Th e diet of an y species is related to aspects of th e local ecology of its pop u lation s as well as th e evolu tion ary h istory of th e taxon . Lizards presen t alim en tary pattern s well defin ed an d con strain ed by th eir ph ylogen y (VITTet al. 2003, VITT & PIANKA 2005). Two widely recogn ized m ain feedin g strategies defin e th ese pattern s: “widely” versu s “sit-an d-wait foragers”
(HUEY & PIANKA 1981). Exam ples of th e first strategy are lizards of th e gen u s Am eiva Meyer, 1795 an d Cnem idophorus Wagler, 1830, wh ich seek food by actively probin g th e leaf litter with th eir ton gu e (ARAUJO 1991, ZALUAR & ROCHA 2000, ROCHAet al. 2000). Th e Tropidu rid gen u s Tropidurus Wied-Neu wied, 1825 is a good exam ple of th e sit-an d-wait strategy. Th is speciou se
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André L. G. de Carvalho; Hélio R. da Silva; Alexandre F. B. de Araújo;
Ricardo Alves-Silva & Roberta R. da Silva-Leite
Laborat ório de Herpet ologia, Depart ament o de Biologia Animal, Inst it ut o de Biologia, Universidade Federal Rural do Rio de Janeiro. Rodovia BR 465, km 47, Caixa Post al 74524, 23851-970 Seropédica, Rio de Janeiro, Brasil.
E-mail: andreluizherpet o@gmail.com; helio@ufrrj.br; araujo@ufrrj.br; biologiaricardo@yahoo.com.br; robert arsl@gmail.com
ABSTRACT. We compared the effects of habitat disturbance on the feeding ecology of two local populations of Tropidurustorquatus (Wied, 1820) in restinga habitats on Ilha da Marambaia, Southeastern Brazil. Both populations presented a relatively diverse diet and differ in the number (DMax = 0.22, DF = 2, p = 0.00) and volume of prey consumed (DMax = 0.82, DF = 2, p = 0.00). Ants were the most frequent and numerous preys in both areas, and plant matter and coleopterans were the most important alimentary items in volume. The population from the disturbed area presented a larger niche breadth for number of prey items consumed (Binum = 3.06) and volume (Bivol = 2.98), when compared to the other population (Bjnum = 2.44, Bjvol = 1.52). Niche overlap was higher for number of items consumed between the populations (Oijnum = 0.82) but lower for volume (Oijvol = 0.05). Our data suggest dietary differences between the two lizard populations, and these differences may be associated with environmental modifications. Our main hypothesis to explain the dietary differences is based in the Optimal Foraging Theory. However, the alimentary behavior, and consequently the diet of these lizards, seems to be constrained phylogenetically, with a pattern that may have evolved in the ancestor of all iguanids. KEY WORDS. Diet; lizards; Optimal Foraging Theory.
RESUMO. Ecologia Ecologia Alimentar de Ecologia Ecologia Ecologia Alimentar de Alimentar de Alimentar de Alimentar de TTTTTrrrrropidurusopidurusopidurusopidurusopidurus torquatustorquatustorquatustorquatustorquatus (W (W (W (W (Wied) (Sauried) (Sauria,ied) (Sauried) (Sauried) (Sauria,ia,ia, Tia,TTTTrrrrropiduropiduropiduridae) em Duas Áropiduropiduridae) em Duas Áridae) em Duas Áridae) em Duas Áridae) em Duas Áreas com Difeas com Difeas com Difeas com Difeas com Dife-e-e-e- e-rrrrrentes Níventes Níveis de Conserventes Níventes Níventes Níveis de Conserveis de Conserveis de Conserveis de Conservação na Ilha da Maração na Ilha da Maração na Ilha da Maração na Ilha da Marambaia,ação na Ilha da Marambaia,ambaia,ambaia,ambaia, Rio de J Rio de J Rio de J Rio de J Rio de Janeiraneiraneiraneiro,aneiro,o, Sudeste do Bro,o, Sudeste do Br Sudeste do Br Sudeste do Brasil. Sudeste do Brasil.asil.asil.asil. Comparamos os efeitos da degradação ambiental sobre a ecologia alimentar de duas populações de Tropidurustorquatus (Wied, 1820) em áreas de restinga da Ilha da Marambaia, sudeste do Brasil. Ambas as populações apresentaram dieta relativamente diversa e deferiram em número (DMax = 0,22; DF = 2; p = 0,00) e volume de presas consumidas (DMax = 0,82; DF = 2; p = 0,00). Formigas foram os itens mais freqüentes e numerosos em ambas as áreas, e material vegetal e coleópteros os mais importantes em volume. A população da área perturbada apresentou maior largura de nicho alimentar para número de presas consumidas (Binum = 3,06) e volume (Bivol = 2,98), quando comparada à outra população (Bjnum = 2,44; Bjvol = 1,52). A sobreposição de nicho foi mais marcante para o número de itens consumi-dos entre as populações (Oijnum = 0,82) e menos marcante para volume (Oijvol = 0,05). Nossos dados sugerem diferenças nas dietas entre as duas populações de lagartos, e estas diferenças parecem estar associadas à degradação ambiental. Nossa principal hipótese para explicar as diferenças nas dietas embasa-se na Teoria do Forrageamento Ótimo. Entretanto, o comportamento alimentar, e conseqüentemente a dieta dos lagartos, mostrarem-se limitadas filogeneticamente, com um padrão que pode ter evoluído no ancestral de todos os iguanídeos.
gen u s of open form ation s dwellers fou n d in tropical an d su b-t ro p ical So u b-t h Am erica, in clu d in g b-t h e G alap ago s Islan d s (RODRIGUES 1987, LOPESet al. 1992, FROSTet al. 2001), presen ts several ad ap tation s related to th is strategy, su ch as crip tical coloration , robu st body, territorial beh avior, an d diet based on m obile p rey (RAND & RAND 1966, RO DRIGUES 1987, BERGALO & ROCHA 1994). In Brazil, Tropidurus is p resen t in th e Am azon , Cerrado, Caatin ga an d Atlan tic Forest, in clu din g th e coastal Restin gas (RODRIGUES 1987, ROCHA 2000).
Tropidurustorquatus (Wied, 1820) is th e m ost widely dis-tribu ted lizard in th e gen u s, occu rrin g from Sou th eastern Bra-zil to th e n orth of Argen tin a (RODRIGUES 1987). Its diet is based on h igh ly vagile arth ropods, prim arily an ts, an d vegetal m at-ter, esp ecially fru its (ARAÚJO 1991, ROCHA & BERGALO 1994, FIALHO et al. 2000). Several aspects of its diet, foragin g m ode, an d be-h avioral ecology were stu died (RAND & RAND 1966, BERGALO & ROCHA 1994, TEIXEIRA & GIOVANELLI 1999, ROCHAetal. 2000). How-ever, th e effects of en viron m en tal m odification s to its feedin g ecology h ave n ot been con sidered yet.
Ch an ges in th e en viron m en tal con dition s can affect prey offer, pattern s of diet, an d costs of foragin g. Accordin g to th e Optim al Foragin g Th eory (MACARTHUR & PIANKA 1966), lizards from species-rich feedin g areas wou ld presen t n arrower food n ich e breadth , wh ile lizards from areas with less den se an d less n u tritive prey wou ld presen t larger alim en tary n ich e breadth in o rd er t o co m p en sa t e fo r t h e lo w q u a lit y o f t h e p rey (MACARTHUR & PIANKA 1966, SCHOENER 1971). Con seq u en tly, liz-ards livin g in a stressed en viron m en t wou ld redu ce th eir spa-tial n ich e to u se on ly th e best p atch es (MACARTHUR & PIANKA 1966). Th erein , we tested th is assu m ption by com parin g th e effects of h abitat distu rban ce on th e feedin g ecology of two local p op u lation s of Tropidurustorquatus th at occu r in restin ga h abitats on Ilh a da Maram baia, Rio de Jan eiro, Sou th eastern Brazil, on e preserved, an d th e oth er distu rbed.
MATERIAL AND METHODS
Th is work was developed on Ilh a da Maram baia, located in Rio d e Jan eiro , So u t h east ern Brazil (23º04’S, 43º53’W ). Maram baia presen ts a m arked spatial h eterogen eity, in cludin g rain forests on th e h ills (Atlan tic Forest), restin ga (lon g san d bars con n ectin g th e h ills to th e con tin en t, covered with open veg-etation ), an d m an grove (MENEZES & ARAUJO 2005). Two sites in th e restin ga h abitat with differen t degrees of con servation were ch osen for sam plin g th e lizards (Fig. 1). Th e first, called Bravo-m eia (23º04’16.36”S, 43º56’43.43”W) presen ts a well-preserved m osaic of sh rubs. Th e oth er, called Heliporto (23º03’40.50”S, 43º59’24.47”W) is a sm all an d isolated restin ga area, bordered by secon dary forest outgrow, both deforested an d burn ed peri-odically; trash is also dum ped in th e area by residen ts.
Fieldwork was u n dertaken from Febru ary 2003 to Sep tem ber 2004. Th e diets were com pared an alyzin g stom ach con -ten ts. We collected 37 specim en s from Heliporto an d 32 from Bravo-m eia, totalizin g 69 lizards. Th e an im als were eu th an ized
with an overdose of lidocain 0,5% an d fixed in form alin 10% wh ile in th e field. Th e collected lizards were posterior sexed an d m easu red (sn ou t-ven t-len gth ) in laboratory. Th e an im als were th en dissected for th e in vestigation of th e stom ach con -t en -t s, u sin g a s-t ereo m icro sco p e. All alim en -t ary i-t em s were cou n ted an d m easu red with a caliper (len gth an d breadth , ± 0.05 m m ). Th e volu m e of th e alim en tary item s were estim ated th rou gh th e ellipsoid form u la (COLLIetal. 1992).
Volume
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We calcu lated th e alim en tary n ich e breadth for n u m ber of item s an d volu m e con su m ed, for th e two areas, followin g PIANKA (1973). W h ere P is th e proportion al u tilization of th e prey item i an d n is th e n u m ber of prey categories u sed.
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To evalu ate th e sim ilarity between th e diets of th e two lizard p op u lation s, we u sed th e n ich e overlap in d ex (PIANKA 1973), sym bols th e sam e as above, bu t j an d k represen t each lizard popu lation .
f
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We tested d ifferen ces in size between th e two lizard ’s
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Figure 1. M ap of the State of Rio de Janeiro, with the location of M arambaia (square). Enlarged, M arambaia Island with part of the restinga and indication of study areas: (H) Heliporto, (B) Bravo-meia.
popu lation s u sin g th e T Test. Diet differen ces were tested u sin g Kolm orogovSm irn ov. Th e Z Test was perform ed to evalu -ate differen ces in th e con su m ption (n u m ber an d volu m e) of each p rey category sep arately. To evalu ate differen ces in th e m ean n u m ber, size an d volu m e of prey item s con su m ed in th e two areas, we u sed th e Man n -W h itn ey Test. Th e relation sh ip between th e sn ou t-ven t-len gth (SVL) an d n u m ber an d volu m e of prey item s con su m ed was evalu ated u sin g lin ear regression an alysis. Th e statistical m eth odology followed ZAR (1999).
RESULTS
Accordin g to T Test, th e two lizard population s do n ot differ in size (T = 0.33, DF = 67, p = 0.74). Both population s presen ted a relatively diverse diet, with 19 prey categories con -sum ed by th e population from Bravo-m eia an d 18 prey catego-ries con sum ed by th e population from Heliporto. However, just 15 prey categories are com m on to both popu lation s (Tab. I). Despite th e sim ilarity in th e n um ber of prey categories con sum ed, th e population s differed in n um ber (Kolm ogorov-Sm irn ov, DMax = 0.22, DF = 2, p = 0.00) an d vo lu m e o f p reys co n su m ed (Kolm ogorov-Sm irn ov, DMax = 0.82, DF = 2, p = 0.00). An ts were
th e m ost frequen t an d n um erous prey item s in both areas. Volu-m etrically, vegetal Volu-m atter was th e Volu-m ost iVolu-m portan t aliVolu-m en tary category for th e population from Bravo-m eia, wh ile for Heliporto it was Coleoptera (Tab. I). Th e Z Test in dicated differen ces in th e proportion of volum e con sum ed for th e m ajority of prey item s an d to h alf of th em , in n um erical proportion (Tab. II). Sim ilar-ity in th e n um erical con sum ption of th e prey item s was observed just for th ose less-con sum ed categories.
Th e population from Heliporto presen ted a larger n ich e breadth for n um ber of prey item s con sum ed (Bin um = 3.06) an d volum e (Bivol = 2.98) wh ile th e population from Bravo-m eia pre-sen ted lower values (Bjn um = 2.441, Bjvol = 1.52). Th e n ich e over-lap was superior for n um ber of item s con sum ed between th e population s (Oijn um = 0.82) an d lower for volum e (Oijvol = 0.05). Th e results of th e Man n -Wh itn ey Test in dicated th at th e m ean n um ber (U = 525.50, 2 = 0.64, DF = 1, p = 0.42), m ean
size (U = 444.00, 2 = 3.17, DF = 1, p = 0.07) an d m ean volum e
(U = 610.00, 2 = 0.05, DF = 1, p = 0.83) of prey item s con sum ed
by th e lizards from th e two areas were n ot statistically differen t. Th e lin ear regression an alysis in dicated n o relation sh ip between body size an d n u m ber (Heliporto: r2 = 0.01, F = 0.22, p
Table I. Number of items, volume (mm3) and frequency of each prey consum ed by the two populations of Tropidurustorquatus studied
on Ilha da M arambaia, Rio de Janeiro.
Prey Bravo-m eia (n = 32) Heliporto (n = 37)
= 0.64 – Bravo-m eia: r2 = 0.02, F = 0.52, p = 0.48) or volu m e
(Helip orto: r2 = 0.09, F = 0.78, p = 0.38 – Bravo-m eia: r2 = 0.09,
F = 3.05, p = 0.09) of prey item s con su m ed.
breadth , in dicatin g a m ore gen eralist diet. However, th e total n u m ber of prey categories con su m ed was sim ilar for th e two popu lation s, with 15 (ou t of 22) item s in com m on , in dicatin g th at, in gen eral, diets were n ot radically differen t. Th e h igh con su m ption of an ts by th e two popu lation s, for exam ple, re-in fo rce a well-kn o wn p at t ern o f it em co n su m p t io n am o n g tropidu rids, th at are sit-an d-wait lizards (HUEY & PIANKA 1981) an d presen t diet dom in ated by som e com m on m obile item s like an ts (VITTetal. 2003, VITT & PIANKA 2005).
Several works report large usage of an ts by sit-an d-wait lizards, especially am on g Tropidurus. Th e con sum ption of an ts is extrem ely com m on in species of differen t h abitats an d locali-ties. Good exam ples of th e frequ en cy of th is beh avior are T. oreadicus Rodrigues, 1987 (ARAUJO 1987, FARIA & ARAUJO 2004), T. itam bere Rodrigues, 1987 (VAN-SLUYS 1993, FARIA & ARAUJO 2004), an d T.spinulosus (Cope, 1862) (COLLIetal. 1992) from Brazilian Cerrado; T. hispidus (Spix, 1825) (VITTet al. 1996) in isolated rock outcrops in Am azon ia; or T.m elanopleurus Boulen ger, 1902, an An d ean sp ecies (PEREZ-MELLADO & DE-LA-RIVA 1993). Th e tropidurid gen us Liolaem us Wiegm an n , 1834 is an oth er exam ple of a sim ilar dietary pattern . L.lutzae Merten s, 1938, a san d dun e lizard of coastal form ation s from Rio de Jan eiro, Brazil, in addi-tion to h igh con sum paddi-tion of vegetal m atter, uses in sect larva, coleopteran s, an d an ts in large am oun ts (ROCHA 1989).
Th e am ou n t of vegetal m atter con su m ed was n otably altered, bein g lower in Heliporto wh en com pared to Bravo-m eia. Th ese results suggest th at m odification s in th e local flora caused by bu rn , deforestation , an d soil overtu rn m ay affect th e diet of t h e lizard s b y alt erin g t h e availab ilit y o f it em s. Tropidurus torquatus is kn o wn fo r it s co n su m p t io n o f veget al m at t er (TEIXEIRA & GIOVANELLI 1999, FIALHO et al. 2000). In Maricá, an -oth er area of restin ga in Rio de Jan eiro, FIALHOetal. (2000) docu -m en t ed h igh u t ilizat io n o f fru it s b y t h is sp ecies. Fru it s o f Erythroxylonovalifolium Peyritisch , 1879 (Eryth roxylaceae) rep -resen ted abou t 50% of th e diet of adu lt m ales sam pled du rin g th e su m m er, th e peak of th is plan t’s fru it-bearin g season .
FIALHOetal. (2000) also sh owed th at vegetal m atter con -su m ption was positively correlated to th e body size of th e liz-ards, poin tin g to an in crease of h erbivory du rin g lizard growth . Con siderin g th e sign ifican t u se of plan ts reported for Tropidurus torquatus, an d con firm ed in th is work for th e lizards from th e m ore in tact restin ga area, we can su ppose th at ch an ges in th e com position of plan ts m ay h ave been cau sed by en viron m en -tal distu rban ces – ch an gin g or depressin g th e offer of eatable fru its, leaves an d flowers – an d th at it m ay h ave affected th e lizard’s diet, perh aps alterin g on togen etic feedin g preferen ces. Th e im portan ce of vegetal m atter in th e diet of th e Tropidurus is rein forced by an alyzin g th e n ich e overlap valu e for volu m e. Th e low valu e obtain ed in dicates th at vegetal m atter is volu -m etrically on e of th e -m ost i-m p ortan t ite-m s, d ifferin g sen si-tively in th e diet of th e two Tropidurus popu lation s stu died.
Ou r resu lts also in dicate th at besides th e depression of h erbivory, ch an ges in th e freq u en cy of con su m ption of som e
DISCUSSION
Ou r data su ggest dietary differen ces between th e two liz-ard popu lation s. Given th at lizliz-ards from th e two popu lation s do n ot differ in size, th ese differen ces m ay be associated with en viron m en tal differen ces between th e two areas an d n ot to m orfom etrical differen ces between th e popu lation s. Ou r m ain h yp oth esis to exp lain th e dietary differen ces is based in th e Optim al Foragin g Th eory (MACARTHUR & PIANKA 1966, SCHOENER 1971). Accordin g to th is th eory, as prey offer (an d q u ality) deteriorates in a given area, th e alim en tary n ich e breadth is en -larged (MACARTHUR & PIANKA 1966).
Th e valu es for alim en tary n ich e breadth fou n d for th e two lizard popu lation s su pport th ese assu m ption s. Lizards from th e disturbed area (Heliporto) presen ted larger alim en tary n ich e
Table II. Results of the Z Test for differences in the proportion of consum ption (number and volume) of each prey category used by t he t w o populat ions of Tropidurust orquat us studied on Ilha da M aram baia, Rio de Janeiro. Significant p values are in bold face.
Taxon Num ber of items Volum e Z P Z P Acarina 1.336 0.091 1.630 0.052
Aranae 1.932 0.027 28.798 0.000 Blattaria 1.662 0.048 3.066 0.001 Chilopoda 0.587 0.279 1.862 0.031 Coleoptera -2.504 0.006 0.000 0.000 Diplopoda -0.749 0.227 -11.189 0.000 Diptera 3.341 0.000 -4.316 0.000 Egg -0.749 0.227 -6.945 0.000 Formicidae 7.887 0.000 -43.356 0.000 Gastropoda -0.094 0.462 -1.640 0.050
Hemiptera -2.208 0.014 -42.243 0.000 Homoptera -0.407 0.342 -40.001 0.000 Hym enoptera -0.113 0.455 -7.301 0.000 Isopoda -2.124 0.017 -17.902 0.000 Isoptera -11.389 0.000 -32.412 0.000 Larva -1.696 0.045 -42.251 0.000 M antodea 1.336 0.091 3.489 0.000 Neuroptera 1.336 0.091 5.155 0.000 Odonata 1.336 0.091 1.630 0.052
oth er item s were recorded in th e distu rbed area. It is possible th at th ese even ts are associated. Th erefore, in order to com -pen sate for th e redu ction of vegetal m atter in th e diet, th e liz-ards apparen tly su bstitu te oth er item s in h igh er freq u en cy, for exam p le coleop teran s, as registered for th e p op u lation from Heliporto. Despite som e differen ces regardin g prey item s in th e diet of th e two popu lation s, th e com m on prey do n ot differ in size or volu m e, in dicatin g th at th e distu rbed area, in gen eral, su stain s a sim ilar su pply of arth ropods th at possibly differ in freq u en cy, as observed in th e stom ach con ten ts of th e lizards. An oth er h ypoth esis th at can accou n t for th e differen ces in t h e freq u en cy o f co n su m p t io n o f d ifferen t gro u p s o f arth rop ods am on g th e two areas relates to th e p ossibility of in com in g an d/or in creasin g of som e arth ropod population s th at wou ld n ot be p resen t, or in h igh den sity, in th e area u n der n orm al, u n d istu rbed , con d ition s. Un fortu n ately, d u rin g th e fieldwork, sam ples of th e available arth ropods were n ot taken . Th erefore, we can n ot properly evalu ate th is h ypoth esis.
In addition to th e aforem en tion ed, in Heliporto, th e liz-ards are com m on ly observed visitin g th e forested area n ear th e restin ga, u sin g tru n ks an d bran ch es as perch es for th erm oregu -lation , an d possibly as foragin g sites. Th e u se of th is bou n dary m ay be reflected in th e diet of th e lizards from Heliporto.
No relation sh ip between lizards body size an d prey n u m -ber an d volu m e were observed in both areas stu died, con trary to wh at was expected accordin g th e Optim al Foragin g Th eory (MACARTHUR & PIANKA 1966, SCHOENER 1971). Th is th eory p dicts th at an optim al prey size wou ld be con su m ed with re-du ced tim e of p rey h an dlin g by th e p redator. However, it is p o ssible t h at t h is exp ect ed effect o n gen eralist lizard s like Tropidurustorquatus is less stron g th an in m ore specialized on es; h owever, th is will req u ire fu rth er in vestigation .
In su m m ary, an alysis of th e stom ach con ten ts in addi-tion to con sideraaddi-tion of lizards as good sam plers of th e food offer, th e data in dicate th at en viron m en tal m odification s m ay affect th e feedin g h abits of Tropidurustorquatus, esp ecially in p lan t co n su m p t io n , as p red ict ed b y t h e O p t im al Fo ragin g Th eory. Th e availability of prey item s n eeds to be m easu red in fu tu re stu dies. Overall, h owever, th e dietary pattern of th is liz-ard species seem s to be con servative, followin g a diet ph yloge-n etically determ iyloge-n ed perh aps iyloge-n th e com m oyloge-n ayloge-n cestor of all igu an ids (VITTet al. 2003, VITT & PIANKA 2005), sin ce th is beh av-ior h as been observed even in altered con dition s.
ACKNOWLEDGEMENTS
To th e Brazilian Navy, in special th e com m an d of CADIM (Cen tro de Adestram en to da Ilh a da Maram baia), th at allowed u s to access th e Islan d of Maram baia an d u se som e of th e fa-cilities of CADIM du rin g field work an d to CNPq (Th e Nation al Cou n cil for Scien tific an d Tech n ological Developm en t) by th e Scien tific In ciation Fellowsh ip given to ALGC an d by th e Gran t 471081/ 2004-3 given to HRS. Mary S. Lin n (Un iversity of Okla-h om a) for reviewin g tOkla-h e m an u script.
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