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Size- and Sex-Dependent Variation in Diet of Rhinella Arenarum (Anura: Bufonidae) in a Wetland of San Juan, Argentina

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Size- and Sex-Dependent Variation in Diet of Rhinella Arenarum (Anura: Bufonidae) in a Wetland of San Juan, Argentina Size- and Sex-Dependent Variation in Diet of Rhinella arenarum (Anura: Bufonidae) in a Wetland of San Juan, Argentina Authors: Quiroga, Lorena B., Sanabria, Eduardo A., and Acosta, Juan C. Source: Journal of Herpetology, 43(2) : 311-317 Published By: Society for the Study of Amphibians and Reptiles URL: https://doi.org/10.1670/07-117R2.1 BioOne Complete (complete.BioOne.org) is a full-text database of 200 subscribed and open-access titles in the biological, ecological, and environmental sciences published by nonprofit societies, associations, museums, institutions, and presses. Your use of this PDF, the BioOne Complete website, and all posted and associated content indicates your acceptance of BioOne’s Terms of Use, available at www.bioone.org/terms-of-use. Usage of BioOne Complete content is strictly limited to personal, educational, and non - commercial use. Commercial inquiries or rights and permissions requests should be directed to the individual publisher as copyright holder. BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit publishers, academic institutions, research libraries, and research funders in the common goal of maximizing access to critical research. Downloaded From: https://bioone.org/journals/Journal-of-Herpetology on 12 May 2020 Terms of Use: https://bioone.org/terms-of-use Access provided by Bing Search Engine SHORTER COMMUNICATIONS Journal of Herpetology, Vol. 43, No. 2, pp. 311–317, 2009 Copyright 2009 Society for the Study of Amphibians and Reptiles Size- and Sex-Dependent Variation in Diet of Rhinella arenarum (Anura: Bufonidae) in a Wetland of San Juan, Argentina 1 LORENA B. QUIROGA, EDUARDO A. SANABRIA, AND JUAN C. ACOSTA Departamento de Biologı´a, Instituto y Museo de Ciencias Naturales, Universidad Nacional de San Juan, Avenida Ignacio de la Roza 590 (O), Co´digo Postal:5400, Argentina ABSTRACT.—We studied body-size and sex-dependent variation in the diet of Rhinella arenarum in a wetland of San Juan, Argentina. We hypothesized that prey size would be positively correlated with toad size and that the guts of larger toads would contain fewer prey items. Toads from this population eat primarily ants and, secondarily, beetles, indicating a feeding strategy that is intermediate between specialist and generalist. This feeding strategy may be influenced by prey availability. Contrary to our expectation, prey size was not related to toad body size, and the relationship between the prey number and toad body size was positive. Our findings, coupled with similar diet studies of toads, suggest geographically widespread phylogenetic conservatism in the diet of bufonids. Both extrinsic factors, such as seasonal food remain dry (Victoria, 1999). The region belongs to the abundance and the presence of competitors and Monte phytogeographic province, which has an arid predators, and intrinsic factors, such as morphological climate and a mean annual temperature of 17.3uC, a constraints resulting from ontogeny, body size, and mean maximum annual temperature of 25.7uC, a specialization (Toft, 1980; Duellman and Trueb, 1986) mean minimum annual temperature of 10.4uC, and a influence an amphibian’s diet. For example, the size of mean annual rainfall of 89 mm, which falls mainly in a frog (or its gape) is often a good predictor of the size summer (Cabrera, 1994). The dominant plants of this of prey it will take. However, larger individuals tend Monte region include Cortaderia sp., Typha dominguen- to consume prey of a wider range of body sizes (Toft, sis, Malvella leprosa, Phyla canescens, Melilotus indicus, 1980; Hirai and Matsui, 1999, 2000, 2002b; Menendez- Gnaphalium sp., Cyperus sp., Prosopis strombulifera, Guerrero, 2001; Hirai, 2002). Prosopis sp., Atriplex sp., and Larrea spp. We studied the diet of a population of the common Toads were collected from 28 November 2001 to 5 toad Rhinella arenarum, in an arid region of San Juan, October 2002, except during winter (May to July), Argentina. Rhinella arenarum is widely distributed in when the animals are not active. A total of 149 Argentina from northern Jujuy Province to the Rı´o individuals was haphazardly collected during day- Chubut in Patagonia. Despite being one of the most and nighttime visual-encounter surveys (Heyer et al., common anurans in this region, most aspects of the 2001), which took place approximately every 10 days ecology of this toad are not well known (Cei, 1980; during the sampling period. Specimens were imme- Gallardo, 1987). Bufonids are widely considered diately sacrificed, fixed in 10% formalin, and pre- generalist predators (Lajmanovich, 1995; Parmelee, served in 70% alcohol. Body length (snout–vent 1999; Hirai and Matsui, 2002b; Moseley et al., 2005). length; SVL) and mouth width (MW) were measured However, most of the toads studied to date eat with digital calipers (0.01 mm precision) and mass primarily beetles and ants, the latter of which can be with a digital balance (Denver Pk-1201; 0.1 g preci- ingested in large quantities because of their often- sion). Sex was determined by dissection; postmeta- clumped distribution (Menendez-Guerrero, 2001). morphic, yet sexually immature individuals were R. arenarum Our objective was to analyze the diet of , considered subadults up to 8 cm. providing basic information on the biology of this Stomachs were excised between the cardiac and widespread species. We predicted that larger R. pyloric sphincters, and their contents were analyzed arenarum from our population would eat larger and under a binocular dissecting microscope (3 2–40). The a wider of range prey items than would smaller length and width of each prey item was measured individuals. Specifically, we hypothesized that (1) prey size would increase with toad size and that (2) with digital calipers and prey volume was estimated 5 p 2 the number of prey would decrease with the size of using the equation [V 4/3 (L/2) (W/2) ] for an the toad. elliptical sphere (Dunham, 1983). Prey items were classified to the lowest taxonomic level allowed by MATERIALS AND METHODS their state of digestion. We studied a population of R. arenarum from a The percentage of plant tissues was estimated with seasonal wetland produced by the Ullu´m Reservoir respect to the total amount of food. An index of 25 km west of San Juan, Departamento Zonda, relative importance (IRI 5 %FO [%N + %V] [Pinkas et Provincia de San Juan, Argentina (31u559S, 68u709W; al., 1971]) was used to compare the dietary contribu- 800 m). The wetland has uneven terrain, with the tion of each food category; where %FO is the lower areas flooding in summer, while higher areas frequency of occurrence of food category, %N is the numerical percentage, and %V the volumetric per- centage, of the analysis toad. A dietary hierarchy (DH) 1 Corresponding Author. E-mail: quiroga_lore@ was obtained by adding the IRI of each order yahoo.com.ar contribute, and then the percentage was calculated Downloaded From: https://bioone.org/journals/Journal-of-Herpetology on 12 May 2020 Terms of Use: https://bioone.org/terms-of-use Access provided by Bing Search Engine 312 SHORTER COMMUNICATIONS TABLE 1. Diet composition of Rhinella arenarum by size class. Abbreviations: N, number of prey; IRI, index of relative importance; DH, dietary hierarchy; Ni, not identified. The sum of the IRI for each taxonomic category is in bold. Adults Subadult Prey category N IRI DH N IRI DH VERTEBRATA Squamata Leptotyphlopidae 2 0.08 0.004 Rodentia Mus musculus 1 0.34 0.02 INSECTA Coleoptera 612.92 36.4 155.52 5.50 Curculionidae 459 231.90 6 67.22 Coccinellidae 23 1.24 Cicindelidae 17 3.44 Cerambycidae 2 0.12 Carabidae 302 343.55 5 35.07 Elateridae 50 7.82 Lampyridae 7 0.31 Larvae 6 0.50 3 3.01 Meloidae 2 0.03 Scarabaeidae 54 23.49 2 48.86 Scolytidae 1 0.01 Ni 7 0.51 2 1.36 Orthoptera 11.17 0.6 10.9 0.38 Gryllotalpidae S. borellii 1 0.02 Acrididae 4 0.58 1 5.35 Gryllidae 17 10.57 1 4.74 Thysanoptera Thrips 2 1.27 0.04 Blattaria 0.22 0.01 Blattidae 2 0.10 Ootheca-P. americana 1 0.01 Philodromidae 2 0.11 Diptera 0.74 0.04 21.84 0.77 Larvae 1 0.01 Culicidae 31 0.71 16 19.14 Calliphoridae Calliphora sp. 1 0.01 4 2.40 Ni 2 0.01 1 0.30 Dictyoptera Mantidae 1 0.32 0.01 Hemiptera 1.29 0.07 5.17 0.18 Belostomatidae 1 0.04 1 1.51 Notonectidae 1 0.07 Pentatomidae 5 0.58 Ni 4 0.09 4 3.66 Homoptera 11 0.51 0.03 27 7.97 0.28 Hymenoptera 1,679.66 100 2,824.15 100 Pupae 1 0.01 Apidae 7 1.27 Formicidae 5,248 1,677.53 99.87 455 2,823.56 99.97 Sphecidae 1 0.02 Vespidae 13 0.83 1 0.59 Isoptera 260 2.72 0.1 2 0.59 0.02 Lepidoptera 0.74 0.04 0.89 0.03 Ni 4 0.04 1 0.30 Larvae 5 0.44 2 0.59 Pupae 5 0.26 ARACHNIDA Acarii 1 0.002 0.0001 29 50.99 1.80 Downloaded From: https://bioone.org/journals/Journal-of-Herpetology on 12 May 2020 Terms of Use: https://bioone.org/terms-of-use Access provided by Bing Search Engine SHORTER COMMUNICATIONS 313 TABLE 1. Continued. Adults Subadult Prey category N IRI DH N IRI DH Aranae 6.79 0.4 Aracnidae 36 6.76 1 0.62 0.02 Solifugae 1 0.03 CRUSTACEA Isopoda 1,002 349.30 18 56 178.55 6.32 ANNELIDA Oligochaeta Haplotaxidae 2 0.01 0.0005 MYRIAPODA 2 0.01 0.0005 1 0.34 0.01 OTHERS 2 0.88 0.05 TOTAL 7,608 623 that contributed each order to the diet.
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