Diet of Salvator Merianae (Squamata: Teiidae): New Prey Item and Review of Predation Records
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Use the following type of citation: North-western Journal of Zoology 2021: e217504 *Handling editor: Yurii Kornilev *Manuscript Domain: Herpetology of the Americas *Manuscript code: NwJZ_20_HA_07 *Submission date: 19_04_ 2020 *Revised: 14_06_2021 *Accepted / Rejected: 01_08_2021 *No. of words (without abstract, acknowledgement, references, tables, captions): 1460 Title of the paper: Diet of Salvator merianae (Squamata: Teiidae): New prey item and review of predation records Running head: New prey item and a diet review of Salvator merianae Authors (First LAST - without institution name!): Heithor Simão DINIZ;Zoology Renato Neves FEIO; Clodoaldo of Lopes DE ASSIS; Jhonny José Magalhães GUEDES proofing Key Words (at least five keywords): Atlantic Forest, Dipsas, feeding behavior, prey-predator Journal interaction, Tegu lizard until No. of Tables: 1 No. of Figures: 0 paper No. of Files (landscape tables should be in separate file): 1 North-western Accepted Use the following type of citation: North-western Journal of Zoology 2021: e217504 1 Diet of Salvator merianae (Squamata: Teiidae): New prey 2 item and review of predation records 3 Heithor Simão DINIZ1,*, Renato Neves FEIO1, Clodoaldo Lopes DE ASSIS1 and Jhonny José 4 Magalhães GUEDES2 5 1. Departamento de Biologia Animal, Museu de Zoologia João Moojen, Universidade Federal 6 de Viçosa, Viçosa, Minas Gerais 36570-900, Brazil. 7 2. Programa de Pós-Graduação em Ecologia e Evolução, Departamento de Ecologia, Campus 8 Samambaia – Universidade Federal de Goiás, Goiânia, Goiás, 74001-970, Brazil. 9 *Corresponding author, H. S. Diniz, E-mail: [email protected] 10 Running title: Zoology 11 Abstract of 12 The Tegu lizard Salvator merianae is a large species proofingwith a wide geographic distribution 13 in South America. Regarding its feeding habits, this lizard is omnivorous, generalist and Journaluntil 14 opportunistic, preying upon a wide variety of food items, such as fruits and seeds, 15 invertebrates and vertebrates. In this study,paper we present the first predation event by the Tegu 16 on a snake from the genus Dipsas, which occurred in the state of Minas Gerais, Brazil. We 17 also performed a comprehensiveNorth-western literature review, compiling data on published food items of 18 S. merianae. Here we demonstrateAccepted the diet of the Tegu lizard is dominated by plants, followed 19 by other reptiles, invertebrates, mammals, birds, and amphibians. Also, we discuss about the 20 generalist and opportunistic diet of S. merianae and its role in the ecosystem. 21 Key words: Atlantic Forest, Dipsas, feeding behavior, prey-predator interaction, Tegu 22 lizard. 23 24 The Tegu lizard Salvator merianae Duméril & Bibron, 1839 (Squamata: Teiidae) has 25 a wide geographical distribution, occurring from northern South America to northern 26 Argentine Patagonia (Barreto-Lima & Camilotti 2009). Recently, the species has been Use the following type of citation: North-western Journal of Zoology 2021: e217504 27 introduced in the USA, in the southern states of Florida and Georgia (Mazzotti et al. 2015, 28 Wood et al. 2018, Haro et al. 2020). It is found throughout most of Brazil (Costa & Bérnils 29 2018), inhabiting natural environments, such as open areas, clearings and forest edges (Avila- 30 Pires 1995), as well as urban areas (e. g., Costa et al. 2009, Guedes et al. 2017). Salvator 31 merianae is one of the largest Neotropical lizards, reaching up to 1.6 m in length and 32 weighing up to 5 kg (Fitzgerald 1992), which makes it an attractive food source for humans in 33 many regions (Avila-Pires 1995, Caldironi & Manes 2006). It is an active generalist and 34 omnivore, consuming a wide variety of invertebrates and vertebrates, carcasses, eggs, 35 mushrooms and plants (Kiefer & Sazima 2002). Despite the diverse known food items in the 36 diet of this species, records of new prey species are still being reported, indicating the 37 existence of knowledge gaps about its diet. Here, we i) reportZoology a predation on a new snake of 38 species of the genus Dipsas (Squamata: Dipsadidae) by S. merianae in the Atlantic Forest proofing 39 from the southeastern region of Minas Gerais State, and ii) perform a much needed literature Journal 40 review on the diet of this species, which data are sountil far scattered throughout the literature. 41 On 30 December 2018 at 12:27 h, one of us (JJMG) witnessed a predation event of a 42 Brazilian Snail-eating snake (Dipsas sp.)paper by the Tegu (S. merianae) in the district of Barretos, 43 a rural area in the municipality of Alvinópolis, state of Minas Gerais, Brazil (20.05940S, 44 43.19308W; datum North-westernWGS84; 680 m a.s.l.). The Tegu was found digging a pre-existing hole in Accepted 45 a garden; after two minutes of digging, the lizard took the snake out of the soil by holding it in 46 its mouth. Prey subjugation began about three minutes after the snake was removed from its 47 refugium, with the lizard biting and shaking its prey laterally. The snake did not offer any 48 apparent resistance to its predator, being ingested head first. 49 We did not disturb or capture the individuals during the predation event. Species 50 identification was done based on color pattern and geographic distribution, and was 51 confirmed by Dr. Henrique Caldeira Costa, a specialist in reptile taxonomy. A video Use the following type of citation: North-western Journal of Zoology 2021: e217504 52 recording is available online at the audiovisual collection of the Fonoteca Neotropical Jacques 53 Vielliard (http://www2.ib.unicamp.br/fnjv/; ZUEC-VID 784). 54 A comprehensive literature review was made about the diet of S. merianae. We 55 searched through peer-reviewed articles, short notes and books available at online repositories 56 such as Google Scholar, Web of Science, Scopus, and websites such as ResearchGate and 57 Academia. We used the following keywords: “Tupinambis” (old genus name; see Harvey 58 2012, Murphy 2016) OR “Salvator” AND “merianae” AND “diet” OR “predat*” OR “prey*” 59 OR “food”. We also checked the references in the pertinent literature found to look for 60 possible reports missing from the online searches. Thus, we produced a list of published diet 61 items. Although more than one publication may have listed the same food item, we decided to 62 keep the review qualitative and to cite only the first mention Zoologyof a given food item. of 63 The literature review about the diet of S. merianae produced 43 publications with proofing 64 distinct records and 174 food items, distributed in 56 orders and 93 families, including plants, Journal 65 animals and fungi (Table 1). Most food items consisteduntil of plant contents (n = 70 records; 66 40.2%), followed by reptiles (n = 39; 22.4%), invertebrates (n = 27; 15.5%), mammals (n = 67 13; 7.5%), birds (n = 12; 6.9%), amphibianspaper (n = 11; 6.3%), fish (n = 1; 0.6%) and fungi (n = 68 1; 0.6%) (see Table 1). 69 Herein, we presentNorth-western the twentieth snake species as food item of S. merianae and the Accepted 70 first predation record upon a snake from the genus Dipsas (formerly Sibynomorphus; see 71 Arteaga et al. 2018). From all known defensive behaviors ascribed to the species of this 72 genus, a very common strategy is mimicking venomous snakes, such as Bothrops spp. 73 (Dell’Aglio et al. 2012). This snake species has a variegated color pattern and when 74 threatened displays a S-shaped posture and triangulation of the head. Other snakes in the diet 75 of S. merianae such as Xenodon neuwiedii also display a similar behavior and mimic viperids 76 (Tozetti et al. 2009). These data suggest that mimicking venomous snakes may not be entirely 77 effective against some predators (Guimarães & Sawaya 2011). It is worth mentioning that, so Use the following type of citation: North-western Journal of Zoology 2021: e217504 78 far, there is no record of viperids as prey of S. merianae in natural conditions. However, a 79 predation upon Bothrops sp. has been reported in captivity (Kasperoviczus et al. 2015). Also, 80 Silva et al. (2014), based on local people knowledge, presented a list with several tentative 81 items in the Tegu’s diet, including Bothrops spp. and Crotalus durissus. This finding indicates 82 that a predation event on venomous snakes by the Tegu could indeed occur in nature. The 83 tendency of generalist predators is to eat what they see first when dominant food sources are 84 not available (Ballinger & Ballinger 1979). Thus, we hypothesize that under stressfull 85 conditions (i.e., lack of food) the Tegu might risk attempting predation on viperid snakes. 86 The diversity of food items observed in the diet of S. merianae highlights its highly 87 generalist eating habits, including opportunistic necrophagy (Kiefer & Sazima 2002, Machado 88 2007, Sazima & D'Angelo 2013). It is worth mentioning thatZoology the Tegu has a high voracity, of 89 being able to neutralize prey with the same, or even larger, body size (Vieira et al. 2018), in proofing 90 which the benefits of predation may be surpassed by the energy expenditure and risk of Journal 91 injuries. The diet and foraging behavior of this speciesuntil indicate a possible advantage of the 92 Tegu to adapt to different environmental conditions, and hence, to a rapid changing 93 landscape. Indeed, the recent establishmentpaper and spread of this species in the states of Florida 94 and Georgia, USA, seems to corroborate that (Kaiser et al. 2013, Wood et al. 2018, Haro et al. 95 2020). The invasionNorth-western by the large, generalist , and opportunistic S. merianae into new habitats Accepted 96 warrants attention because of the potential severe harmful effects upon a large proportion the 97 native fauna (Jarnevich et al.