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Recent research suggests that non-native Purple Loosestrife (Lythrum salicaria) invasions may negatively affect the performance of larval American Toad tadpoles (Bufo americanus), and that compounds leached from L. salicaria leaves play a direct or indirect role in this effect.2We raised individual B. americanus and Gray Treefrog (Hyla versicolor) tadpoles on high-quality diets in aqueous extracts of senescent leaves from L. salicaria, native Broad-Leaf Cattail (Typha latifolia), and control water to determine whether loosestrife extracts directly affect anuran tadpole performance.3Even at high artificial food levels, B. americanus survival was significantly lower in L. salicaria extracts compared with T. latifolia extracts and a water control. Food level strongly affected B. americanus development, but tadpoles raised in L. salicaria extract were less developed compared with conspecifics raised in cattail extract or water. Unlike B. americanus, Hyla performance was not affected by exposure to any plant extract compared with the water control.4Our study implicates secondary plant compounds as a mechanism underlying the impact of an invasive plant on some but not all native fauna.

We hypothesize that high tannin concentrations of L. salicaria leaves have the potential to create environments that are directly toxic to B. americanus tadpoles. We hypothesize that obligate gill breathers such as B. americanus tadpoles are highly sensitive to gill damage caused by high concentrations of phenolics. Other anuran species such as H. versicolor that develop well-functioning lungs early may be less affected by high tannin concentrations.The increasing spread of non-native plant species raises concerns about the impacts of plant invasions on habitat quality for native fauna. Generally, we expect that invasive plants will affect animal communities by altering resource availability (trophic effects) or habitat structure. However, some invaders may have particular traits that contribute to their effects on native fauna in unexpected ways. For example, Purple Loosestrife (Lythrum salicaria) is a highly invasive plant in wetlands throughout North America. L. salicaria invasions are linked to declines in habitat quality for breeding birds, mammals and turtles (reviewed by Blossey 1999).

Recent evidence shows that L. salicaria invasions may also reduce growth and survival of American Toad tadpoles (Bufo americanus) (Brown et al. 2005). Brown et al. (2005) present evidence of changes in tadpole diets associated with L. salicaria invasions (a trophic effect), which might be the underlying cause of reduced performance in L. salicaria habitats. They also demonstrated raising B. americanus tadpoles in aqueous extracts of L. salicaria leaves could produce similar levels of mortality, raising the possibility that soluble compounds in L. salicaria leaves affect the productivity or composition of food resources or act directly on developing tadpoles.Alteration of the chemical environment is known for a number of invasive plant species (Blossey 1999; Caraco & Cole 2002; Ehrenfeld 2003), but little is known about the effects of L. salicaria invasion on the chemistry of wetland habitats. Templer, Findlay & Wigand (1998) document changes in sediment chemistry associated with L. salicaria invasion.

Rauha et al. (2001) found that L. salicaria leaves have particularly high concentrations of phenolics (tannins), but the fate of those compounds and their potential effects on host communities or ecosystems has not been addressed. We recently collected water samples from L. salicaria-invaded habitats in 13 wetlands in New York, Rhode Island, and Minnesota, and found reactive phenolic concentrations during the spring when anurans were breeding as low as 1 mg l−1 and as high as 11 mg l−1 (B. Blossey and J. C. Maerz, unpublished data). These results show that while not ubiquitous, habitats invaded by L. salicaria can have high tannin concentrations that might affect aquatic fauna.High tannin concentrations can have direct and indirect effects on aquatic fauna. A growing body of literature shows that phenolics are retained at high concentrations in plant leaves, and through effects on decomposition rates (Rier et al. 2002; Tuchman et al. 2003b; Cornelissen et al. 2004; Parsons et al. 2004;

Schweitzer et al. 2004; Rier, Tuchman & Wetzel 2005) and allelopathy (Ervin & Wetzel 2003), inhibit bacteria, phytoplankton and periphyton growth, which consequently affects the performance of organisms at higher trophic levels (Tuchman et al. 2002, 2003a). In addition to trophic effects, aquatic organisms in wetland habitats are literally bathing in leached plant compounds that they may ingest while feeding or that pass over gills as the animals respire; yet there has been little attention to the direct effects of tannins or other secondary plant compounds on aquatic fauna. Applebaum & Birk (1979) note that saponins leaching from aquatic plants poison fish by lysing blood cells, a process that has been recognized and exploited by fisherman since ancient times. Temmink et al. (1989) found that tannins leached into water from tree bark had lethal and subacute effects on fish at high and low concentrations, respectively. Lethal effects were related to gill lesions caused by phenolic reactions with sensitive tissues, and subacute effects may have been related to a combination of gill damage and phenolics reducing the nutritional quality of food or interfering with digestive enzymes.

Though the results of Brown et al. (2005) suggests that compounds, most likely high concentrations of tannins, leached from L. salicaria may contribute to the plant's effect on B. americanus tadpole performance, the design of that study cannot distinguish between the indirect trophic effects of L. salicaria extracts on algal growth or direct toxicity as the mechanism.The purpose of this study was to determine whether compounds leached from L. salicaria leaves directly affect two species of larval anuran. Using high-quality artificial food supplements to reduce indirect trophic effects, we compared survival and development of tadpoles raised in extracts of L. salicaria, T. latifolia and control water. To follow up on previous research, we tested B. americanus tadpoles (hereafter referred to as Bufo). We tested a second amphibian species because studies of larval amphibian responses to pesticides (Relyea 2003) and different plant substrates (Skelly, Freidenburg & Kiesecker 2002) show that species can differ in their sensitivity similar compounds.