Caribbean Journal of Science, Vol. 39, No.

3, 321-326, 2003
Copyright 2003 College of Arts and Sciences
University of Puerto Rico, Mayaguez

Hemidactylus mabouia (Sauria: Gekkonidae), Host of Geckobia

hemidactyli (Actinedida: Pterygosomatidae), throughout the
Caribbean and South America
CARLOS CE SAR MARTINEZ RIVERA1,3, ALBERTO GONZA LEZ NEGRO N1, MICHEL BERTRAND2,
1
AND JAIME ACOSTA
1

Departamento de Biologa, Universidad de Puerto Rico, Recinto de Mayaguez, Mayaguez, Puerto Rico 00680 and
2
Laboratoire de Zoogeographie, Universite Montpellier III, route de Mende, 34199 Montpellier Cedex 5, France
3
Present Address/Corresponding Author: Division of Biological Sciences, 105 Tucker Hall, University of Missouri,
Columbia, MO 65211, USA ccmwvf@mizzou.edu
ABSTRACT.Hemidactylus mabouia (Sauria: Gekkonidae) from Puerto Rico often harbor tiny orange colored mites which we identified as Geckobia hemidactyli (Actinedida: Pterygosomatidae). We also found this
mite on museum specimens of H. mabouia from South America and the Caribbean. Geckobia is a widespread
genus of reptilian ectoparasites, found mostly on the Eastern Hemisphere. Geckobia hemidactyli has been
reported from southern Africa on H. mabouia and H. tasmani and from the Mediterranean and Asia in
H. frenatus and H. mercatorius. Other species of Geckobia have been found on Tarentola americana and in
American geckos of the genus Platydactylus (possibly Thecadactylus) and Phyllodactylus. This is the first
report of G. hemidactyli, from the western hemisphere and of any pterygosomatid mite for Puerto Rico and
its satellite islands.

INTRODUCTION
Many studies have been conducted on
the gekkonid lizard Hemidactylus mabouia in
South America and the Caribbean. Most are
related to human-mediated dispersal, ecology, natural history, and cytology (Powell
et al. 1998), but research on parasites is limited. Previous research on the internal and
external parasites of H. mabouia from
Puerto Rico has revealed only one parasite,
a pentastomid, Raillietiella hemidactyli (Self
and Garca Daz 1961). Dyer et al. (1999)
identified other parasites from reptiles in
Puerto Rico, but none in H. mabouia while
Goldberg and Bursey (2000) found two
nematodes in H. mabouia from Antigua, and
Baker (1987) found a diversity of nematodes on H. mabouia. Research on the external parasites of reptiles in South America
and the Caribbean have revealed Geckobia,
an African pterygosomatid mite, on neotropical geckos, including G. leonilae, from
Phyllodactylus lanei rupius (Hoffmann and
Morales Malacara 1985); G. guyanensis and
G. manaensis from a Platydactylus (possibly
Thecadactylus, Floch and Abonnenc, 1944,

1945); G. cayennensis from Phyllodactylus

sp. (Floch and Fauran 1955); G. tarentolae
from Tarentola americana (De la Cruz 1973)
and an unidentified Geckobia from Thecadactylus rapicauda (Martnez Rivera unpublished). Although there are no formal published reports of G. hemidactyli from South
America or the Caribbean, various workers
have seen this mite on H. mabouia (W. E.
Meshaka Jr; J. A. Rivero pers. comms.) and
published photographs of neotropical H.
mabouia clearly show Geckobia (see Malhotra and Thorpe 1999; Rivero 1978). Thus
we addressed the following questions, did
Geckobia follow H. mabouia in its geographic
expansion throughout the Americas? And
if so, how wide is its range? Would it be an
effective tool to answer biogeographic
questions about the colonization of new
tropical regions by H. mabouia?
Pterygosomatid mites, such as Geckobia,
are commonly found on geckos and other
lizards (Bauer et al. 1990). Various Geckobia
are known to occur on geckos from the Paleotropics, including other hemidactyls,
(Tarentola mauritanica Girot 1969), diplodactylines (Naultinus and Hoplodactylus Robb

321

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C. C. MARTINEZ-RIVERA ET AL.

1986) and gekkonines, (Gehyra oceanica,

Lepidodactylus euaensis Bertrand and Ineich
1986, Bertrand 1987). Other Geckobia have
been found in Hemidactylus garnotii (Oliver
and Shaw 1953), Hemidactylus sp. in the pet
trade (pers. obs.), and tortoises (Geckobia
enigmatica from Geochelone yniphora; Bertrand and Perdon o 2000).
Pterygosomatid mites were reviewed by
Lawrence (1935, 1936) and revised by
Womersley (1941) and Radford (1943).
Lawrence (1936) diagnosed Geckobia as having a dorsal scutum, short eyes, mouth
parts almost wholly exposed on the anterior dorsal surface of the body, and coxae
armed with spurs (also see ReichenbachKline and Elkan 1965). G. hemidactyli is
identified by its orange coloration, an inconspicuous dorsal shield and dorsum covered with many short hair (setae), eyes are
inconspicuous, situated laterally. Males
and nymphs have a rounder body and
fewer setae than females. See Lawrence
(1936) for description of females and Jack
(1961) for descriptions of males and
nymphs.
Mites of the genus Geckobia, although
mostly parthenogenic, are sexually dimorphic with males being rare and considered
neotenic deutonymphs (2nd stage larvae).
The sexes exhibit distinct behavior and congregate at various locations at different levels of maturity (Bauer et al. 1990). Mobility
is reduced in various life cycles; protonymphs (1st stage larvae), tritonymphs
(3rd stage larvae) and adults are calyptostasic (quiescent stage), while deutonymphs
(2nd stage larvae) are the only active stage
(Girot 1969).
MATERIALS AND METHODS
Field study.We collected thirty-one
Hemidactylus mabouia and three Hemidactylus haitianus {H. brookii haitianus, Powell et
al. 1996} at night during regular foraging
activities in or near buildings in Puerto
Rico, Culebra, and Isla de Mona. The thirtyone H. mabouia were collected as follows:
twenty-two from mainland Puerto Rico
(West: An asco, Cabo Rojo and Mayagu ez;
South: Guayanillas and Ponce; North/

Central: Naranjito; North: Arecibo and

East: Humacao); four from the park office
at Playa Flamingo, Culebra, five from the
kitchen building in Playa Sardinera, Isla de
Mona. The three H. haitianus were collected
on western Puerto Rico (An asco and Mayagu ez).
We identified the geckos to species,
sexed them, examined and measured them
to the nearest millimeter and assigned them
to age categories based on snout to vent
length (SVL) adult >50 mm, sub-adults 40
mm to 50 mm, or juvenile/neonate <40
mm. All geckos from Culebra and Isla de
Mona and most geckos from Puerto Rico
were released after examination. We kept
12 H. mabouia from mainland Puerto Rico
and removed mites using tweezers and alcohol swabs; in the latter case we rinsed
geckos with distilled water afterwards. We
housed geckos in trios in 37.85 L glass enclosures and fed them small crickets (Acheta
domesticus) ad libitum every other day. All
geckos were released at their capture site
and mites were deposited at the University
of Puerto Rico, Mayagu ez, for further
study. We identified the mites using the descriptions in Lawrence (1936), Jack (1961)
and Reichenbach-Kline and Elkan (1965).
Collection revision.We examined containers of H. mabouia in the collections of
the University of Puerto Rico, Mayagu ez
(UPR-M) and the Natural History Museum
and Biodiversity Research Center at the
University of Kansas, Lawrence (UK-L).
Three 10 ml samples from the bottom of
each container were withdrawn and placed
on a Petri dish for observation under a dissection microscope. When present, mites
were identified, using the aforementioned
description, placed in a smaller vial and returned to the specimen container. For a list
of the specimens see Appendix 1 (UPR-M)
and 2 (UK-L).
RESULTS
Field results.All adult and sub-adult H.
mabouia had mites, but no juveniles or neonates harbored mites. The number of mites
ranged from three to >300, with both extremes from Mayagu ez. Mites were at-

HEMIDACTYLUS MABOUIA HOST OF GECKOBIA

tached anywhere from the rostrum to the

caudal third of the tail, with higher concentrations in the axilla and groin. Mites on the
ventral surface were larger (up to 0.5 mm
long at the widest part of their body) and
attached beneath the scales. Mites on the
dorsum were usually smaller (ca. 0.3 mm)
and rounder, often moved freely, and were
not firmly attached to the skin. Interestingly, mites remain in place when a gecko
sheds (pers. obs.). The three H. haitianus
were free of mites, even when captured in
syntopy with H. mabouia.
Laboratory results.We found Geckobia
hemidactyli in containers of H. mabouia from
Culebra and Isla de Mona, but not from
Isabela, Puerto Rico (Appendix 1). We
found mites in containers of H. mabouia
from various localities in South America
and the Caribbean: Antigua, Brazil, Colombia, Cuba, Culebra (P.R.), Dominica,
Grenada, Guadeloupe, Isla de Mona (P.R.),
St Croix (B.V.I.), St. Eustatuius, St Lucia,
Tortola (B.V.I.), Vieques (P.R.), the US Virgin Islands (St John, St. Thomas, and other
localities) and one African locality Uganda
(Appendix 2). However mites were absent
from the containers of specimens collected
from Brazil, Martinique, Montserrat, Peru ,
Saba, Saint Lucia, USA (Florida), Kenya,
Liberia, South Africa, Swaziland, Tanzania
and Zambia (African specimens observed
for comparison).
DISCUSSION
In this study we present biogeographic
evidence suggesting that G. hemidactyli is a
common parasite of H. mabouia from Puerto
Rico and most of its range in South
America and the Caribbean. We suggest
that H. mabouia arrived together with Geckobia hemidactyli in Trans-Atlantic dispersal
incidents. Because we observed G. hemidactyli only on sub-adult and adult H. mabouia,
we suggest that non-juvenile geckos must
have been present in the colonization
events of H. mabouia in Puerto Rico and localities where mites are present. However
we do not rule out colonization events by
juveniles, or arrival in the form of egg
masses as suggested by Kluge (1969), or

323

colonization from mite-free gecko populations in Africa or other localities in the New
World. These findings also serve as evidence for the importance of ectoparasites as
a tool for biogeographic and ecological
studies.
To think that G. hemidactyli can freely disperse itself from one individual to the next
at any given time, or that geckos can acquire adult mites from the substrate seems
unlikely. Geckobia shows no adaptation for
jumping or clinging onto passing hosts
(Bertrand 2001), especially at maturity. In
an experimental study of Geckobia, Girot
(1969) observed that larvae search for a favorable attachment zone, becoming fixed to
their host at this stage. Motility largely decreased at the nymph and adult stages,
where individuals loosing their host were
unable to find it again.
We propose that transmission through
sexual contact or other prolonged and direct physical interaction (i.e. fighting, diurnal retreats, communal nesting) is the most
likely way for a host to acquire Geckobia.
We base our argument on the evidence that
mites were present only in adult and subadult H. mabouia and adult mites were particularly abundant in body parts that come
in contact during mating (ventral surface,
axilla, groin and tail). Krysko et. al. (in
press) and Rivero (1978) found that individuals of H. mabouia share nesting cavities
in the lower Florida Keys and Puerto Rico
respectively, and Kearny et al. (2001) found
heterospecific aggregation of mature geckos in diurnal retreats. Accordingly, no
neonates or juveniles had mites, and none
were either captured or observed close to
older conspecifics. Juvenile geckos of various species seldom forage or rest in the
same areas as adults since they risk physical aggression and cannibalism (Hanley et
al. 1998; Kearney et al. 2001). Howard et al.
(2001) found that juveniles of H. mabouia on
Anguilla forage closer to the ground than
adults, presumably for similar reasons.
Not all the containers of H. mabouia had
G. hemidactyli, this may be attributed to
many factors. First, some individuals were
collected in the early 1950s and 1960s and
have been used for various studies, while

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C. C. MARTINEZ-RIVERA ET AL.

others show evidence of invasive research

and are anatomically incomplete (e.g. open
stomach, missing body parts and skin). Second, some geckos from the same container
and geopolitical region have been placed in
a single container. Regrouping specimens
from various containers, requires the elimination of the preserving solution along
with any mite present on it. Third, Caribbean and South American specimens
without mites were from containers with
clear alcohol and little debris, mites were
almost exclusively found on containers
with visible debris at bottom suggesting
that mites can be flushed everytime the preserving solution is replaced.
Research is being conducted in Puerto
Rico to establish the distribution of G. hemidactyli and H. mabouia around Puerto Rico
and its adjacent islands and find the genetic
relatedness of various populations. Future
studies might look at possible interactions
and differences (for example, ecological,
genetic, morphological and biogeographical) between mites from the Western and
Eastern hemispheres in order to compare
the ecological and phylogeneic relationships of G. hemidactyli and their host.
Acknowledgments.We thank Amandy
Carranza for involving us with the geckos,
A. Berrios, I. Sastre, D. Segu, L. Williams,
J. A. Rivero, and F. Bird, for counseling,
G. Breckon and D. Kolterman provided
laboratory space and equipment (UPR lvarez,
Mayagu ez), M. A. Garca, A. O. A
and G. Pons helped in Isla de Mona (P. R.
DRNA). R. Cocroft, G. Ho bel, Y. Jang, and
V. Marshall, (UM-Columbia, MO) commented on the manuscript. We thank R.
Powell (Avilla College, Kansas City, MO),
W. E. Meshaka, Jr. (State Museum of PA),
and K. Krysko (FL Museum of Nat. Hist.)
for their revisions and comments. J. Guzma n, H. Martnez, and O. Monsegur (UPRMayagu ez) helped collect specimens. We
thank J. M. Guayasamn, H. Alamillo, and
L. Trueb for the loan from KU-Lawrence.
The manuscript was written at the laboratory of H. C. Gerhardt (UM-C) by CCMR
under a GAANN Scholarship from the Department of Education.

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Appendix 1
Hemidactylus mabouia from University of
Puerto Rico, Mayagu ez Campus: Collected
by the class of Herpetology 1972; Isabela,
Puerto Rico; Playa Flamingo, Culebra; and
Playa Sardinera, Isla de Mona.
Appendix 2
Hemidactylus mabouia from the Natural
History Museum and Biodiversity Research Center, University of Kansas,
Lawrence:
South America. Brazil: Ilha Madre de
Deus (29429-29437); Belem (127137-127142),
IPEAN, 3 km E Belem (128130-128168);
Obidos (130191-13095); Manaus (13096).
Colombia: Leticia (124916-124922). Peru :
Tarapoto (212605); Moyobamba (212606212609); Iquitos, Hotel Turistas (220484);
San Jacinto (222139); Teniente Lo pez
(222140). Caribbean. Antigua: Dutchman
Bay (225133), Dutchman Bay (229535229552). Cuba: Baha de Guanta namo
(229515-229534). Dominica: Clark Hill Estate (100529-100534). Grenada: Salt Pond
(229553); St Georges (229554-229564); Point
Saline lighthouse (229565-229566); Tyrrell
Bay (229567); NA (229568-229569). Guadaloupe: Grande Terre, Grosier (229570229571); Basse Terre, 2 km SE Vieux Habitants, Grande-Terre, 1 km W Pointe des
Chateaux (229573-229596); Terre de Bas
(229597); Ilet a Cabrit, Fort Josephine
(229598); Terre-de-bas, Pointe a Vache
(229599), Terre-de Haut, southern end
(229600-229601). Montserrat: Carrs Bay
(229606-229608). Puerto Rico: Culebra
(180566); Vieques, Cayo de Tierra, eastern
most of two (229636-229639), (229666); Isla
de Mona, Anclaje Sardinera (229667229669). Saba: The Bottom (229609-229612).
St Croix: Christiansted (229640), (229642229644), (229647-229648); Cape forming
western side of Prune Bay (229641). St Eustatius: Ornajestad (229613-229626). St

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C. C. MARTINEZ-RIVERA ET AL.

John: Caneel Bay (229649); Lameshur

Range Station (45675), (46678-46683); 1/5
mi S & 2/5 mi W Lameshur (45685); 1/5 mi
WSW Lameshur (46684-446686); NA
(94418). St. Lucia: Castries (229627-229633);
Reduit (229634). St. Thomas: Water Bay
(229650-229651); S side of Lovango Cay
(45632); 0.2 mi SW Lameshur (45673); 0.5
mi SW Lameshur (45674); 0.1 mi SW Lameshur (45678-45682), (45684). United States:
Key Largo (220254); 2 mi S Miami International Airport (221732); Fort Myers
(222285); Florida Bay, Flamingo (223405223408); Long Pine Key, Dan Beard Re-

search Center (223408). U.S. Virgin Islands:

Reef Bay, Estate House (45676-77), Reef Bay
Estate House (45683), Annaberg (4568645687). Tortola: 1 mi W Pasea Hall Estate
(229508-229510); Lloyds Pond (229511229514). Africa.
Kenya: Tsavo National Park (172933);
Meru, Samburu Lodge, Ewaso Ngiro River
(172934-172936). Liberia: Sanniquelle
(63595). South Africa: St. Luca (195945195946). Swaziland: 40 km NW Big Bend
(196099-196100). Tanzania: Minaki, St Andrews College (97101-97113). Uganda: NA
(97114). Zambia: Livingstone (175558).