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Lizards and the City: a Community Study of Lacertidae and Gekkonidae from an Archaeological Park in Rome

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Lizards and the City: a Community Study of Lacertidae and Gekkonidae from an Archaeological Park in Rome Zoologischer Anzeiger 283 (2019) 20e26 Contents lists available at ScienceDirect Zoologischer Anzeiger journal homepage: www.elsevier.com/locate/jcz Research paper Lizards and the city: A community study of Lacertidae and Gekkonidae from an archaeological park in Rome * Simbula Giulia a, Luiselli Luca b, Vignoli Leonardo a, a Dipartimento di Scienze, Universita Roma Tre. Viale G. Marconi 446, 00146 Rome, Italy b IDECC Institute for Development, Ecology, Conservation and Cooperation, via Giuseppe Tomasi di Lampedusa 33, I 00144 Rome, Italy article info abstract Article history: The study of animal communities is mainly concerned with how the groupings of species are distributed Received 20 December 2018 in nature and the ways in which the members of these communities assemble and interact with each Received in revised form other. Pianka suggested that animals partition environmental resources in three basic ways: temporally, 18 July 2019 spatially, and trophically. Such differences in activities separate the niches, reduce competition, and Accepted 2 August 2019 presumably allow the coexistence of a variety of species in the communities. The present work compares Available online 9 August 2019 the spatial, trophic and temporal niches of four reptile species, two geckos (Tarentola mauritanica and Corresponding Editor: Alexander Kupfer Hemidactylus turcicus) and two lizards (Podarcis muralis and Podarcis siculus) living in a same roman-age archeological park situated area inside Rome metropolitan area (central Italy), and sharing the same two- Keywords: dimensional habitat (the roman aqueduct walls). The results showed an overall overlap among species Urban ecology higher than expected by chance for all considered ecological dimensions except for the spatial resource Lizards related to the vertical position on the wall. Surprisingly, despite living in a two-dimension habitat would Niche dimensions expect an increase of interspecific interaction rate, no partition was observed between species with the Overlap most convergent ecological requirements. Although our hypotheses about lizard communities struc- Competition turing and interactions in the urban habitat require further investigation, we think that our lizard community is based more on the ecological needs of each species rather than on species’ interactions. The observed spatial segregation between diurnal geckos and lizards would not support alone the interaction hypothesis. We speculated that the coexistence of ecologically overlapping species at the study area is allowed by a non-limiting availability of resources (above all food) that would result in the reduction of heterospecific competitive interactions and an overall wide overlap of resource use. © 2019 Elsevier GmbH. All rights reserved. 1. Introduction that allowed co-existence by partitioning one or more niche di- mensions (Vitt & Zani 1998; Chase & Leibold 2003; Grbac & Brnin An animal ecological niche is a multifactorial concept that can 2006; De Pinho et al. 2009). Resource partitioning strategies, as a be subdivided into several dimensions (Pianka 1974; Schoener response to past pressures, have been documented in many co- 1974a, 1982). Differences in the ecological and physiological re- existing, ecologically similar animal species as a general mecha- quirements of an individual will determine its position in the nism to mitigate or avoid competition (Chase & Leibold 2003; different spatial and temporal components of a niche. Therefore, Luiselli 2006). Indeed, plasticity in resource use may be an impor- the co-existence of species may result in an overlap in optimal tant characteristic in the process of co-evolution in sympatric habitat requirements (i.e. spatial niche) and represents one of the populations and enables flexibility in spatial, trophic or temporal most common forms of species interactions (Schoener 1974b). To niches, thus allowing successful coexistence (Lisicic et al. 2012). overcome the potential negative effects of overlapping exploitation During the last twenty years, interest in herpetological com- of limiting resources, syntopic species may have evolved strategies munities has grown enormously and several meta-analyses have been published (Luiselli 2006, 2008; Vignoli & Luiselli 2012; Vignoli et al. 2017). Even concerning the urban habitats, several datasets on * their herpetological communities have been published (Germaine Corresponding author. & E-mail addresses: [email protected] (S. Giulia), [email protected] Wakeling 2001; Jellinek et al. 2004; Perry et al. 2008), (L. Luca), [email protected] (V. Leonardo). including in Mediterranean towns (Luiselli & Capizzi 1999; Rugiero https://doi.org/10.1016/j.jcz.2019.08.001 0044-5231/© 2019 Elsevier GmbH. All rights reserved. S. Giulia et al. / Zoologischer Anzeiger 283 (2019) 20e26 21 & Luiselli 2006, 2007; Vignoli et al. 2009). Urban habitats suitable for herpetofauna are, usually, severely fragmented and present a number of important environmental alterations: natural vegetation is reduced and its structure simplified; food resource availability decrease in variety and is often spatially concentrated; disturbance from humans increases; the community of predators changes and pollution is more frequent with regards to night-time lights, acoustic noise and chemicals. Nevertheless, a few species seem not only to be little affected by the urbanization process, but appear to perceive urban habitats as ecological opportunities, and proliferate and expand their range by showing altered behaviour or life- history strategies in urban areas and thrive (Ditchkoff et al. 2006; Sol et al. 2013). Mortality rates of urban dwelling species may be elevated because of altered predator communities (e.g., dogs and cats) and road killing. On the other hand, habitats with drastic anthropogenic changes might harbour synanthropic generalist species with broad preferences ecologically associated with anthropogenically modified environments and inefficient preda- tors (Tyler et al. 2016). Predation pressures, disturbance intensity, and trophic dynamics in urban areas are indeed complex and may keep populations at lower number than the environmental carry capacity, thus reducing the effect of potential competition due to the exploitation of shared resource. In this paper we compare the spatial, trophic and temporal niches of four syntopic reptile species similar in size, two geckos (Tarentola mauritanica, Linnaeus, 1758, and Hemidactylus turcicus Fig. 1. General landscape of the study area, showing the Roman age aqueduct, and the Linnaeus, 1758) and two congeneric lizards (Podarcis muralis, four main habitat. Symbols: Rv ¼ wall with climbing vegetation; Re ¼ exposed wall Laurenti, 1768 and Podarcis siculus, Rafinesque, 1810) inhabiting a without climbing vegetation; Rf ¼ walls with some crevices available; F ¼ ruined spots very simplified urban habitat (bidimensional brick wall) in a with high abundance of crevices in the wall. roman-age archeological park situated inside Rome metropolitan area (central Italy). For convenience, we will use verbal shorthand and speak of the food, time and spatial niches rather than refer to protocol involved to capture the individuals following a transect “the trophic, temporal and spatial dimensions of the niche,” as along the walls of the Felice aqueduct, always starting from Tor proposed by Pianka (1973). Fiscale park(total sampled wall length, 650 m), and continuing for In the present paper we address the following questions: i) Are another 800 m inside the Acquedotti park. Sites were revisited over there any overlaps along the three niche dimensions among the 3 days in each season to ensure that a high proportion of the in- study species? ii) If so, is there any resource partitioning among the dividuals were sampled. The animals were caught by noose during species in order to allow co-existence? Due to the very simplified daylight and by hand during night hours. For each sampling activity habitat structure (i.e. low spatial heterogeneity) shared by the we registered date, time, weather conditions and air temperature, study species, we should expect a lower overlap of resource use by whereas for each collected animal we recorded species, sex, snout- the species on the other main niche dimensions. However, in a vent length (SVL), weight (W), height above ground at first sight same way, the simplified habitat should reduce the prey availability and microhabitat. After being caught, lizards were placed in plastic for the species, thus a strong partition of the food spectrum is boxes, each individual was marked by means of non-toxic marker unlikely. Therefore, we cannot predict how the coexisting species pen of different colours for short time identification, while digital exploit the three main dimension of the ecological niche. photographs of ventral and dorsal sides were taken for long-time identification by examining the number and position of scales 2. Materials and methods (Sacchi et al. 2010). We were able to individually identify most of the lizards and geckos just by using the pen marking due to the high 2.1. Study area frequency of sampling that prevent moulting events to fade the colour marker. When we caught a non-marked individual we used Fieldwork was carried out in two contiguous urban green areas, the photographic database to assess if it was a new or already with Roman age monuments, situated in the city of Rome: Tor captured individual. Once they produced faeces for food niche Fiscale (about 3 ha, 4151030.8100N,
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