Transport of Helminths to Hawaii via the Brown Anole, Anolis sagrei

Transport of Helminths to Hawaii via the Brown Anole, Anolis sagrei (Polychrotidae)Author(s): Stephen R. Goldberg and Charles R. BurseySource: The Journal of Parasitology, Vol. 86, No. 4 (Aug., 2000), pp. 750-755Published by: The American Society of ParasitologistsStable URL: http://www.jstor.org/stable/3284959 .

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J. Parasitol., 86(4), 2000, p. 750-755 ? American Society of Parasitologists 2000

TRANSPORT OF HELMINTHS TO HAWAII VIA THE BROWN ANOLE, ANOLIS SAGREI (POLYCHROTIDAE)

Stephen R. Goldberg and Charles R. Bursey* Department of Biology, Whittier College, Whittier, California 90608

ABSTRACT: Sixty-two brown anoles, Anolis sagrei, from Oahu, Hawaii were examined for helminths. Anolis sagrei was introduced to Hawaii, presumably from the Caribbean. Two species of trematodes, Mesocoelium monas and Platynosomum fastosum, 3 species of nematodes, Atractis scelopori, Physaloptera squamatae, and Physocephalus sp., 1 acanthocephalan, Acanthocephalus bufonis, and 1 pentastome, Raillietiellafrenatus, were found. Atractis scelopori and P. squamatae, previously unknown in Hawaii, are widely distributed in the Caribbean and were most likely transported to Hawaii with the introduced anoles. Mesocoelium monas, P. fastosum, Physocephalus sp., A. bufonis, and R. frenatus have been previously reported from Hawaiian herptiles; A. sagrei most likely acquired infections of these parasites from Hawaiian populations. This study indicates that helminths can be transported with their introduced hosts and become established in the colonized areas and that introduced lizards may quickly acquire species of previously established helminthes.

Introduced lizards have received little attention with regard to the species of helminths harbored. Of the 137 species of lizards in the continental United States and Canada, 32 have been introduced (Powell et al., 1998). Of these, helminth lists exist for 3: the bark anole Anolis distichus, the brown anole Anolis sagrei, and the Mediterranean gecko Hemidactylus turcicus, (Pence and Selcer, 1988; McAllister et al., 1993; Gold- berg et al., 1994). Of 4 lizard species in Bermuda, 3 were introduced (Wingate, 1965), the panther anole Anolis bimaculatus, Graham's anole Anolis grahami, and savannah anole Anolis roquet; these 3 species have been examined for helminths (Goldberg, Bursey, and Tawil, 1995). All 18 lizard species in Hawaii have been introduced (McKeown, 1996). Four, the green anole Anolis carolinensis, the house gecko Hemidactylus frenatus, the Indo-Pacific gecko Hemidactylus garnotii, and the mourning gecko Lepidodactylus lugubris, have been examined for helminths (Loo, 1971; Brown et al., 1995; Bursey and Gold- berg, 1996a; Hanley et al., 1998). These helminth lists raise an intriguing question. Are these helminths acquired after introduction or are they transported with the host and establish new populations?

Anolis sagrei Dum6ril and Bibron, 1837 is native to Cuba, Jamaica, and the Bahamas and has been introduced to Florida, Grand Cayman Island, and Hawaii (Schwartz and Henderson, 1991; McKeown, 1996). The origin of the Hawaiian (Oahu) population is unknown, but it is believed that these anoles are descendents of released pets; they were first noticed as part of the herpetofauna in 1980 (McKeown, 1996). On Oahu, Hawaii, A. sagrei is found in urban areas where it is sympatric with the cane toad Bufo marinus, A. carolinensis, H. frenatus, L. lugubris, and the metallic skink Lampropholis delicata. Helminths of native A. sagrei have been reported, i.e., Cuba (Coy Otero, 1970; Coy Otero and Baru', 1973, 1979), Jamaica (Bundy et al., 1987), and the Bahamas (Goldberg et al., 1994), as well as from introduced populations, e.g., Florida (Price and Under- wood, 1984; Sellers and Graham, 1987; Goldberg et al., 1994) and Grand Cayman Island (Goldberg, Bursey, and Cheam, 1995).

The purpose of the present study was to examine A. sagrei

from Hawaii to compare helminths of this introduced population with helminths harbored by native conspecific anoles from Cuba, Jamaica, and the Bahamas and introduced conspecific anoles from Florida and Grand Cayman Island.

MATERIALS AND METHODS

Sixty-two A. sagrei (mean snout-vent length [SVL] = 50.1 mm _ 9.4 SD, range = 27-66 mm) were collected by handheld noose Feb-

ruary 1999 at Kailua (21?25'N, 157045'W) Koolaupoko District, Oahu, Hawaii. The anoles were killed within 4 hr of capture by an injection of sodium pentobarbital, fixed in 10% formalin, and preserved in 70% ethanol. The body cavity was opened by a longitudinal incision from vent to throat and the gastrointestinal tract was removed by cutting across the esophagus and rectum. The esophagus, stomach, small intestine, large intestine, lungs, liver, gall bladder, and body cavity were examined separately for helminths. Nematodes were placed in undiluted glycerol, allowed to clear, and examined under a light microscope. Trematodes were stained in hematoxylin, mounted in Canada balsam, and identified. Voucher specimens were sent to the U.S. National Par- asite Collection (USNPC), Beltsville, Maryland: Mesocoelium monas, 89067; Platynosomum fastosum, 89068; Atractis scelopori, 89069; Phy- saloptera squamatae, 89070; Physocephalus sp. (encysted larvae), 89071; Acanthocephalus bufonis, 89072; Raillietiella frenatus, 89073. Anoles were deposited in the herpetology collection of the Natural His- tory Museum of Los Angeles County (LACM nos. 145346-145388, 145460-145478).

RESULTS

Fifty-five anoles (89%) harbored helminths. Two species of Trematoda, M. monas (Rudolphi, 1819) and P. fastosum Kos- sack, 1910, 3 species of Nematoda, A. scelopori (Gedoelst, 1919), P. squamatae Harwood, 1932, and Physocephalus sp., 1 species of Acanthocephala, A. bufonis (Shipley, 1903), and 1 species of Pentastomida, R. frenatus Ali, Riley, and Self, 1981, were found. Prevalence, mean intensity, range, and site of infection for these helminths are given in Table I.

DISCUSSION

General observations

All helminths found in the present study have previously been reported from other hosts. Of these helminths, M. monas, P. fastosum, and Physocephalus sp. have a somewhat cosmopolitan distribution (Alicata, 1935; Loo, 1971; Nasir and Diaz, 1971). Atractis scelopori and P. squamatae are known previously only from the western hemisphere (Baker, 1987). Acan-

Receieved 15 November 1999; revised 20 January 2000; accepted 20 January 2000.

* Department of Biology, Pennsylvania State University, Shenango Campus, Sharon, Pennsylvania 16146.

750



GOLDBERG AND BURSEY-TRANSPORT OF HELMINTHS TO HAWAII 751

TABLE I. Site of infection, prevalence, mean intensity, and range of infection of 7 helminth species in 62 Anolis sagrei.

Prevalence Helminth species Site of infection (%) Mean intensity + SD Range

Trematoda

Mesocoelium monas Small, large intestine 3 1.0 Platynosomum fastosum (larvae) Gall bladder 2 9.0

Nematoda

Atractis scelopori Large intestine 39 179.3 + 108.9 45-425 Physaloptera squamatae Stomach 61 18.0 + 322.5 1-120 Physocephalus sp. (larvae) Stomach wall (cysts) 50 9.6 + 13.1 1-51

Acanthocephala

Acanthocephalus bufonis Small intestine 2 1.0

Pentastomida

Raillietiella frenatus Lung 8 2.2 ? 2.7 1-7

thocephalis bufonis and R. frenatus have an oriental distribution (Ali et al., 1981; Barton and Pichelin, 1999). A helminth list for native A. sagrei is given in Table II. A helminth list for introduced A. sagrei is given in Table III. Known helminths of lizards from Hawaii are listed in Table IV. However, Loo's (1971) observation that helminth parasites resembling tape- worms, roundworms, and lungworms are found in Hawaiian geckos is not recorded in Table IV. Some comments may be made about the helminth species harbored by A. sagrei on Ha- waii from these distribution patterns.

Cosmopolitan species Mesocoelium monas is a parasite of the intestine of amphib-

ians, mainly anurans, and reptiles in tropical and subtropical regions (Prudhoe and Bray, 1982). Mesocoelium monas (=Me- socoelium incognitum Travassos, 1921) was first reported in B. marinus from Hawaii by Yuen (1965). Bufo marinus was introduced into Hawaii from Puerto Rico in 1932 (McKeown, 1996). Because native populations of A. sagrei are not known to harbor M. monas (Table II) but populations of introduced A. sagrei are (Table III), and both A. sagrei and B. marinus are sympatric on Oahu, Hawaii, it seems reasonable to assume that A. sagrei acquired this helminth in Hawaii.

Platynosomum fastosum, the cat liver fluke, requires 3 intermediate hosts to complete its life cycle: mollusk, isopod, and lizard (Maldonado, 1945). It has been found in Puerto Rico, Brazil, Cuba, United States, and Africa (Yamaguti, 1958) and was first reported in Hawaii by Ash (1962). In Hawaii, lizard intermediate hosts were determined by Loo (1971) to be A. carolinensis, H. frenatus, and H. garnotii. Eckerlin and Leigh (1962) reported A. sagrei from south Florida as a host for P. fastosum (Table III). Anolis sagrei is the fourth Hawaiian lizard species known to harbor P. fastosum.

Physocephalus sp. was found only as encapsulated larvae, and in some cases sclerification of the cysts had begun. Adults of Physocephalus are known to occur in the stomachs of suids, equines, bovines, and lagomorphs; infective larvae have been recovered from 20 genera of dung beetles and encapsulated infective larvae are commonly found in the tissues of amphibians, reptiles, birds, and mammals that have ingested infected beetles (Anderson, 1992). Alicata (1964) reported that in Ha-

waii feral pigs are the primary host for Physocephalus sexalatus. Given the life cycle of P. sexalatus, it is reasonable to assume that infection of A. sagrei in Hawaii is a byproduct of the anole's diet, and infection could be expected in any insec- tivorous Hawaiian lizard. Hanley et al. (1998) reported encapsulated larvae of Physocephelus sp. from H. frenatus of Hawaii (Table IV). Brown et al. (1995) found encapsulated nematodes in stomach walls of H. frenatus and L. lugubris but did not attempt identification. Although we have not seen the specimens, we believe there is a high probability that they are Phy- socephalus sp. and have assigned them to this genus in Table IV.

Western hemisphere species Atractis scelopori was originally described by Gedoelst

(1919) from specimens found in a preserved lizard, Sceloporus undulatus, collected in North America (no precise location was given) and deposited in the Natural History Museum of Brus- sels, Belgium. Since then, A. scelopori has been reported from about 50 species of lizards including A. sagrei from the Carib- bean, southern United States, Mexico, and Panama (Baker, 1987). The life cycle of A. scelopori has not been studied; however, eggs of atractids hatch and larvae develop to third stage in utero, thus autoinfection is the rule (Anderson, 1992). Be- cause A. scelopori is known only from the Caribbean and ad- jacent area, it is reasonable to assume transport of this helminth to Hawaii by A. sagrei and continued infection of this host. Hawaii is a new geographical record for A. scelopori.

Physaloptera squamatae was originally described from specimens taken from Scincella lateralis (=Leiolopisma laterale) and Agkistrodon piscivorus (=Agkistrodon mokasen) by Har- wood (1932) from Houston, Texas. It has subsequently been reported from A. sagrei as well as 1 species of lizard from the southern United States and 12 additional species of Caribbean lizards (Baker, 1987). Physalopterans utilize insect intermediate hosts (Anderson, 1992). Because P. squamatae is known only from the western hemisphere, it is reasonable to assume transport of this helmninth to Hawaii by A. sagrei. Hawaii is a new geographical record for P. squamatae. Physalopterid larvae have been reported from Hawaii in H. frenatus (Table IV); however, no species determination was attempted. Physaloptera



752 THE JOURNAL OF PARASITOLOGY, VOL. 86, NO. 4, AUGUST 2000

TABLE II. Helminths of native A. sagrei.

Number anoles infected/number

Helminth Locality anoles examined Reference

Trematoda

Urotrema scabridum Bahamas 3/45 Goldberg et al., 1994 Cuba 5/21 Coy Otero, 1970 Jamaica 2/12 Bundy et al., 1987

Urotrema wardi Cuba 2/21 Coy Otero, 1970

Nematoda

Abbreviata sp. (larvae) Cuba 12/138 Coy Otero and Barug, 1979 Atractis opeatura Cuba 2/138 Coy Otero and Barug, 1979 A. scelopori Bahamas 23/45 Goldberg et al., 1994

Cuba 4/21 Barug and Coy Otero, 1969 Cuba 4/21 Coy Otero, 1970 Cuba 4/19 Coy Otero and Barug, 1973 Cuba 33/138 Coy Otero and Barug, 1979 Jamaica 10/12 Bundy et al., 1987

Oswaldocruzia lenteixeirai Bahamas 3/45 Goldberg et al., 1994 Cuba 3/138 Coy Otero and Barug, 1979

Ozolaimus monhystera Cuba 2/138 Coy Otero and Barug, 1979

Parapharyngodon cubensis Bahamas 1/45 Goldberg et al., 1994 Cuba 1/11 Barug and Coy Otero, 1969 Cuba 1/21 Coy Otero, 1970 Cuba 2/138 Coy Otero and Barug, 1979 Jamaica 1/12 Bundy et al., 1987

P. squamatae Bahamas 20/45 Goldberg et al., 1994 Cuba 3/138 Coy Otero and Barug, 1979

Physalopteridae gen. sp. Cuba 1/138 Coy Otero and Barug, 1979 Porrocaecum sp. (larvae) Bahamas 1/45 Goldberg et al., 1994

Cuba 5/138 Coy Otero and Barug, 1979

Skrjabinoptera phrynosoma Cuba 7/21 Barug and Coy Otero, 1969 Cuba 7/21 Coy Otero, 1979 Cuba 55/138 Coy Otero and Barug, 1979

Strongyloides sp. Bahamas 1/45 Goldberg et al., 1994

Trichospirura teixeirai Cuba 13/138 Coy Otero and Barug, 1979

Acanthocephala

Centrorhynchus sp. Bahamas 1/12 Goldberg et al., 1994 Jamaica 2/12 Bundy et al., 1987

muris-brasiliensis Diesing, 1861, a parasite of rats, and Phy- saloptera praeputialis Linstow, 1899, a parasite of cats, have been reported from Hawaii (Alicata, 1964).

Far East species

Acanthocephalus bufonis has been reported from 5 species of amphibians and 1 species of lizard (Kennedy, 1982; Barton and Pichelin, 1999). The life cycle of A. bufonis is unknown. Natural populations of A. sagrei (Table II) are known to harbor

cystacanths, but adult acanthocephalans have not been reported. Barton and Pichelin (1999) reported Hawaiian populations of B. marinus to harbor A. bufonis; no other hosts were listed. Anolis sagrei is a new host record for A. bufonis.

Rallietiella frenatus was originally described from H. frenatus from Malaysia by Ali et al. (1981), who also reported it in lizards from the Philippine Islands, Taiwan, South Vietnam, and Thailand. Hanley et al. (1998) reported R. frenatus from Hawaii to occur in H. frenatus and L. lugubris. Brown et al. (1995) also reported pentastomid infections from Hawaiian

populations of H. frenatus and L. lugubris but did not identify the species. In Table IV, based upon the revision of Raillietiella

by Ali et al. (1981) in which R. frenatus is described and Rail- lietiella hemidactyli is limited to the Indian subcontinent and, even though we were unable to examine them, we have as-

signed the pentastomids of Brown et al. (1995) to R. frenatus. The life cycle of pentastomids involves 2 hosts; infection occurs with ingestion of infected intermediate hosts (Ali and Riley, 1983). Anolis sagrei is a new host record.

Conclusions

The answer to our original question must be that parasite lists for introduced hosts may contain parasites transported by the introduced host as well as parasites acquired by the host after introduction. This is especially true for Hawaiian lizards, all of which have been introduced. Of the 7 helminth species harbored by A. sagrei, 5 species, M. monas, P. fastosum, Physo- cephalus sp., A. bufonis, and R. frenatus, were previously known to infect Hawaiian herptiles. These parasite species most




TABLE III. Helminths of introduced A. sagrei.

Number anoles infected/number

Helminth Locality anoles examined Reference

Trematoda

P. fastosum (larvae) Florida Not stated Eckerlin and Leigh, 1962 Hawaii 1/62 This study

M. monas Florida 10/100 Price and Underwood, 1984 Florida 2/82 Sellers and Graham, 1987 Hawaii 2/62 This study

Urotrema scabridum Florida 7/82 Sellers and Graham, 1987 Florida 4/25 Goldberg et al., 1994

Nematoda

Abbreviata sp. (larvae) Cayman Islands 1/10 Goldberg, Bursey, and Cheam, 1995 Atractis penneri Florida 22/25 Goldberg et al., 1994 A. scelopori Cayman Islands 10/10 Goldberg, Bursey, and Cheam, 1995

Florida 36/100 Price and Underwood, 1984 Hawaii 24/62 This study

P. squamatae Cayman Islands 3/10 Goldberg, Bursey, and Cheam, 1995 Florida 50/100 Price and Underwood, 1984 Florida 15/25 Goldberg et al., 1994 Hawaii 38/62 This study

Physocephalus sp. (larvae) Hawaii 31/62 This study

Acanthocephala A. bufonis Hawaii 1/62 This study

Pentastomida

R. frenatus Hawaii 5/62 This study

TABLE IV. Helminths infecting Hawaiian lizards.

Host

Anolis Hemidactylus Hemidactylus Lepidodactylus Helminth Anolis sagrei carolinensis frenatus garnotii lugubris

M. monas X* P. fastosum (metacercaria) X* Xt Xt Xt Cylindrotaenia allisonae X# Cylindrotaenia sp. Xt X Oochoristica sp.

Xt,? Xt

A. scelopori X* Gongylonema sp. (larvae) X$ Parapharyngodon sp. Xt Pharyngodon lepidodactylus -- X

,#, P. squamatae X* Physaloptera sp. (larvae) X$ Physocephalus sp. (larvae) X*

Xt,? X? X? Spauligodon hemidactylus Xtll Skrjabinelazia machidai X Xt,# Unidentified oxyurid nematodes X? X? A. bufonis X* R. frenatus X* Xt ? X,?,# * This study. t Loo, 1971.

$ Hanley et al., 1998. ? Brown et al., 1995. 11 Bursey and Goldberg, 1996a. # Goldberg and Bursey, 1997. 1? Bursey and Goldberg, 1996b.



754 THE JOURNAL OF PARASITOLOGY, VOL. 86, NO. 4, AUGUST 2000

likely infected A. sagrei after its introduction to Hawaii and have remained in the introduced populations at low prevalence. Two species, A. scelopori and P. squamatae, reported from Ha- waii for the first time, most likely entered Hawaii with introduced A. sagrei and have since become well established (as indicated by high prevalence) in the Hawaiian population of A. sagrei. Examination of sympatric lizard species should be made to determine if these parasites have infected other hosts.

It is becoming evident that lizards are infected by generalist helminths (helminths capable of completing their life cycles in many hosts). Introduced lizards may acquire previously established helminths and under favorable circumstances transport their helminths to colonized areas as seen in this study. A con- sequence of infection by generalist helminths is that lizard species lack a unique helminth fauna. Thus, ecological conditions favoring egg survival for monoxenous helminths and the distribution of intermediate hosts for heteroxenous helminths are much more important than the presence of a particular lizard species.

ACKNOWLEDGMENT

We thank Cheryl Wong for assistance with dissections.

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Transport of Helminths to Hawaii via the Brown Anole, Anolis sagrei