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Enhanced Interspecific Territoriality and the Invasion Success of The Biol Invasions (2010) 12:865–874 DOI 10.1007/s10530-009-9507-3 ORIGINAL PAPER Enhanced interspecific territoriality and the invasion success of the spotted tilapia (Tilapia mariae) in South Florida Wesley R. Brooks Æ Rebecca C. Jordan Received: 15 August 2008 / Accepted: 19 June 2009 / Published online: 27 June 2009 Ó Springer Science+Business Media B.V. 2009 Abstract South Florida’s freshwaters are amongst behaviors of T. mariae are an important component the most invaded in the world with 34 naturalized fish of their success in South Florida. The continued spread species. How these non-natives affect the local native of T. mariae populations throughout South Florida into fish populations, however, is largely unknown. Native natural habitats suggests an increasing potential to sunfish of the genus Lepomis are important as preda- affect the quality of spawning habitat available for tors in structuring fish and invertebrate assemblages in Lepomis sunfishes and warrants a renewed focus on the swamps and seasonal wet prairies of the Big T. mariae as a non-native species of special concern. Cypress Swamp and Florida Everglades. The spotted tilapia, Tilapia mariae, is a successful West African Keywords Big cypress swamp Á Competitive invader that exhibits territorial and spawning behavior interactions Á Fish invasions Á Florida Everglades Á that closely matches that of native Lepomis sunfishes. Interspecific competition Á Lepomis sunfish Á We tested the hypothesis that Lepomis sunfishes and South Florida Á Spotted tilapia Á Territorial behavior T. mariae would compete when space was limiting. Additionally, we predicted that T. mariae, because of their aggressiveness, would be more successful in Introduction acquiring space. We collected juveniles of both groups from Big Cypress National Preserve, Everglades South Florida has experienced a high rate of non- National Park, and the South Florida Water Manage- native freshwater fish introductions since the 1960’s ment District canal system for laboratory trials in (Courtenay and Robins 1975; Shafland 1996a, b), with which likely competitive interactions were staged and 34 species categorized as having self-sustaining pop- observed. T. mariae were bolder and more aggressive ulations (Shafland et al. 2008). This is likely due in part than Lepomis sunfishes. T. mariae residents resisted all to low species richness of the native fish community intruders whereas 30% of Lepomis sunfish residents (Moyle and Light 1996; Gido and Brown 1999), as were ejected. We surmise that these enhanced well as vast anthropogenic alterations of the region’s hydrology and aquatic habitats (Moyle and Light 1996; Marchetti et al. 2004). Attempts to prevent such W. R. Brooks (&) Á R. C. Jordan introductions and manage non-native populations that Department of Ecology, Evolution and Natural Resources, become naturalized are often limited by insufficient Rutgers University, 14 College Farm Rd, ENR Bldg, biological knowledge and a lack of environmental and School of Environmental and Biological Sciences, New Brunswick, NJ 08901, USA economic impact data (Simberloff et al. 2005). While e-mail: [email protected] evidence for deleterious ecological effects of 123 866 W. R. Brooks, R. C. Jordan non-native fishes on native aquatic communities has and Spotted Sunfish (L. punctatus) are territorial been documented in many watersheds (Taylor et al. substrate spawners (Loftus and Kushlan 1987). 1984), there are no experimental data from South Additionally, T. mariae and Lepomis sunfishes over- Florida that demonstrate strong negative interactions lap in spawning season (Loftus and Kushlan 1987), by non-native fishes on native fish populations or and have been observed nesting in close proximity to communities (Shafland 1996b; Courtenay 1997; Fuller each other, interacting via displays and chases (Hogg et al. 1999; Trexler et al. 2000). A few qualitative 1974; Annett et al. 1999; Trexler et al. 2000). This assessments of the region’s fish communities have suggests a strong potential for territorial competition reported localized shifts in species composition and that, to date, has not been empirically tested. abundance following the establishment of non-native T. mariae individuals are aggressive and territorial fishes (Courtenay and Hensley 1979; Kushlan 1986; towards conspecifics and interspecifics throughout Loftus and Kushlan 1987). These reports, however, are their native range (King and Etim 2004). Interspecific speculative in nature and likely are more indicative of aggression and territoriality by non-natives can fish communities in the urban canals of Southeast contribute to invasion success (Holway and Suarez Florida (see Shafland 1999) as opposed to communi- 1999; Moyle and Marchetti 2006), and lead to the ties in natural wetlands (Trexler et al. 2000). displacement of natives from preferred microhabitats The Spotted Tilapia, Tilapia mariae, has received such as spawning sites (Taylor et al. 1984). These particular attention in South Florida as a non-native fish spatial alterations can impact individual growth rates of high concern (Shafland 1976; Courtenay and Hensley and population densities leading to a host of direct 1979). Native to lowland rivers of tropical West Africa and indirect ecological effects in the native commu- (Ikomi and Jessa 2003), this algivore/detritivore was nity (Taylor et al. 1984). Moreover, habitat structure likely introduced through escape from aquaculture into can play an important role in determining competi- Miami’s urban canal system in the early 1970’s (Hogg tion intensity between native and non-native com- 1974). Within a few years of introduction, T. mariae petitors (Petren and Case 1998). Specifically, habitat replaced another introduced cichlid, the Black Acara structure may affect the effort that one or both (Cichlasoma bimaculatus), as the most abundant fish by competitors will invest in the acquisition or defense biomass in urban canals (Courtenay and Hensley 1979). of a territory (Johnsson et al. 2000). Consequently, T. mariae has expanded its range via the canal system the scope of the potential impacts of T. mariae on throughout South Florida and is now routinely captured Lepomis sunfishes might be limited to certain habitats in other artificial wetlands including restoration and not applicable across all of their shared range. marshes, rice fields, agricultural ditches (Chimney and We directly tested the notion that non-native Jordan 2008), as well as natural wetlands in the T. mariae and native Lepomis sunfishes will actively Everglades and Big Cypress regions including fresh- compete for space. More specifically, we tested the water marsh (Chick et al. 2004), wet prairie (Trexler hypothesis that individuals of T. mariae would be et al. 2000; Chick et al. 2004), slough (Chick et al. more aggressive towards interspecifics than their 2004), and cypress swamp habitats (Chimney and Lepomis sunfish competitors resulting in enhanced Jordan 2008). In Everglades National Park, where it has territorial acquisition and defense. We also expected most recently invaded, T. mariae already constitutes 4.5 that different habitat structures might have a signif- and 3.1%, respectively, of the abundance of the large icant effect on the potential outcomes of competitive fish catch in Shark River and Taylor sloughs, and has interactions, with one or both groups exhibiting replaced the Mayan Cichlid, Cichlasoma urophthal- modified behavior in each habitat type. mus, as the most abundant non-native species in the former (Chick et al. 2004). Courtenay and Hensley (1979) suggested that Methods T. mariae was likely to impact native fishes, especially Lepomis sunfishes and other Centrarchids, Collection and housing through competition for space and spawning habitat. Like T. mariae, the Lepomis sunfishes including the Juvenile Lepomis sunfishes (L. gulosus, L. macrochi- Warmouth (L. gulosus), Bluegill (L. macrochirus), rus mysticalis, and L. punctatus punctatus) and 123 Enhanced interspecific territoriality 867 T. mariae cichlids, measuring 3–6 cm total length to experimentation in order to establish residency. The (TL) were collected from fifteen sites throughout remaining pool of unused individuals were designated South Florida within a range of urban and natural as ‘intruders’ and were maintained in small same- areas (Fig. 1). Fish were transferred to New Jersey species groups to facilitate easier individual identifi- and conditioned to the laboratory environment. In the cation and capture for measurement and trial usage. laboratory, individuals were maintained in same- Residents were used in each of four consecutive trials species 110-l tanks maintained at 24–28°C, pH 7–8, which varied in terms of competition type (Interspe- and 12-h broad-spectrum light/dark cycle. The fish cific: TM versus LG/LM/LP, Intraspecific) and habitat were fed a combination of flake and stick foods three structure of the tank (microhabitat 1: sandy bottom times daily. Except for feeding, occasional care, and only, microhabitat 2: sandy bottom with small PVC tank maintenance, the fish were not disturbed during tube, microhabitat 3: sandy bottom with artificial the 3 months prior to experimentation. plants blocking swimming in the top half of the tank). Experimental tanks were 18 liters in volume with a Intraspecific trials were performed in order to provide bottom surface area of 0.1 m2. The tank bottoms were a baseline for behavioral data for both T. mariae and covered in sand to a depth of 2 cm and outfitted with the three Lepomis sunfish species utilized in
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