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Behavioral Constraints for the Spread of the Eastern Mosquitofish

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Behavioral Constraints for the Spread of the Eastern Mosquitofish Biol Invasions DOI 10.1007/s10530-007-9109-x ORIGINAL PAPER Behavioral constraints for the spread of the eastern mosquitofish, Gambusia holbrooki (Poeciliidae) Shireen D. Alemadi Æ David G. Jenkins Received: 23 February 2007 / Accepted: 14 March 2007 Ó Springer Science+Business Media B.V. 2007 Abstract Eastern mosquitofish (Gambusia holbro- Keywords Invasive Á Dispersal Á Habitat Á Sex Á oki) are native to the southeastern United States but Water depth Á Leaf litter notoriously invasive elsewhere, and are aggressive predators in ecosystems they inhabit. Information on dispersal behavior is needed to better understand Introduction mosquitofish spread upon introduction and potential means to mitigate that spread. We experimentally Many invasive species are introduced to novel tested the effects of shallow water depths (3–24 mm) regions by human actions (Rahel 2002), and spread and obstacles (leaf litter) on mosquitofish dispersal thereafter by species’ actions. Dispersal behavior is behavior, plus a range of conditions relevant to field critical to the spread of many actively-dispersing situations. Mosquitofish dispersed significantly faster species after introduction (Ehrlich 1986; Holway and in deeper water (p < 0.001) but some dispersed in Suarez 1999; Sakai et al. 2001; Rehage and Sih only 3 mm water depth (i.e., one-half average body 2004). Therefore, dispersal behavior may also be depth). Wetland and upland leaf litter at natural critical to the prevention or control of an invader’s densities strongly interfered with mosquitofish dis- spread. persal behavior. Based on our results, introduced The eastern mosquitofish (Gambusia holbrooki, mosquitofish spread rapidly given unimpeded dis- Poeciliidae) is native to the Southeastern United persal corridors (e.g., mowed ditches), and may do so States but has been widely introduced elsewhere; its at rates >800 m/day. Also, consistent lack of sexual congener, the western mosquitofish (Gambusia affi- dimorphism in dispersal behavior indicates that nis) is listed among the 100 worst invasive species in mosquitofish spread is not strongly dependent on the world (Lowe et al. 2000). Mosquitofish appar- female poeciliid reproductive biology. Our results ently disperse well following introduction to a new support designation of mosquitofish as highly inva- region (Arthington and Lloyd 1989) and have been sive and suggest that barriers to mosquitofish spread introduced to many wetlands outside their native must obstruct dispersal pathways as shallow as 3 mm range for the purpose of mosquito control, though depth. their efficacy for mosquito control has been ques- tioned (Lloyd et al. 1986, Leyse et al. 2004, Courtenay and Meffe 1989). Mosquitofish have & S. D. Alemadi Á D. G. Jenkins ( ) relatively brief generation times and live-bearing Department of Biology, University of Central Florida, Orlando, FL 32816-2368, USA females can store sperm from multiple males; one e-mail: [email protected] fertilized female may start a new population with 123 S. D. Alemadi, D. G. Jenkins little or no damaging founder effects (Chesser et al. Materials and methods 1984;Zaneetal.1999). Thus, mosquitofish potentially establish large populations after dispers- Fish were collected from Lake Claire and its neigh- ing into a new habitat, where they act as major boring pond *50 m to the west (hereafter Shireen’s predators that affect entire food webs (Meffe Pond) on the University of Central Florida (UCF) et al. 1983; Meffe 1985; Courtenay and Meffe Orlando campus. Lake Claire is a permanent sinkhole 1989; Gamradt and Kats 1996; Goodsell and Kats lake with a diverse fish assemblage, whereas 1999; Pyke 2005). Shireen’s Pond is shallow and may dry completely Despite the central role of dispersal behavior to the in drought years; no other fish species were observed understanding and control of invasive spread, too few during the primary study year. Fish were collected studies have examined dispersal ability alone with minnow traps (Aquatic Eco-Systems, Inc., (Rehage and Sih 2004) and relatively little is known Apopka, FL) set overnight, or by umbrella nets or of mosquitofish dispersal behavior. For example, a dip nets, and were transferred in pond water to the recent and comprehensive (26 pages, 261 citations) laboratory. Fish were then transferred to 45.5 l review of Gambusia biology (Pyke 2005) discussed aquaria (Nanotanks; Transworld Aquatic Enterprises, the subjects of dispersal, movement patterns and Inc., Los Angeles, CA) containing dechlorinated tap orientation in only 2 paragraphs and 12 citations. water with standard filtration and fish were fed Short-term behavioral experiments can help predict Tetramin fish flakes ad libitum at least 24 h before long-term population-level responses in natural sys- experiments. This protocol acclimated fish to labora- tems (Gilliam and Fraser 1987; Abrahams and Dill tory conditions and standardized hunger levels. 1989). For example, Rehage and Sih (2004) observed Experimental trials were conducted in twelve interspecific differences in schooling behavior and arenas (3 m long · 0.08 m wide · 0.10 m high), risk and dispersal behavior in four Gambusia species also filled with dechlorinated tap water. A fish was using experimental arenas. transferred by aquarium net into a holding zone We experimentally tested the effects of seven (0.12 m · 0.08 m · 0.10 m), which was partitioned variables on individual G. holbrooki dispersal behav- from the rest of the arena by a Plexiglas door. All ior: sex, hunger state, light, acclimation time, source doors had ten 4-mm drilled flow holes and hinged up habitat, water depth (3–24 mm), and the interaction and away from fish to open simultaneously during of depth · leaf litter type (wetland or upland). The dispersal trials. Each arena was drained after each first four variables were tested in preliminary exper- trial and refilled with dechlorinated tap water to iments and then standardized for more powerful remove any residual chemical cues left by previous subsequent experiments of the latter three variables. fish. Arenas were curtained to ensure that any human Female mosquitofish are larger than males: we movement around them did not affect fish behavior. predicted a priori that females would disperse faster Fish were timed with a stopwatch to the nearest than males. In addition, we expected that hungry fish 1/100 s, beginning when doors were opened until fish would disperse faster than satiated fish, and that fish reached a finish line 2.58 m away from the door. Fish would disperse faster in lighted conditions than in were observed in a mirror attached above the arenas dim light. We expected that fish collected from a to avoid behavioral responses to observer activity. shallow pond without other fish species would Fish were observed for 30 min; all fish that reached disperse faster than those from a permanent lake, the finish line were recorded for total time elapsed. because mosquitofish habituated to predatory fish Body height (mm), standard length (mm), and sex of may be more risk-averse (Rehage et al. 2005). each fish was recorded after a trial. Each fish was We also expected that net dispersal rate would observed only one time, so that individual fish be inversely proportional to water depth, because constituted individual replicates in all experiments. fish would seek greater depth. Finally, we Because each fish was free to meander within the expected that leaf litter would slow dispersal, but arena and because fish varied in size, data were fish would disperse faster through wetland litter expressed as both absolute net dispersal rate (m sÀ1) than upland litter, related to litter density and/or and relative net dispersal rate (body lengths sÀ1) for chemical cues. fish that completed a trial (i.e., reached the finish line 123 Behavioral constraints for the spread of the eastern mosquitofish within 30 min). Differences among treatments in net Lake Claire). Trials were run in 1 day to eliminate dispersal rates were analyzed by analysis of variance temporal variation, fish were randomized among (ANOVA) when log10-transformed values met para- arenas, and sex and fish dimensions were recorded metric assumptions, or by nonparametric methods as above. In addition, we repeated the habitat (Mann–Whitney or Kruskal–Wallis tests) when para- experiment approximately 1 year later; predatory fish metric assumptions could not be met. Some fish did (warmouth; Lepomis gulosus) were collected in not cross the finish line within 30 min, and the minnow traps in Shireen’s Pond in the second study number of those fish was compared among treatments year (but not the first), whereas Lake Claire is known by v2 analysis of contingency tables. All data were to support multiple fish species, including warmouth analyzed data using SPSS V. 11.5 (SPSS Inc., (L. gulosus) and largemouth bass (Micropterus sal- Chicago, IL, USA). Power analysis was conducted moides). Because fish may have responded to slightly with JMP 6.0 (SAS Institute, Cary, NC). different experimental conditions (and a different mix of investigators) between years, we analyzed years as Preliminary experiments an additional experimental variable. We hypothesized that the apparent colonization of predatory fish may Results of preliminary experiments were used to set alter behavior in Shireen’s Pond. Log-transformed parameters for subsequent experiments. In all pre- absolute and relative net dispersal rates were ana- liminary experiments, water depth was standardized lyzed by analysis of variance for habitat, sex, and to 6 cm (10 · average body height), and arenas were year as treatments. free of any obstacles while fish were allowed to disperse in the arenas as described above. The first Water depth experiment preliminary experiment tested for the effects of pre- trial conditions on experimental outcomes. Fish were Based on prior results, fish were collected from held in aquaria for >24 h pretrial and were either fed Shireen’s Pond for this experiment, fish were not fed (ad libitum) or not within the 24 h preceding dispersal for 24 h before the trial, and all arenas were lighted.
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