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Isotopic Niche Overlap Between the Invasive Leiothrix and Potential Native Competitors

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Isotopic Niche Overlap Between the Invasive Leiothrix and Potential Native Competitors Animal Biodiversity and Conservation 41.2 (2018) 427 Isotopic niche overlap between the invasive leiothrix and potential native competitors E. Pagani–Núñez, M. Renom, C. Furquet, J. Rodríguez, F. Llimona, J. C. Senar Pagani–Núñez, E., Renom, M., Furquet, C., Rodríguez, J., Llimona, F., Senar, J. C., 2018. Isotopic niche overlap between the invasive leiothrix and potential native competitors. Animal Biodiversity and Conservation, 41.2: 427–434, Doi: https://doi.org/10.32800/abc.2018.41.0427 Abstract Isotopic niche overlap between the invasive leiothrix and potential native competitors. We analysed niche overlap between the red–billed leiothrix Leiothrix lutea, a spreading exotic invasive bird, and the European robin Erithacus rubecula and the blackcap Sylvia atricapilla, similar native species, which are commonly considered as potential competitors in Mediterranean forests. We analyzed stable isotope ratios (δ13C and δ15N) from feathers of birds sampled in several locations within the Collserola mountain range (Barcelona, NE Spain) where leiothrix have strongly increased their numbers in the last decade and quantified niche overlap with nicheROVER. Blackcap individuals showed the lowest probability to be found in the isotopic niche of the other two species (around 60 %), while leiothrix and robins showed a high probability to share the same isotopic niche (82 %). Our results stress that competition for resources is potentially high and the species shared marked niche asymmetries, which may have implications for community dynamics in the long term. Key words: Interspecific competition, Niche overlap, Niche asymmetries, Red–billed leiothrix, Stable isotopes Resumen Solapamiento de nichos isotópicos entre el ruiseñor del Japón, invasivo, y los posibles competidores nativos. Ana- lizamos el solapamiento de nichos entre el ruiseñor del Japón, Leiothrix lutea, que es un ave exótica invasiva en expansión, y el petirrojo europeo, Erithacus rubecula, y la curruca capirotada, Sylvia atricapilla, que son especies nativas parecidas que se suelen considerar competidores potenciales en los bosques mediterráneos. Analizamos la relación de isótopos (δ13C y δ15N) en las plumas de las aves capturadas en varios lugares dentro de la cordillera de Collserola (Barcelona, NE España) en los que la población de ruiseñor del Japón ha aumentado notablemente en el último decenio, y cuantificamos el solapamiento de nichos con nicheROVER. Los individuos de curruca presentaron la menor probabilidad de encontrarse en el nicho isotópico de las otras dos especies (alrededor del 60 %), mientras que el ruiseñor y el petirrojo mostraron una elevada probabilidad de compartir el mismo nicho isotópico (el 82 %). Nuestros resultados destacan que la competencia por los recursos puede ser elevada y que las especies comparten marcadas asimetrías de nichos, lo que puede tener consecuencias para la dinámica de comunidades a largo plazo. Palabras clave: Competencia interespecífica, Solapamiento de nichos, Asimetrías de nichos, Ruiseñor del Japón, Isótopos estables Received: 29 VI 18; Conditional acceptance: 30 VII 18; Final acceptance: 05 IX 18 Emilio Pagani–Núñez, State Key Lab. of Biocontrol, Department of Ecology/School of Life Sciences, Sun Yat–sen University, No.135 West Xingang Road, 510275 Guangzhou, People's Republic of China.– María Renom, Juan Carlos Senar, Museu de Ciències Naturals de Barcelona, Psg. Picasso s/n., 08003 Barcelona, Spain.– Carles Furquet, Institut Català d’Ornitologia (ICO), Museu de Ciències Naturals de Barcelona, Psg. Picasso s/n., 08003 Barcelona, Spain.– Jordi Rodríguez, Grup d’Anellament Parus, Institut Català d’Ornitologia (ICO), c/ Salze 36, 08186 Lliçà d’Amunt, Barcelona, Spain.– Francesc Llimona, Estació Biològica Can Balasc, Consorci del Parc Natural de Collserola, ctra. de l’Església 92, 08017 Barcelona, Spain. Corresponding author: Emilio Pagani–Núñez. E–mail: [email protected] ISSN: 1578–665 X © 2018 Museu de Ciències Naturals de Barcelona eISSN: 2014–928 X 428 Pagani–Núñez et al. Introduction servation policies in the areas invaded by leiothrix. The aim of our work was therefore to shed new Invasive species are a major threats to biodiversity light on this controversy by using stable isotopes (Lockwood et al., 2007; Kolar and Lodge, 2001) but as an alternative approach. Trophic interactions they also offer an opportunity to study ecological and are fundamental to study the impact of exotic on evolutionary processes (Mooney and Cleland, 2001). native species (David et al., 2017). Comparing the A topic of current interest is the process by which niches of invasive and native species in the invasion animals become successfully established in a newly distribution range has been suggested as a good invaded area and how they accommodate within the approach to assess the occurrence of interspecific host community (Marsico et al., 2010; Henriksson competition (Cornell and Lawton, 1992). Analyses et al., 2016; Sol and Maspons, 2016; Duncan et al., using stable isotopes ratios of δ13C and δ15N, 2003). Two main hypotheses have been suggested: which are mostly indicative of habitat use and diet the competition hypothesis and the opportunistic respectively (Pagani–Núñez et al., 2017), can help hypothesiss (Duncan et al., 2003). The competition understand niche overlap over longer time spans hypothesis suggests that the main factor facilitating than behavioural observations (Pagani–Núñez et successful colonization of novel environments is the al., 2017; Inger and Bearhop, 2008; Layman et al., superior competitive ability of the invader over native 2012). Moreover, stable isotopes are commonly species (Duncan et al., 2003; Petren and Case, 1996; used to assess interspecific competition (see e.g. Duncan, 1997). This assumes that invasive and native Karlson et al., 2015; Smith et al., 2017). This is why species show high niche overlap, which may drive it has been suggested as a powerful tool to study intense competition. The opportunistic hypothesis, niche overlap between native and invasive species however, proposes that the invading species does not (Jackson et al., 2012; Kamenova et al., 2017). compete with natives, but simply takes advantage of In this paper we compared the isotopic niche of the niche opportunities not used by native species (Dun- leiothrix with that of the European robin (hereinafter can et al., 2003; Shea and Chesson, 2002; Batalha robin) and the blackcap, which have been identified et al., 2013). However, it has been difficult to find as the two main native competitors of leiothrix in clear empirical support for either of these hypotheses the Mediterranean area (Pereira et al., 2017). We (Duncan et al., 2003). assessed niche overlap among these species using The red–billed leiothrix Leiothrix lutea (hereinafter nicheROVER, a new statistical tool developed by leiothrix) exemplifies this controversy. A worldwide– Swanson et al. (2015). This allowed us to contrast scale meta–analysis of avian exotic introductions the competition and niche opportunity hypotheses in identified the leiothrix as one of the bird species with wild conditions and using the most advanced research the highest negative local impact on bird communities techniques. (Martin–Albarracin et al., 2015). This species has successfully invaded several regions, from Japan to Hawaii and SW Europe (Herrando et al., 2010; Material and methods Male et al., 1998; Tojo and Nakamura, 2004; Pereira et al., 2017), while it is native to South Asia (from This study was carried out in the Natural Park of West Himalaya to South and Central China) (Collar Collserola (45° 27' N, 2° 8' E; Catalonia, northeast et al., 2018). A recent work in the Iberian Peninsula, Spain). Our study area may be characterized as based on behavioural observations during foraging Mediterranean mixed forest (Pagani–Núñez et al., and on habitat use, concluded that leiothrix exhibited 2014). The three main tree species were Aleppo pine little niche overlap with most native species, and that Pinus halepensis, holm oak Quercus ilex and the oak this invasion had relatively few consequences for the Q. cerrioides. The first leiothrix was captured in this populations of the other species (Vall–llosera et al., area on 14th July 1990 (La Espinagosa, Vallvidrera; 2016). As a consequence, they suggested that the C. Jordà, J. L. Copete and J. C. Senar, pers. obs.). success of leiothrix in this area could be explained by Since then, the species has spread exponentially the opportunistic hypothesis. However, another two along the whole park, across forest and riverside works, also conducted in Iberia, used morphology, zones, now reaching the close mountain range of diet and exploratory behaviour to show a competitive Serra de Marina (Herrando et al., 2010). We used advantage of leiothrix over native species rather than data from field surveys to illustrate and quantify this an opportunistic occupation of an empty ecological pattern (see below). niche (Pereira et al., 2017). Behavioural observations The study is based on 67 leiothrix, 43 robins and also supported interference competition by an active 20 blackcaps captured in the area using mist nets displacement of native species such as European and funnel traps (Senar et al., 1997) between January robins Erithacus rubecula and blackcaps Sylvia and August 2013. We took the 5th most external tail atricapilla through aggressive attacks of leiothrix feather of the right side of all individuals for laboratory (Pereira et al., 2018). analyses. Given the long
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