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significant increase in the biomass of facultative species (Wilcoxon signed rank test: n = 20, W = 44, P = 0.008; Fig. 1c, d). Macroinvertebrate community structure changed significantly following the litter amendment (R-statistic = 0.50, P = 0.001; Fig. 2). Pairwise ANOSIMs indicated that community structure did not differ among study reaches prior to the litter amendment (Pre-CR vs. Pre-MR; R-statistic = 0.10, P = 0.058) but diverged dramatically following the litter amendment (Post-CR vs. Post-MR; R-statistic = 0.62, P = 0.001; Fig. 2). Pre-litter amendment community structure within each reach differed from post-litter amendment structure within each reach (Pre-CR vs. Post-CR and Pre-MR vs. Post-MR; R-statistic = 0.53 – 0.70, P = 0.002; Fig. 2). Seven taxa accounted for 82-91% of the overall dissimilarity among all pair-wise comparisons (Fig. 3). Three taxa, Polypedilum, Oligochaeta, and Ephemeroptera, increased in biomass in both study reaches following the litter amendment, but the strongest increase occurred in the manipulation reach. A fourth taxon, Paraphaenocladius, increased similarly in both study reaches following the amendment, while biomass of two taxa, Tanypodinae genus A and B, increased only in the manipulation reach following the amendment. The biomass of a final taxon—the obligate cave isopod Caecidotea—was similar between both study reaches before and after the amendment. Organic matter storage explained a large and significant amount of the variation in macroinvertebrate biomass within the combined cave and surface stream data sets (Fig. 5; F 30 = 268, R 2 = 0.90, P < 0.001). Annual mean organic matter storage and macroinvertebrate biomass were generally lowest in the cave streams and highest in the forested headwater surface streams without experimental litter-exclusion. The litter exclusion experiment by Wallace et al. (1999) reduced both organic matter storage and macroinvertebrate biomass to levels similar to that of high-detritus cave streams, while the litter amendment experiment in this study increased organic matter storage and 49

macroinvertebrate biomass to levels similar to the forested headwater surface stream with litter exclusion (Wallace et al. 1999). Crayfish and salamanders Larger consumers in the food web (crayfish and salamanders) followed the same general patterns as those of macroinvertebrates (Fig. 1e, f, g, h). The biomass of the obligate cave crayfish C. hamulatus did not respond to the litter amendment (t-test: df = 8, t = 0.4, P = 0.71), while the biomass of the facultative crayfish C. tenebrosus (Wilcoxon signed rank test: n = 20, W = 33, P = 0.04) and salamander Eurycea sp. (Wilcoxon signed rank test: n = 20, W = 36, P = 0.01) showed a significant positive response. Patterns could not be discerned for the obligate cave salamander G. palleucus due to low capture rates. Carbon flow Three species, five groups that include multiple species (e.g. Ephemeroptera or Oligochaeta), and four types of organic matter were included in the stable isotope analyses (Table 3). Within the control reach, δ 13 C values for all types of organic matter ranged from -27 to -28‰, indicating a C3 plant origin. The δ 13 C values of wood and coarse particulate organic matter (CPOM; e.g., leaves from ambient organic matter) in the manipulation reach also indicated a C3 plant origin, while the corn litter δ 13 C value was characteristic of C4 plants (~- 11‰; Fry 2006). Fine particulate organic matter (FPOM) in the manipulation reach appeared to be partially composed of corn litter due to its higher δ 13 C value compared with FPOM from the control reach. With the exception of C. tenebrosus, the δ 13 C values for consumers within the control reach (-25 to -27‰) were similar to those of organic matter, indicating that growth of consumers within the control reach was supported by carbon from C3 plant detritus. All consumers within the manipulation reach, however, were partially supported by carbon from the corn litter, because the δ 13 C values of their tissues were higher and more similar to the δ 13 C 50

macroinvertebrate biomass to levels similar to the forested headwater surface stream with litter<br />

exclusion (Wallace et al. 1999).<br />

Crayfish and salamanders<br />

Larger consumers in the food web (crayfish and salamanders) followed the same general<br />

patterns as those <strong>of</strong> macroinvertebrates (Fig. 1e, f, g, h). <strong>The</strong> biomass <strong>of</strong> the obligate cave<br />

crayfish C. hamulatus did not respond to the litter amendment (t-test: df = 8, t = 0.4, P = 0.71),<br />

while the biomass <strong>of</strong> the facultative crayfish C. tenebrosus (Wilcoxon signed rank test: n = 20, W<br />

= 33, P = 0.04) and salamander Eurycea sp. (Wilcoxon signed rank test: n = 20, W = 36, P =<br />

0.01) showed a significant positive response. Patterns could not be discerned for the obligate<br />

cave salamander G. palleucus due to low capture rates.<br />

Carbon flow<br />

Three species, five groups that include multiple species (e.g. Ephemeroptera or<br />

Oligochaeta), and four types <strong>of</strong> organic matter were included in the stable isotope analyses<br />

(Table 3). Within the control reach, δ 13 C values for all types <strong>of</strong> organic matter ranged from -27 to<br />

-28‰, indicating a C3 plant origin. <strong>The</strong> δ 13 C values <strong>of</strong> wood and coarse particulate organic<br />

matter (CPOM; e.g., leaves from ambient organic matter) in the manipulation reach also<br />

indicated a C3 plant origin, while the corn litter δ 13 C value was characteristic <strong>of</strong> C4 plants (~-<br />

11‰; Fry 2006). Fine particulate organic matter (FPOM) in the manipulation reach appeared to<br />

be partially composed <strong>of</strong> corn litter due to its higher δ 13 C value compared with FPOM from the<br />

control reach. With the exception <strong>of</strong> C. tenebrosus, the δ 13 C values for consumers within the<br />

control reach (-25 to -27‰) were similar to those <strong>of</strong> organic matter, indicating that growth <strong>of</strong><br />

consumers within the control reach was supported by carbon from C3 plant detritus. All<br />

consumers within the manipulation reach, however, were partially supported by carbon from the<br />

corn litter, because the δ 13 C values <strong>of</strong> their tissues were higher and more similar to the δ 13 C<br />

50

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