Biota Neotropica ISSN: 1676-0611 [email protected] Instituto Virtual da Biodiversidade Brasil

Lima, Vandeberg F.; Brito, Samuel V.; Araujo Filho, João A.; Teles, Diêgo A.; Ribeiro, Samuel C.; M. Teixeira, Adonias A.; A. Pereira, Antonio M.; Almeida, Waltécio O. Helminth parasites of Phyllodactylidae and Gekkonidae lizards in a Caatinga ecological station, northeastern Brazil Biota Neotropica, vol. 17, núm. 4, 2017, pp. 1-7 Instituto Virtual da Biodiversidade Campinas, Brasil

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Helminth parasites of Phyllodactylidae and Gekkonidae lizards in a Caatinga ecological station, northeastern Brazil

Vandeberg F. Lima 1*, Samuel V. Brito 4, João A. Araujo Filho 2, Diêgo A. Teles 2, Samuel C. Ribeiro 3, Adonias A. M. Teixeira 2, Antonio M. A. Pereira 1, Waltécio O. Almeida 1

1Programa de Pós-Graduação em Bioprospecção Molecular, Departamento de Química Biológica, Universidade Regional do Cariri – URCA, Rua Cel. Antônio Luiz, 1161, Campus do Pimenta, 63105-000 Crato, CE, Brazil 2Programa de Pós-Graduação em Ciências Biológicas (Zoologia), Departamento de Sistemática e Ecologia – DSE, Centro de Ciências Exatas e da Natureza – CCEN, Universidade Federal da Paraíba – UFPB, Cidade Universitária, Campus I, 58059-900 João Pessoa, PB, Brazil 3Instituto de Formação de Educadores - IFE, Universidade Federal do Cariri - UFCA, Campus Brejo Santo, 63260-000 Brejo Santo, CE, Brazil 4Centro de Ciências Agrárias e Ambientais, Universidade Federal do Maranhão – UFMA, Boa Vista, 65500-000 Chapadinha, MA, Brazil *Corresponding author: Vandeberg F. Lima, e-mail: [email protected]

 . .  . .   € . . •• ‚. .  • . ƒ. •„• . . . …•• . . . •‚ †. . Helminth parasites of Phyllodactylidae and Gekkonidae lizards in a Caatinga ecological station, northeastern Brazil. iot eotropic.  : e  . http://
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Abstract: We investigated the parasites of ve lizard species belonging to Phyllodactylidae ( Phyllopezus pollicaris n
Gymnodactylus geckoides  n
‡eˆˆoni
e  Hemidactylus agrius  Lygodactylus klugei n
Hemidactylus brasilianus  families in a semiarid region of Brazil. Six nematode species were identied: Parapharyngodon alvarengai n
Spauligodon oxkutzcabiensis (Pharyngodonidae), Physaloptera lutzi (Physalopteridae), Skrjabinelazia intermedia e‰rti
e Trichospirura sp. hb
ochoni
e n
Piratuba sp. nchocerci
e n
 cesto
e species Oochoristica sp. instowii
e. he Šost pre‹lent species were Spauligodon oxkutzcabiensis  which inŒecte
P. pollicaris  • n
Parapharyngodon alvarengai  which inŒecte
G. geckoides (29%). South American lizards were identied as bein new hosts Œor the Trichospirura en‰s  ‰s‰l prsite oŒ ŠŠŠls n
there were  new occ‰rrences oŒ parasite species in the ve lizard species studied herein. Keywords: Squamata, Nematoda, Cestoda, geckos, semiarid

Helmintos parasitas de lagartos Phyllodactylidae e Gekkonidae em estação ecológica na Caatinga, nordeste do Brasil

Resumo: Nós investigamos os parasitas de cinco espécies de lagartos pertencentes às famílias Phyllodactylidae ( Phyllopezus pollicaris e Gymnodactylus geckoides  e ‡eˆˆoni
e  Hemidactylus agrius  Lygodactylus klugei e Hemidactylus brasilianus  eŠ reiŽo seŠi•ri

o rsil. eis esp•cies
e neŠtoi
es ŒorŠ encontr
os: Parapharyngodon alvarengai e Spauligodon oxkutzcabiensis (Pharyngodonidae), Physaloptera lutzi (Physalopteridae), Skrjabinelazia intermedia e‰rti
e Trichospirura sp. hb
ochoni
e Piratuba sp. nchocerci
e e ‰Š esp•cie
e cest‘
eo Oochoristica sp. instowii
e. s esp•cies
e Šiores pre‹l’ncis ŒorŠ S. oxkutzcabiensis   “‰l inŒecto‰ P. pollicaris  • e P. alvarengai  “‰l inŒecto‰ G. geckoides  .•. ‘s
oc‰ŠentŠos no‹o reistro
e hospe
eiro pr lrtos n Š•ric
o ‰l pertencente o ’nero Trichospirura  o “‰l • coŠ‰Š eŠ ŠŠ”Œeros e  no‹s ocorr’ncis
e esp•cies prsits ns cinco esp•cies
e lrtos “‰i est‰

s. Palavras chave: Squamata, Nematoda, Cestoda, geckos, semiárido

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iŠ .. et l.

Introduction families have been conducted by nos et l.   il  il   il et l.   il  il   rito et l.    os et l. €elŠinths re prsites tht inŒect the internl n
eternl orns oŒ   Cazorla & Morales Moreno (2015) n Bezerra et al. (2016) . Šost in‹ertebrte n
‹ertebrte ro‰ps  er‰nson 
on 
Parasites are good indicators of healthy ecosystems, which is essential l o lon o et l. 
  nn et l.   b
il et l.   . to sties o€ conser tion n  inten nce o€ host popl tions rco liese Among this wide diversication of hosts, there is a great diversity of parasites    ), which may reect the anthropic inuence in the environment in South American lizards ( il  il   n seer l sties h e in which they reside (   n et l. b ). Thus, characterization of the recorded variation in parasite richness and diversity ( Bursey & Goldberg p r site popl tion o€ cert in re o€ the ‚ tin bioe is essenti l 
Bursey et al. 2005
nos et l. 
r o ilho et l.  
especially for future studies on species conservation. rito et l.   b
 lino et l. 
os et l.  . The current study characterizes the helminth richness of parasitic species Knowledge of the lizard-associated helminth fauna has increased through and the parameters of parasitic infection (prevalence, mean intensity of rese rch concernin i recors o€ new hosts Bursey & Goldberg 2004
infection and range) in lizards of the Phyllodactylidae and Gekkonidae Bursey et al. 2005
il  il 
cllister et l. 
il et l. families collected at the Aiuaba Ecological Station, northeastern Brazil.   ii escriptions o€ new p r site species rs ey et al. 200 
•ereir et l.  ), and (iii) inuence of biotic and abiotic variables on helminth diversity and abundance ( h rpilo et l. 
rito et l.   b
Material and Methods  lino et l.  . Lizards were collected at the Aiuaba Ecological Station (ESEC Aiuaba), Phyllodactylidae and Gekkonidae are two lizard families of the Gekkota municipality of Aiuaba, state of Ceará, northeastern Brazil (6°36’27”S and clade that are phylogenetically closely related taxa ( ites et l.  . 40°08’00.9”W, 466m asl, datum SIRGAS 2000) ( i re . ‰he s ple re Both families occur in Brazil with, respectively, 13 and six nominal species is within the Caatinga biome, which is characterized by xerophytic plants, ‚ost  ƒrnils  ), and only Gymnodactylus geckoides pi„  shrubs, thorny trees and open areas ( nr e ‹i  . ‰he cli te Phyllopezus periosus ori es  Phyllopezus pollicaris pi„  in this bioe is sei ri hot tropic l with n er e nn l r in€ ll o€ (Phyllodactylidae) , Hemidactylus mabouia ore  e …onnes  n Hemidactylus agrius Vanzolini, 1978 (Gekkonidae) occur within the Caatinga 562.4 mm, average temperature ranging from 24 ºC to 26 ºC and a rainy bioe. Hemidactylus brasilianus  r l  n Lygodactylus klugei season from February to April ( Œ••‚•  . ith rtin  w in †† e‡‡oni e o€ten inh bit the ‚ tin Že concte  n l collections thro h ctie se rchin oer €or bioe ˆitt 
och et l. 
nr e et l.  . ‰hese species s plin e„peitions two in  epteber n •oeber n two in re sit n w it €or ers h e noctrn l h bits e„cept €or the irn l 2015 (February and April) with authorization provided by the “Sistema de L. klugei  n n insectiore iet ˆitt 
‚olli et l. 
och  Autorização e Informação em Biodiversidade” (SISBIO order number 43753 -1). ori es 
esŠit et l. 
os 
ecorer et l. 
The lizards were euthanized with a lidocaine lethal dose. Their snout-vent lbŠerŠe et l. 
• ssos  och 
• ssos et l.  . len th ˆ‹ w s e sre with i it l c liper • .. ‰here €ter ‰he sties reportin helinth p r sitis in oth eric €or both they were labelled, xed with 10% formaldehyde and preserved in 70%

Figure 1. Geographic map featuring the Aiuaba Ecological Station (AIA), state of Ceará, northeastern Brazil. In gray, the Caatinga biome as conceived by Œ•  . http://www.scielo.br/bn http://
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Helminth parasites of lizards in a Caatinga ethyl alcohol. Lizard specimens were deposited in the Herpetological Results ‚ollection o€ the e ion l University of Cariri, municipality of Crato, state of Ceará, Brazil. The hosts were dissected under a stereomicroscope We sampled a total of 355 lizards specimens, distributed in the ve focused and their body cavity, lung, stomach, small intestine and large intestine species of this study. Phyllodactylidae: Phyllopezus pollicaris   speciens was analyzed in search of helminths. The lizard’s gonads were analyzed  les   .  .  n  eles   .  .  to determine their gender and sexual maturity. We considered as mature Gymnodactylus geckoides   speciens € les   .  .  males those with developed testicles and a convoluted epididymis, and as n  eles   .  . . •e‚‚oni e: Hemidactylus  tre €e les those th t h  itello enic €ollicles in their oicts n/ agrius   speciens  les   .  .  n  eles or e s in their o ries.   .  .  Lygodactylus klugei   speciens  les The nematodes were cleared in a Hoyer solution and the cestodes were   .  .  n € eles   .  .  n colored in Carmim and cleared in a Hoyer solution as well for identication Hemidactylus brasilianus   speciens  les     .  n •erh rt † ). The parasites were mounted on temporary slides for 13 females (SVL = 43.6 ± 5.7 mm). All lizards were in full sexual maturity. identication, and their morphology was compared with specimens present ƒe o n si„ neto e species … Parapharyngodon alvarengai †reits in the p r sitolo ic l collection o€ the ‘niersi e e ion l o ‚ riri  Spauligodon oxkutzcabiensis hitwoo   Physaloptera lutzi
 n biblio rphic re erence on the escription o the prsite risto ro • ir‡es ˆ o ri es  Skrjabinelazia intermedia †reits species found that did not have specimens in the collection. Subsequently, € Trichospirura sp. ith ˆ hitwoo   n Piratuba sp. ent they were deposited in the Parasitological Collection of the Regional ˆ †reits  ‰ n one cesto e o the en s Oochoristica Šhe  University of Cariri (Appendix 1). The prevalence and the mean intensity that could not be identied at the species level ( ‹ble . of infection for each endoparasite species were calculated for each lizard Helminth parasites found in this study have different life cycles. While species ccor in to sh et l.   . P. lutzi  Oochoristica sp. Piratuba sp. Trichospirura sp. n S. intermedia

Table 1. Helminth parasites collected in lizards belonging to Phyllodactylidae and Gekkonidae families from the Aiuaba Ecological Station, municipality of Aiuaba, state of Ceará, northeastern Brazil. Number of Infected hosts = (NI), Prevalence = P (%), Mean Intensity of Infection = MII and Range (R). (*) New records of hosts. Parasite Location in host Host (NI) P (%) MII (R) Œ•Ž‹•• Pharyngodonidae Parapharyngodon alvarengai lr e intestine Phyllopezus pollicaris  .‘ . ’ lr e intestine Hemidactylus brasilianus*  .‘   sll intestine lr e intestine Hemidactylus agrius  .‘ . ’ sll intestine lr e intestine Gymnodactylus geckoides  .‘ . ’ Spauligodon oxkcutzcabiensis sll intestine lr e intestine Phyllopezus pollicaris  ‘ .€ € lr e intestine Hemidactylus brasilianus*  .‘ . ’ lr e intestine Hemidactylus agrius*  .‘ . ’ lr e intestine Gymnodactylus geckoides  .€‘   lr e intestine Lygodactylus klugei*  .‘  ’ Physalopteridae Physaloptera lutzi stoch lr e intestine Phyllopezus pollicaris*  .€‘ . ’ stoch Hemidactylus brasilianus*  .‘   stoch Hemidactylus agrius  .‘   stoch lr e intestine Gymnodactylus geckoides  .‘  ’ stoch Lygodactylus klugei  .‘  

Seuratidae Skrjabinelazia intermedia sll intestine lr e intestine Phyllopezus pollicaris*  €.‘   lr e intestine Hemidactylus brasilianus*  .‘ . ’ lr e intestine Hemidactylus agrius*  .‘ . ’ Rhabdochonidae Trichospirura sp. llbl er Phyllopezus pollicaris*  .‘ . ’ llbl er Hemidactylus brasilianus*  .‘ . ’ llbl er Hemidactylus agrius*  .€‘ . ’ llbl er Gymnodactylus geckoides*  .‘ . ’ Onchocercidae Piratuba sp. body cavity Gymnodactylus geckoides*  .€‘  

•‹•• Linstowiidae Oochoristica sp. sll intestine Phyllopezus pollicaris  .‘ . ’ sll intestine Hemidactylus brasilianus*  .‘   sll intestine Hemidactylus agrius*  .‘   http://„.oi.or /./†  •   http://www.scielo.br/bn  Biota Neotrop . : e€€ €

i .†. et l.

Figure 2. Helminths of the gastrointestinal tract of geckos from the Aiuaba Ecological Station (AIA), state of Ceará, northeastern Brazil: (a) Skrjabinelazia intermedia posterior iewƒ b Spauligodon oxkutzcabiensis entire specienƒ c Oochoristica sp. nterior iewƒ  Physaloptera lutzi nterior iewƒ e Piratuba sp. nterior iewƒ  Parapharyngodon alvarengai nterior iewƒ  Trichospirura sp. posterior iew. have heteroxenic life cycles, which requires more than one host to complete Physaloptera lutzi infected all lizards sampled in the current study. its life cycle, P. alvarengai n S. oxkutzcabiensis h“e ono„enic li e Although it had been originally described as a parasite of A. ameiva  cycles, not requiring more than one host ( ”ll en
ƒilc‚e et l.   we recore both P. pollicaris n H. brasilianus s new hosts or this n erson €€€  heritte et l. €€   †i re . species. †ther recors o hosts or P. lutzi were lso reporte in the litertre  •n‚os et l.  ƒ „il et l.    ƒ rito et l. b : Discussion Ameivula abaetensis ‡is ˆoch € ‰rcibric    Cnemidophorus abaetensis  Ameivula littoralis ˆoch •rŠ‚o ‰rcibric € ost  Among the parasite species identied, there were 16 new records  Cnemidophorus littoralis  A. ocellifera  Enyalius bilineatus ‡‹ril of hosts distributed among the ve lizard species sampled in this study. € ibron Œ  Eurolophosaurus nanuzae ˆories   Liolaemus ƒe o n three prsite species in P. pollicaris  o r in H. agrius  two in alticolor rbor    L. ornatus Koslowsky, 1898, L. quilmes •therie G. geckoides  si„ in H. brasilianus n one in L. klugei .  Œ Tropidurus guarani  T. semitaeniatus  T. hispidus  T. itambere Spauligodon oxkutzcabiensis was rst recorded with Thecadactylus ˆories    T. torquatus …ie   H. agrius  L. klugei  G. geckoides  rapicauda (Houttuyn, 1782) (Phyllodactylidae). In South America, Micrablepharus maximiliani ˆeinhrt € Žt•en   n Salvator S. oxkutzcabiensis was recorded in association with the following lizards merianae ‡‹ril € ibron Œ   Tupinambis merianae . •ien •“il et l. €€ •ol ber et l. €€ Goldberg & Bursey 2010 tht P. lutzi has a heteroxenic life cycle, its intermediate host is probably o s et l. €  : G. geckoides  Hemidactylus mabouia Žore e a food item that is consumed by all sampled lizards in our study area.
onns  Microlophus occipitalis eters   Phyllodactylus reissi ˆerin the Trichospirura ens or t reele P. pollicaris  eters   P. inaequalis ope   P. johnwrighti Dixon & Huey, G. Geckoides  H. agrius n H. brasilianus lizards as new host records   P. microphyllus  ope   Phyllopezus lutzae oerie   for South America. Only primates from the following genera had been  Bogertia lutzae  n Tropidurus guarani Alvarez, Cei & Scolaro, 1991 previously recorded as hosts for Trichospirura : Callicebus ‘hos  T. spinulosus . Spauligodon oxkutzcabiensis was present in all ve lizard  Œ  †rihel € eibol   ƒ checo et l. Œ  Callithrix •rleben species sple in this reserch. t is thereore consiere here s the   ith € hitwoo  ƒ ˆesene et l.    Saimiri ‰oit Œ lizard’s generalist endoparasite for Phyllodactylidae and Gekkonidae, adding †rihel € eibol    Aotus llier   †rihel € eibol    H. agrius  H. brasilianus n L. klugei to the list o new recors o hosts. Callimico ’irn ˆibeiro   †rihel € eibol    n Saguinus Parapharyngodon alvarengai was recorded as being hosted by Ameivula “onnse   osroe et l.   . ocellifera pi    Cnemidophorus ocellifer  Ameiva ameiva ‘he Skrjabinelazia ens ws on inectin ec•os  heritte et l. innes   Amphisbaena ridleyi olener   Brasiliscincus  ƒ •n‚os et l.  . ”eertheless recors or S. intermedia were restricte heathi chit € ner   G. geckoides  Phyllopezus pollicaris  so r to  tei Ameivula nativo ˆoch erllo € eccinini•ele   P. periosus  Trachylepis atlantica chit   Tropidurus hispidus  Cnemidophorus nativo   Menezes et al. 2004  twice in ‘ropirie pi   T. semitaeniatus pi   n Hemidactylus agrius Tropidurus torquatus n T. guarani  ‰icente et l.  Œ  n once or Vanzolini, 1978 ( •n‚os et l.  ƒ „il et l.  ƒ rito et l. b . Dactyloidae Dactyloa punctata ‡in    Anolis punctatus   „il …e obsere P. alvarengai inectin P. pollicaris  G. geckoides  H. agrius € il  . †r t proie three host recors or S. intermedia : n H. brasilianus . P. pollicaris  H. agrius  n H. brasilianus . http://www.scielo.br/bn http:// „. oi.or /€.€/’€’ Œ’€’€ Biota Neotrop .  : e   Œ  

Helminth parasites of lizards in a Caatinga

Little is known about the biology of the genus Oochoristica . ’ost sties Cientíco e Tecnológico - CNPq (PQ-302429/2015-8) for the research on the ens were liite to escribe new species n recor new hosts ellowship ien to ….†. •lei n scholrship to
.•. •r‚o–˜ilho „il € il  ƒ rito et l. b ƒ os et l.  ). Our study n ‡.•. ‘eles. reports three recors o hosts  P. pollicaris  H. agrius n H. brasilianus  with two new inection recors or the ens Oochoristica . Author Contributions Piratuba is a genus that belongs to the Onchocercidae family. It has a wiespre istribtion throhot oth •eric in which other sties •onis •phoen ’rtins ‘eieir: contribtion to t collectionƒ reported that lizards from the genera Tropidurus  Kentropyx pi   contribution to identication of parasites and lizards; contribution to data Plica innes   Polychrus ’erre   Dactyloa ‡in analysis and interpretation; contribution to manuscript preparation and   Anolis  n Ameiva Meyer, 1795 were infected by parasites criticl reision. o the reerre ens  „il € il  . …e recore  specien •ntonio ’rcos •les ereir: contribtion to t collectionƒ contribtion Piratuba sp. inectin Gymnodactylus geckoides  which correspons to  to identication of parasites and lizards; contribution to data analysis and new recor or Piratuba . interprettionƒ contribtion to nscript preprtion n criticl reision. Currently, there are 23 genera of helminths recognized as parasites of ‡išo •les ‘eles: contribtion to t collectionƒ contribtion to the lizard families Phyllodactylidae and Gekkonidae ( •n‚os et l.  ƒ identication of parasites and lizards; contribution to data analysis and „ il € il   ƒ „il et l.  ƒ •il € il Œ ƒ rito et l. interprettionƒ contribtion to nscript preprtion n criticl reision.  ƒ os et l.  ƒ Cazorla & Morales Moreno 2015 ƒ Bezerra et al.
o›o •ntonio •r‚o ˜ilho: contribtion to t collectionƒ contribtion  ; this study); for the Caatinga biome, about 10 species of helminths to identication of parasites and lizards; contribution to data analysis and have already been reported with a prevalence above 50% ( •n‚os et l. interprettionƒ contribtion to nscript preprtion n criticl reision.  ƒ •il et l.  ƒ rito et l. b ƒ os et l.  ; this study), Samuel Cardozo Ribeiro: contribution to data collection; contribution n there were or helinth species with prelences below — or to identication of parasites and lizards; contribution to data analysis and eserts n costl res in er  Goldberg & Bursey 2010 ƒ two species interprettionƒ contribtion to nscript preprtion n criticl reision. had prevalences of 33.3% in the Brazilian Amazon forest ( „il € il el ‰ieir rito: contribtion to t collectionƒ contribtion to Œ ƒ two species h prelences p to .— in the •tlntic ˜orest identication of parasites and lizards; contribution to data analysis and „il et l.  ƒ •lei–•oes et l.  ƒ n three helinth interprettionƒ contribtion to nscript preprtion n criticl reision. species had prevalences below 15% for the tropical thorny mountain ‰neber ˜erreir i: contribtion to t collectionƒ contribtion areas of Venezuela ( Cazo rl € ’ orles ’oreno  . ”otwithstnin to identication of parasites and lizards; contribution to data analysis and that some genera and species of helminths were previously reported for interprettionƒ contribtion to nscript preprtion n criticl reision. ierent bioes s prsites o ec•os s S. oxkutzcabiensis n ll …lt‹cio e †lieir •lei: contribtion to t collectionƒ contribtion species o the ener Oochoristica n Parapharyngodon  lost ll to identication of parasites and lizards; contribution to data analysis and prelences reistere or the were below tht obtine or the tin interprettionƒ contribtion to nscript preprtion n criticl reision. bioe e.. Oochoristica sp. n species o the ens Parapharyngodo n presente p to — prelence while S. oxkutzcabiensis i not ecee Conicts of interest 8% in coastal environments of the tropical thorny mountain areas in Peru The authors declares that they have no conict of interest related to and Venezuela ( Goldberg & Bursey 2010 ƒ Cazorla & Morales Moreno the pbliction o this nscript.  . ‘hese reslts sest tht the ierent enironentl conitions of distinct biomes may favor one parasitic species over another, such as S. oxkutzcabiensis . ‘he presence o Trichospirura in lizards of the genus References Sceloporus in ’eico n H. brookii haitianus ’eerwrth   in ALBUQUERQUE  ”.ˆ.‡. COSTA-URQUIZA  •.‡.. †•ˆ•  ’.. •‰•  “ispniol  •olber et l.  Œ ƒ owell et l.    sests tht the new .. € URQUIZA , M.V.S. 2013. Diet of two sit-and-wait lizards, Phyllopezus host records documented here may not be species-specic of the studied pollicaris (Spix, 1825) (Phyllodactylidae) and Hemidactylus mabouia ’ore biome. Therefore, additional studies are necessary in the different biomes e
onns  •e••onie in  perinthropic re o ’to •rosso o to determine the true helminth diversity of these two families of lizards. Sul, western Brazil. Biota Neotrop. 13(4):376-381. http://.oi.or/. / The current study increases to 16 the number of new host records  – Œ ŒŒ lst ccess on //  in association with the families Phyllodactylidae and Gekkonidae in the •’•‡•–•†’•  ’. ‰ˆ  ˆ•‡  ‡. ’••– •ˆ”•ˆ†  ‘. € ˆ† “•  tin bioe. “oweer the eelopent o new reserch on other C.F.D. 2011. Diet and endoparasites of the lizard Gymnodactylus darwinii (Gekkota, Phyllodactylidae) from an Atlantic Rainforest area in southeastern areas of the biome is essential to enhance our scientic knowledge of the Brazil. Biotemas 25(1):203-206. doi: . /  – .   np Œ diversity of helminths associated with lizards. lst ccess on  //  •”‡•ˆ†”  ˆ. . . ”etoe prsites o ertebrtes: ‘heir eelopent Supplementary material n trnsission. e. •  nterntionl …llinor. ‘he ollowin online teril is ilble or this rticle: •”‡ˆ•‡•  ’.
.’.‡. ••  ˆ.˜.‡. € ˜ˆ•ˆ• , E.M.X. 2013. Ecology and diversity of a lizard community in the semiarid region of Brazil. Biota Neotrop. •ppeni : ‰ocher speciens o helinth prsites n respectie ŒŒ: –  . http://.oi.or/. / – Œ ŒŒ Œ lst host lizards collected in the Aiuaba Ecological Station, northeastern ccess on //  Brazil. •”‡ˆ•‡•–’•  ‡.  . ‘he ctins oini. ˆe. brs. bot. : – Œ. •”
†  .•. ˆ† “•  .˜.‡. ‰ˆ  ˆ•‡  ‡. € ‰ •”‘• 
.
. . Acknowledgements Helminths of the exotic lizard Hemidactylus mabouia ro  roc• otcrop re in southeastern Brazil. J. Helminthol. 79(4):307-313. http://.oi.or/. / We thank the support of the Coordenação de Aperfeiçoamento de
†“   lst ccess on //  essol e ”™el perior  •• or the scholrship o ‰.˜. i •.•.’. •”
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Lygodactylus Klugei (Gekkonidae), from Caatinga biome, Northeastern Brazil. ‰ Œ .‰.  ˆ–  . . ƒ. —pteis br…sileiros: ist… e esp—cies. ”eotrop. “elinthol.  : – . erpetolo•i… ˆr…sileir… „:‚ƒ. •”
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.   Œ ˆ.. ƒ‚. otes on the helinths oŽ Pseudemys scripta elegans 2013. Gastrointestinal nematodes of the lizard Tropidurus hispidus œt: ie „ in …re…s oŽ Œe€…s …n ˜l…ho…. ‹roc. ˜l…. c… . ci. :„„. Tropiduridae) from a semi-arid region of northeastern Brazil. J. Helminthol. 
 . . . e•elopent oŽ the et…cerc…ri…e oŽ Diplostomum  :Œ– . http://.oi.or/. /  •   . lst ccess exicaudum in the lenses oŽ Žro•s t•rtles bir s …n ……ls. ‡. ‹…r…sitol. on //  Š:. •ˆ•ž
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.•.•. „‰•  ˆ.…. € •’•‡•  ….†. . rsitic netoes o ZANCHI-SILVA D. 2014. Helminths infection patterns in a lizard ( Tropidurus Polychrus acutirostris (Polychrotidae) in the Caatinga biome, Northeastern hispidus  pop•l…tion Žro … sei…ri eotropic…l …re…: …ssoci…tions between Brazil. Braz. J. Biol. 74(4):939-942. http://.oi.or/. / – .ŒŒ Že…le repro •cti•e …lloc…tion …n p…r…site lo… s. ‡. ‹…r…sitol. Š:ŠŠ‚. lst ccess on / /  http:// €. oi.or•/.Šƒ/„Š. l…st …ccess on Š/Š/ƒ „‰•  ˆ.…. € ‰• , R.J. 2010. Checklist of helminths from lizards and
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. ‰eno. •ni. Gastrointestinal helminths of seven species of Sceloporine lizards from ‘oins.  :Œ– . http://.oi.or/. / –   e€ico. o•thwest. …t. :‚. http:// €. oi.or•/./„ lst ccess on //  „™:
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ec˜o. n op…r…sites. erpetol. e•. :„. lutzae (Loveridge, 1941) from the Atlantic forest in north-eastern Brazil. J. elinthol. :. http:// €. oi.or•/.‚/ ƒ„  .. GONZÁLEZ ‰..  ” , A.I. 2006a. Helminth community l…st …ccess on // str•ct•re oŽ the o•en Žro• Leptodactylus latinasus (Anura, Leptodactylidae) Žro ‰orrientes r•entin…. ct…. ‹…r…sitol. ƒ:. † .. ‡ .. ˆ  .‰.   ..   .‡.   , W.O. 2012. Nematodes of lizards (Reptilia: Squamata) from  .. ” ..  GONZÁLEZ , C.E. 2006b. Species affinity and Caatinga biome, northeastern Brazil. Comp. Parasitol. 79(1):56-63. http:// infracommunity ordination of helminths of Leptodactylus chaquensis n•r…: €. oi.or•/.Šƒ/ƒ. l…st …ccess on // Leptodactylidae) in two contrasting environments from northeastern Argentina.  ..    , R.J. 2013. Helminths of lizards from the municipality of ‡. ‹…r…sitol. Š:‚‚. Aripuana in the southern Amazon region of Brazil. J. Helminthol. 87(1):12-16. ˆ
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  , D.J. 2004. Parasites: small players with crucial roles in the ˆ • ‰. .
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. .  Œ   ‡ . . „. Rhabdias anolis ecolo•ic…l the…ter. coe…lth :ƒŠ. http:// €. oi.or•/.‚/s„„ n. sp. (Nematoda: Rhabdiasidae) from the lizard, Anolis frenatus  …•ri…: „ l…st …ccess on Š// Polychrotidae), from Panama. J. Parasitol. 89(1):113-117. http:// €. oi. or•/.Šƒ/„„ƒ„‘„:   ’..‰“ l…st …ccess on  ‰
  , D.J. 2005. Parasites of the superorganism: are they indicators of // ecosystem health? Inter. J. parasitol. 35(7):705-716. http:// €. oi.or•/.Š/ž. ižp…r….ƒ..ƒ l…st …ccess on Š// ˆ • ‰. . 
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. .  ‰ ‰. . ‰op. ‹…r…sitol. ‚:ƒŠ. http:// €. oi.or•/.Šƒ/„ l…st …ccess on 2004. Helminths infecting the parthenogenetic whiptail lizard Cnemidophorus // nativo in a restinga habitat of Bahia State, Brazil. J. Helminthol. 78(4):323-328. ˆ  ..   Œ• ”.. LOTZ ‡..   Œ” , A.W. 1997. Parasitology MESQUITA .. ‰
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.  ŒŒ , L.J. 2006. Ecology of a meets ecology on its own terms: Margolis et al. revisited. J. Parasitol. 83(4):575-583. Cerrado lizard assemblage in the Jalapão region of Brazil. Copeia 2006(3):460-471. CAZORLA .      ‹. ƒ. ‹…r•sitos intestin…les e   Œ.‰.  ˆ . . ‚. Œrichospir•rosis in o•th eric…n Thecadactylus rapicauda (Reptilia: Squamata, Phyllodactylidae) en Coro, monkeys. J. Parasitol. 57(6):1366-1368. estado Falcón, Venezuela. Rev. Científ. 25(4):346-351. ‹‰‰ . .   ..  .Œ.   .. „. ‹…r…sitiso ‰
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..   ˆ.  GENGOZIAN . Š. elinth p…r…sites and climbing in the darkness: microhabitat use, daily activity and thermal ecology oŽ the t……rin Saguinus fuscicollis . …b. ni. ‰…re. Š:ŠƒŠƒŠ. oŽ the •ec˜o Phyllopezus periosus Rodrigues, 1986. Herpetozoa 25(3/4):171-174. http://www.scielo.br/bn http://.oi.or/. / – – ”–  – Œ Biota Neotrop . ‚: eŠŠ„ ‚ ‚

Helminth parasites of lizards in a Caatinga

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„ƒ. l…st …ccess on //ƒ evolutionary novelties in lizards and snakes: sex, birth, bodies, niches, and ‹ .  ‹.‡.
 ‡..  Œ .. . cc•rrence oŽ venom. Annu. Rev. Ecol. Evol. Syst. 42(1):227-244. http:// €. oi.or•/ .Š/ Trichospirura teixeirai  pir•ri …: h…b ochoni …e in Hemidactylus brookii annurev-ecolsys-102710-145051 l…st …ccess on /Š/ƒ haitianus  …•ri…:
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. . str•t•r… … ‰o•ni … e e …•…rtos e • re…nescente ƒ‚:‚‚ƒ. de mata atlântica do Estado do Rio Grande do Norte, Brasil. Dissertação de ‰  . Œ  . ‰‰ .   
 .Œ. . estr… o. ni•ersi … e e er…l o io
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. ˆ Œ .. † .. Œ ..   ‡  ƒ:ƒ. ‡.. Œ  ... ‡ ..   .. . elinths   ... ‹   ..  ... Œ  ..    and Pentastomida of two synanthropic gecko lizards, Hemidactylus mabouia ..ˆ.    , C.L.D. 1994. Parasitism by Primasubulura jacchi …rcel …n Phyllopezus pollicaris , in an urban area in Northeastern Brazil. Braz. J. ƒ‚ n•lis ƒ …n Trichospirura leptostoma ith …n ‰hitwoo Š‚ in ˆiol. ‚:„. http:// €. oi.or•/.ƒ/ƒŠ.„ l…st …ccess Callithrix penicillata …rosets tr…ppe in the wil en•ironent …n …int…ine on /„/ƒ in captivity. Mem. Inst. Oswaldo Cruz 89(1):123-125. Œ ..  ‰Œ .ˆ. Š‚. Trichospirura leptostoma •en. et sp. ROCHA, C.F.D., ANJOS, L.A. & BERGALLO, H.G. (2011). Conquering Brazil: n. (Nematoda: Thelazioidea) from the pancreatic ducts of the white-eared the invasion by the exotic gekkonid lizard Hemidactylus mabouia (Squamata) …roset Callithrix jacchus . ‡. ‹…r…sitol. ƒ„Š:‚‚. in Brazilian natural environments. Zoologia (Curitiba) 28(6):747-754. http:// ‰Œ ‡.‡.  
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 .‰.  ‹Œ . . „. e…tŸi es €. oi.or•/.ƒ/ Š‚Š‚ //‚ do Brasil. Parte III: Nematóides de Répteis. Rev. Bras. Zool. 10(1): 19-168.   .‰. Š. ‰hec˜ list oŽ the helinth p…r…sites oŽ Žric…n ……ls oŽ ŒŒ , L.J. 1995. The ecology of tropical lizards in the caatinga of northeast Brazil. the orders Carnivora, Tubulidentata, Proboscidea, Hyra-coidea, Artiodactyla and Occasional Papers of the Oklahoma Museum of Natural History. 1(1995)1-29. Perissodactyla. 1 ed. Farnham: Commonwealth Agricultural Bureaux, St. Albans. 

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  .  MENDOZA-PALMERO ‰. ƒ. Received: 19/09/2016 elinth p…r…sites in Žreshw…ter Žish Žro the ‹…p…lo…p…n ri•er b…sin e€ico. Revised: 26/09/2017 ‹…r…sitol. res. Š:Š. http:// €. oi.or•/.‚/s„Šƒ„ƒ l…st Accepted: 01/10/2017 …ccess on /Š/ƒ Published online: 26/10/2017

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