discuss the known geographical Lesser Antilles. However, its range has diminished dramatically, and thus distributions of the Caribbean iguana-associated ticks, including the iguana is listed as ‘‘endangered’’ by the International Union for the intriguing apparently segregated distributions of A. albopic- Conservation of Nature (Breuil et al., 2010). tum and A. torrei in the Bahamas. Morphological identification of ticks Adult ticks were identified based on morphological characters MATERIALS AND METHODS following Robinson (1926), Whittick (1939), Clifford and Kohls Tick collections (1962), and Kohls (1969b). Some immature ticks were identified according to characters or illustrations in Whittick (1939), Cerny´ As part of a larger study on reptile ecology, ectoparasites were collected (1966, 1969b), Keirans (1985), and Keirans and Oliver (1993). Identities from reptiles in the Bahamas and on Dominica from 2006 to 2011. Ticks of several immature ticks were confirmed or determined using molecular were removed with fine forceps or counted from live, restrained reptiles. methods (see following section). Voucher tick specimens from this study Collections focused on iguanas, whereas removal from other reptiles was have been deposited in the U.S. National Tick Collection (curated at opportunistic. Ectoparasites were removed and stored in labeled vials Georgia Southern University) under accession numbers RML124486– containing 70% ethanol. Instead of removing all ticks from each iguana, RML124296 and RML124442–RML124448.

Received 2 July 2014; revised 22 September 2014; accepted 30 September Molecular identification of immature ticks 2014. Because the nymphal stage of A. albopictum has not been described, and * Daniel P. Haerther Center for Conservation and Research, John G. original descriptions of the immature stages of some other tick species that Shedd Aquarium, 1200 S. Lake Shore Drive, Chicago, Illinois 60605. we collected are inadequate, we used molecular techniques to confirm or † U.S. National Tick Collection, Institute of Coastal Plain Science, determine identifications of these stages by matching DNA sequences with Georgia Southern University, Statesboro, Georgia 30460-8056. identified adults. This technique has proven successful with a number of ‡ Museum of Biological Diversity, The Ohio State University, 1315 exotic Amblyomma species (Mukherjee et al., 2014). Immature stages of A. Kinnear Road, Columbus, Ohio 43212-1192. albopictum and A. torrei will be formally described or redescribed, as DOI: 10.1645/14-602.1 appropriate, in future publications, which will also include molecular

24 DURDEN ET AL.—REPTILE TICKS AND MITES, DOMINICA, BAHAMAS 25

TABLE I. Ticks collected from reptiles from Dominica and the Bahamas, 2006–2011 (tick sample sizes from iguanas were derived from a subsample of ticks from randomly chosen iguanas).

Location Host Tick species* (no.) Mean intensity† 6 SD (range)

Dominica: Batali, Champagne Iguana delicatissima Laurenti (Lesser Amblyomma antillorum (158?,91/, 75N, 19.6 6 40.5 (1–267) Bay, Coulibistri, Sunset Bay, Antillean iguana) (ticks recorded 197L) and Vidal farm from 517 of 1,149 individuals) Boa nebulosa Lazell (boa Amblyomma antillorum (2?) constrictor) (n ¼ 2) Alsophis sibonius Cope (Antilles Amblyomma antillorum (6L) snake) (n ¼ 1) Ameiva fuscata Garman (Dominican Amblyomma antillorum (6L) ground lizard) (n ¼ 1) Mabuya mabouya Bonnaterre Amblyomma rotundatum (2N,10L) (Lesser Antillean skink) (n ¼ 1) Bahamas: Andros Island Cyclura cychlura cychlura (Cuvier) Amblyomma albopictum (202?,78/, 21.8 6 27.1 (1–200) (Andros iguana) (ticks recorded 68N, 22L) from 131 of 152 individuals) Leiocephalus carinatus coryi Schmidt Amblyomma albopictum (2N, 1L) (Andros curlytail lizard) (n ¼ 1) Bahamas: Central Exumas (Bitter Cyclura cychlura figginsi Barbour Amblyomma albopictum (208?,93/, 9.1 6 9.2 (1–79) Guana and Gaulin Cays) (Exuma iguana) (ticks recorded 96N, 21L) from 271 of 292 individuals) Bahamas: Central Exumas Cyclura cychlura figginsi Barbour Amblyomma torrei (37?,16/, 24N) 19.8 6 16.1 (2–54) (Pasture Cay) (Exuma iguana) (ticks recorded from 17 of 17 individuals) Bahamas: Southern Exumas Cyclura cychlura figginsi Barbour Amblyomma torrei (131?, 117/, 130N, 12.5 6 14.2 (1–80) (Noddy, North Adderly, and (Exuma iguana) (ticks recorded 39L) White Bay Cays) from 238 of 244 individuals)

*N¼ nymph(s), L ¼ larva(e). † Mean intensity ¼ mean per infested host following Bush et al. (1997) (only calculated for host sample sizes of 8 or greater). aspects of their phylogenetic and taxonomic placement. Photographs of Hyperparasitic erythraeid mites erythraeids attached to A. torrei were taken using a Visionary Digital K2/ SC long-distance microscope (Infinity Photo-Optical Company, Boulder, Larval prostigmatan mites belonging to the Leptus Colorado). Selected unattached larval erythraeids were cleared in (family Erythraeidae) were found attached to 8 A. torrei ticks lactophenol, mounted in Hoyer’s medium, and ringed with Glyptal (Fig. 1) from the Bahamas, with an additional 5 larvae found following standard techniques (Krantz and Walter, 2009). unattached in vials that contained A. torrei. These unattached larvae presumably had been dislodged from A. torrei specimens RESULTS during transit in these vials. Based on morphological characters, Ticks all erythraeid specimens retrieved are conspecific. Ticks (A. antillorum) were recorded from 517 of 1,149 (45%) DISCUSSION captured iguanas and from 3 snakes and 2 lizards on Dominica (Table I). Another tick species (A. rotundatum) was recorded from In this paper, we provide new host association data for A. 1 lizard on Dominica (Table 1). Ticks (A. albopictum or A. torrei) antillorum from I. delicatissima, B. nebulosa, A. sibonius, and A. were recorded from 131 of 152 (86%) Andros Island iguanas and fuscata on Dominica and for A. albopictum from L. carinatus from 238 of 244 (98%) Exuma Island iguanas inhabiting the coryi in the Bahamas. Our collection data for both of these tick southern Exuma Islands, and from 271 of 292 (93%) Exuma species and for A. torrei, all of which were much more abundant Island iguanas inhabiting the central Exuma Islands. We also on iguanas than on other reptiles, confirm that these 3 ticks are collected ticks (A. albopictum) from a curlytail lizard (Leiocepha- principally ectoparasites of Caribbean iguanas, although A. lus carinatus coryi) from Andros Island (Table I). Mean intensities albopictum has also been reported from parts of Central and of ticks parasitizing iguanas ranged from 9.1 for A. albopictum on South America (Guglielmone et al., 2003). To our knowledge, our C. cychlura figginsi in the central Exumas to 21.8 for A. record of A. rotundatum from Mabuya mabouya on Dominica is a albopictum on C. cychlura cychlura on Andros Island in the new host association and island record, although this widespread Bahamas (Table I). In the Bahamas, A. albopictum was recorded tick has been recorded from a fairly large number of reptiles and from iguanas on Andros Island and in the central Exuma Islands, several amphibian species in the Neotropical region (Keirans and whereas A. torrei was recorded from iguanas in the southern Oliver, 1993; Guglielmone et al., 2003). Exuma Islands. An exception to this trend was that A. torrei was Geographically, we recorded A. antillorum only from Dom- recorded from C. cychlura figginsi on Pasture Cay in the central inica, an island from which Kohls (1969a, 1969b) and Gugliel- Exumas (Table I). mone et al. (2003) also reported this tick. In agreement with most 26 THE JOURNAL OF PARASITOLOGY, VOL. 101, NO. 1, FEBRUARY 2015

geology and vegetation structure. The most southern A. albopictum and most northern A. torrei populations in the Exumas are separated by 46 km of sea and cays, which are uninhabited by iguanas. An exception to this biogeographic demarcation is the C. c. figginsi population on previously uninhabited Pasture Cay in the central Exumas, which is parasitized by A. torrei instead of A. albopictum. This anomaly, however, is due to a 2002 translocation of 16 iguanas (with attached A. torrei) originating from an island adjacent to North Adderly Cay in the southern Exumas (Knapp, 2002). An additional species of Caribbean iguana-associated tick that we did not record in this study is A. cruciferum. Given the restricted geographical distributions of 3 of the 4 known species of Caribbean iguana-associated ticks, combined with the rarity, endangered or threatened status of some of their hosts, these tick species (A. antillorum, A. cruciferum, and A. torrei) could be candidates for conservation action (Durden and Keirans, 1996). However, as long as their host populations remain viable and FIGURE 1. Hyperparasitic larval erythraeid (Leptus sp.) attached to the intersegmental membrane between the femur and tibia on the right ticks are not eliminated from iguanas, these tick species should fourth leg of a female Amblyomma torrei that was parasitizing an Exuma survive in adequate numbers. A fourth Caribbean iguana- iguana (Cyclura cychlura figginsi), southern Exumas, Bahamas. associated tick species, A. albopictum, has a wider range than the other 3 species and is presumably not in need of conservation of our collections, Kohls (1969b) recorded the endangered I. status. delicatissima as the host of A. antillorum on Dominica, although To our knowledge, hyperparasitic erythraeid mites have not we also recorded 2 snakes and 1 lizard species as secondary hosts been recorded previously from ticks. However, larval erythraeids (Table I). Previous records of A. antillorum are from the of various species are known to parasitize members of other endangered Anegada ground iguana (Cyclura pinguis Barbour) groups such as opilionids, scorpions, spiders, and from the island of Anegada in the British Virgin Islands (Kohls, insects (Young and Welbourn, 1987; Southcott, 1999; McAloon 1969b; Keirans, 1985) and from an iguana (likely the Turks and and Durden, 2000; Krantz and Walter, 2009). Further, there are Caicos rock iguana, Cyclura carinata carinata Harlan according records of larval erythraeids parasitizing oribatid mites (Norton et to Keirans [1985]) on East Caicos Island in the Turks and Caicos al., 1988), other prostigmatan mites (Wendt et al., 1992), and even Islands. other erythraeids (Young and Welbourn, 1987; Wendt et al., In this study, A. albopictum was mainly recorded from iguanas 1992). Larvae of some species of erythraeids are also ectoparasitic on Andros Island and the central Exuma Islands in the Bahamas, on vertebrates or are free-living (Southcott, 1999; Krantz and although 3 immature specimens were also recorded from 1 L. Walter, 2009). carinatus coryi on Andros Island (Table I). This tick has also been Clearly, Caribbean reptiles are parasitized by a unique recorded from iguanas on Cuba, the Dominican Republic, and community of acarine ectoparasites. Further research on these Haiti in the Caribbean region and also from Brazil, Costa Rica, host–parasite relationships is warranted. and Honduras (Clifford and Kohls, 1962; Morel, 1967; Cerny,´ 1969a, 1969b; Guglielmone et al., 2003). ACKNOWLEDGMENTS Amblyomma torrei has a restricted Caribbean distribution. Although Cerny´ (1966) stated that A. torrei is confined to Cuba, it We thank the Bahamas Environment, Science, and Technology has also been recorded from the Cayman Islands, Puerto Rico, Commission for permission to conduct the study in the Bahamas. We thank the Dominica Forestry, Wildlife and Parks Division for permission and the Bahamas (southern Exuma Islands and Pasture Cay in to conduct research on Dominica. We are especially grateful to Lindon the central Exumas) as an ectoparasite of various species of Prince, who was an invaluable assistant on Dominica. Shedd Aquarium iguanas (Whittick, 1939; Thompson, 1950; Maldonado Capriles and student volunteers provided immense assistance in the field. John G. and Medina Gaud, 1977; Guglielmone et al., 2003; Durden and Shedd Aquarium provided support for the study, as did San Diego Zoo Knapp, 2005). We recorded interesting distributions for A. torrei Global through a Conservation Research Postdoctoral Fellowship. This work was supported in part by NSF grant DEB-REVSYS 1026146. and A. albopictum relative to each other in the Bahamas on the same host species, C. cychlura (Table I). There is a sharp LITERATURE CITED geographical delineation between these 2 tick species in the Bahamas, with A. albopictum parasitizing C. c. cychlura on BREUIL, M., M. DAY, AND C. KNAPP. 2010. Iguana delicatissima. In IUCN Red List of Threatened Species. Version 2010.4. Available at: www. Andros Island and C. c. figginsi in the central Exumas. iucnredlist.org. Accessed 1 October 2014. Conversely, the geographical distribution of A. torrei is restricted BUSH, A. O., K. D. LAFFERTY,J.M.LOTZ, AND A. W. SHOSTAK. 1997. to C. c. figginsi iguana populations in the southern Exumas (Table Parasitology meets ecology on its own terms: Margolis et al. revisited. I). 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