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Digestive Ecology of Two Omnivorous Canarian Lizard Species

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Digestive Ecology of Two Omnivorous Canarian Lizard Species Digestiveecology of two omnivorous Canarian lizardspecies (Gallotia,Lacertidae) AlfredoV alido 1,ManuelNogales Departamento deBiología Animal (Zoología), Universidadde La Laguna,E-38206, Tenerife, Canary Islands, Spain 1Authorfor reprint request; current address: Dept. of Ecologyand Genetics, AarhusUniversity, Ny Munkegade Building540 DK-8000 Aarhus C, Denmark e-mail: [email protected] Abstract. Omnivorousendemic Canarianlacertids ( Gallotiaatlantica and G. galloti)donotpresent any speci c digestiveand physiological adaptations to herbivorous diet, compared to species andpopulations with a different degree ofherbivory in the Canarian archipelago. The only characteristics thatcould be related tothe type of diet were thenumber of cusps per tooth (between species) andthe number of small stonescontained in droppings (between species andpopulations). The rest ofmeasured traits were correlated withlizard size andfor this reason G. galloti has longerintestines, heavier stomachs andlivers, more teethand cusps, and longer gut passage. These data suggestthat body size isamajor determinantof the reliance onplant food (mainly eshyfruits) in these lizards andfacilitates mutualisticinteractions with eshy-fruitedplant species. Introduction Structuraland functional adaptations of the digestive system for optimalprocessing of plantmatter have been described for severalomnivorous and herbivorous vertebrates (see reviewsin Lö nnberg, 1902; Ziswiler and Farner, 1972; Skoczylas, 1978; Chivers and Hladik,1980; Guard, 1980; Sibly, 1981; Jordano, 2000). Althougha lowproportion (about 2-3%) of knownlizard species eat some quantities of plantfood (Pough, 1973; Iverson, 1982; King, 1996), several morphological, anatomical andphysiologicalmodi cations have also been described for herbivorouslizards (Gekkota, Iguania,and Scincomorpha), which may increase the digestiveef ciency of lizardsfeeding onthis resource (Throckmorton, 1973; Johnson and Lillywhite, 1979; Zimmerman and Tracy,1989). The mostimportant of thesetraits are: larger body size(Szarski, 1962; Sokol, 1967;Pough, 1973; Iverson, 1982; Schluter, 1984, butsee V anDamme, 1999);tendency to showa largerintestine/ SVL ratio(Skoczylas, 1978; V anDevender, 1982; Dearing, 1993); © KoninklijkeBrill NV ,Leiden,2003 Amphibia-Reptilia24: 331-344 Alsoavailable online - www.brill.nl 332 AlfredoV alido,Manuel Nogales longertransit time that permits a suitabledegradation of plant tissues (Christian et al., 1984;Troyer, 1984; Karasov, 1986; W aldschmidtet al., 1986; Zimmerman and Tracy, 1989;V anMarken Lichtenbelt, 1992); smaller stomachs (Skoczylas, 1978); three-cusp or polycuspdentition (Hotton, 1955; Sokol, 1967; Montanucci, 1968; Greer, 1976;Iverson, 1982;Mateo and Ló pez-Jurado, 1992); intestinal caeca and compartmentalizati onofthe colon(Iverson, 1982; Zug, 1993); and the tendency to ingest small stones (Sokol, 1971; Skoczylas,1978). Modications to lizard digestive systems have been described mainly in folivorous speciesand their presence has been justi ed, by the need to process ber-richplant components,more dif cult to digest (Iverson, 1982; Cooper and V itt,2002). However, littleis known about the digestive modi cation of lizards that primarily feed on other, comparativelyeasy to digest, plant components (basically fruits, owers andnectar) although,in some cases, the proportion of thisplant component in the diet is considerable. Generally,lizards inhabiting island habitats have a tendencytowards a moreherbivorous dietthan lizards on mainlands (V anDamme, 1999;Cooper and V itt,2002); this is the caseof theendemic Canarian lacertids (g. Gallotia),whichpresent the most herbivorous specieswithin Lacertidae (V alido,1999; V anDamme, 1999). The seven extant Canarian lacertidscan be considered omnivorous, but the contribution of plantmatter in their diet variesaccording to lizard species, body size, habitat, and season of theyear (e.g. Mateo andLó pez-Jurado, 1992; V alidoand Nogales, 1994; Pé rez-Mellado et al., 1999; V alido, 1999).However, in some cases (e.g. G. galloti from xerophyticzones in Tenerife), the plantcomponent (basically eshyfruits) is predominantthrough the year (V alido,1999). Theprimary aim of thisstudy is to characterizethe assemblageof traitsin some Canarian lizards.W eselecteda pairof speciesthat are more distant their diet along an omnivorous gradient:G.atlantica from Fuerteventura,which is omnivorousbut basically insectivorous, andthree distant populations of G. galloti from Tenerife(T enoBajo, Bajamar e Izaña), whichvary in the importance of the plant component in their diet (from omnivorous, basicallyinsectivorous to omnivorous, mainly herbivorous). Lizardsof the genus Gallotia (Lacertidae)represent an oceanic radiation of reptiles endemicto CanaryIslands and are resolved as abasaltaxon in the family Lacertidae (Fu, 1998).This genus is characterized by seven extant species and one extinct taxon. This genushas a monophyleticorigin and the radiation took place along an eastern-western geographicgradient from themainland (Gonzá lez et al.,1996). Gallotiaatlantica inhabitsFuerteventura and Lanzarote; it isthesmallest species of the genus(adult SVL mean s 60:8 9:9mm;V alido,1999) and shows an omnivorous § D § diet,although animal components are the main preys. Gallotiagalloti inhabitsthe islands ofTenerifeand La Palma; it isa medium-sizedlizard (SVL: mean 106:4 12:1; Valido, D § 1999)and is characterized by an omnivorous diet in which eshyfruit is the principal componentin those populations inhabiting altitudinal lower zones (V alidoand Nogales, 1994;V alido,1999). Omnivorouslizards 333 If thetraits described above represent adaptations for anherbivorous diet, we would expectthe more insectivorous populations ( G.atlantica from Fuerteventuraand G. galloti from Izaña inTenerife)to have similar characteristics (shorter intestines and transit period, heavierstomachs, presence of mono/bicuspiddentition) and to differ from thetwo more herbivorouspopulations ( G. galloti from TenoBajo and Bajamar). However, an alternative hypothesisis thatthese lizards do notneed special anatomical traits like true herbivorous lizards(Iverson, 1982; Cooper and V itt,2002), since the principal parts of theplant in the dietare eshyfruits and owers. Theuse of these species can also help us to understand the mechanisms in uencing dietaryvariations among insular lizard populations, speci cally the general trend for relativelymore herbivorous diets when compared to continentalareas (V anDamme, 1999; Cooperand Vitt, 2002). This insular trend to increase herbivory (mainly fruits) has also importantecological and evolutionaryimplications for mutualisticinteractions with eshy- fruitedplant species in islands(V alido,1999), a phenomenonwhich seems to occurmore frequentlyin islandecosystems (Olesen and V alido,2003). Materialand methods Thelizards understudy were capturedin July 1994 at fourdifferent localities on two islands: G.atlantica in Fuerteventura(V alle deTetir — 400m a.s.l.;Tetir hereafter) and G. galloti inthree differenthabitats of Tenerife (Barranco deLas Cuevas,Teno Bajo — 150m a.s.l.;Barranco deV argas, Bajamar —110m a.s.l.;and Corral del Niño, Izañ a —2293m a.s.l.;hereafter TenoBajo, Bajamar andIzañ a, respectively). Theclimate ofTetir is characterized byan annual rainfall of less than200 mm andmean temperature of 21±C.Characteristic plantsof these xeric zonesare Launaeaarborescens (Compositae), Salsolavermiculata (Chenopodiaceae),and Lyciumintricatum (Solanaceae). InTeno Bajo, the climate is dry,with an annual rainfall < 300mm, anda mean temperature ofabout 21 ±C.Themore abundantplants in this habitat include some species belongingto the genus Euphorbia: E.obtusifolia , E.canariensis , E.balsamifera .Inaddition, Plocama pendula, Rubiafruticosa (Rubiaceae) andthe introduced Opuntiadillenii (Cactaceae) are abundant.The other localityon Tenerife (Bajamar) presentsa mean annualrainfall < 400mm, amean temperature of21 ±C and is characterized byscrubvegetation where Asparaguspastorianus , A.umbellatus (Liliaceae), and Boseayervamora (Amaranthaceae), are abundant.The third locality on Tenerife, Izañ a, is ahighmountain site witha cool,dry climate andannual rainfall of about600 mm andmean temperature of10 ±C;inwinter, rainfall can turnto snow. Spartocytisussupranubius and Adenocarpusviscosus (Fabaceae) composeshrubs basically. Procedures. We collectedrecent lizarddroppings ( n 150,at each site) ineach ofthe four localities, in July D 1994,in order to study the diet composition. Therefore, in order to cover those different microhabitats within each localityand to have a representativeapproximation of the lizard diet at populationlevel, linear transects were carried out.To avoid seudoreplication during the sampling collection, no more than velizard droppings were collectedat thesame site.Each droppingwas keptindividually and analyzed under 16 magnication after £ thematerial hadbeen soaked in water. We visuallyestimated thepercentage volumein each faecal sample made upby animal andplant matter (tothe nearest 10%),and the number of animal items andseeds were recordedand identied. Small stonesinside droppings were alsonoted. Although this method is less accurate thanstomach contentsto trying to knowa taxonomicdescription of invertebratescomponents, in the approximation carried out inthis study (animal vs vegetalcomponent), feces analyses gavesimilar resultsto those from stomach (angular transformationon percentages ofplant matter; ANOVA, F 0:016, df 1, 118, P 0:89forcomparative data D D D obtainedfrom
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