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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
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THE INFLUENCE OF ENERGY AVAILABILIT
Page 3 and 4:
ABSTRACT Detritus from surface envi
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LIST OF ABBREVIATIONS AND SYMBOLS m
Page 7 and 8:
NC IL ON U.K. VIAT VIE Fig. North C
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AIC K-S Akaike information criterio
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laboratory work and was an excellen
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LIST OF TABLES TABLE 2.1 TABLE 2.2
Page 15 and 16: LIST OF FIGURES FIGURE 2.1 (a) Box
Page 17 and 18: FIGURE 4.3 Growth models for Orcone
Page 19 and 20: chemolithoautotrophy-based systems;
Page 21 and 22: ecology, 51, 31-53. Gibert J. & Cul
Page 23 and 24: CHAPTER 2 EFFECTS OF ORGANIC MATTER
Page 25 and 26: community structure in cave “pits
Page 27 and 28: communities and how variation in co
Page 29 and 30: the different source locations was
Page 31 and 32: of natural-log transformed data (%
Page 33 and 34: peak in organic matter in Big Mouth
Page 35 and 36: Figs. 5a, b). The breakdown rate of
Page 37 and 38: per litter bag. Similarly, Huntsman
Page 39 and 40: ags was the greater retention of li
Page 41 and 42: Historically, limited resource inpu
Page 43 and 44: Culver, D.C. & Pipan, T. (2009) The
Page 45 and 46: Merritt, R.W., Cummins, K.W. & Berg
Page 47 and 48: Table 1. Mean (1 S.D.) macroinverte
Page 49 and 50: Table 2. Mean (±1 S.D.) daily temp
Page 51 and 52: Figure 1. (a) Box and whisker plot
Page 53 and 54: Figure 3. Non-metric multidimension
Page 55 and 56: Figure 5. Box and whisker plot of l
Page 57 and 58: streams, while the obligate-cave sp
Page 59 and 60: More recent observational and exper
Page 61 and 62: of netting (mesh size 2.5×1.5-cm)
Page 63 and 64: the two end-members. This conservat
Page 65: crop organic matter significantly i
Page 69 and 70: and surface streams (20-35,000 g m
Page 71 and 72: these factors, the stable isotope a
Page 73 and 74: surface streams. Thus, it is likely
Page 75 and 76: subsides to ecosystem dynamics have
Page 77 and 78: populations. Journal of Applied Eco
Page 79 and 80: Poulson T.L. & Lavoie K.H. (2001) T
Page 81 and 82: Wood P., Gunn J. & Perkins J. (2002
Page 83 and 84: Table 2. Mean (±1 standard deviati
Page 85 and 86: Table 2. Continued Chironomini Para
Page 87 and 88: Figure 1. Mean (bars are standard e
Page 89 and 90: Figure 3. Non-metric multidimension
Page 91 and 92: CHAPTER 4 REXAMINING EXTREME LONGEI
Page 93 and 94: individuals, he predicted that it w
Page 95 and 96: A phylogeographic study by Buhay &
Page 97 and 98: differences in size structure among
Page 99 and 100: and females in Tony Sinks Cave were
Page 101 and 102: Estimates of life span for O. austr
Page 103 and 104: available size-classes were well re
Page 105 and 106: References Anonymous (1999) Cave Sc
Page 107 and 108: Huryn A.D. & Wallace J.B. (1987) Pr
Page 109 and 110: Whitmore N. & Huryn A.D. (1999) Lif
Page 111 and 112: Table 2. Estimated life span (years
Page 113 and 114: Figure 1. Annual growth increment (
Page 115 and 116: Figure 3. Growth models for Orconec
Page 117 and 118:
CHAPTER 5 CONSUMER-RESOURCE DYNAMIC
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following incidental inputs of orga
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Temperature data were not available
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minimum (Venarsky et al., 2012b). A
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100% organic matter (i.e., maximum
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Macroinvertebrate biomass varied si
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Discussion The energy-limitation hy
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crayfish captured in most months we
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systems is likely high, especially
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function of microbial films in grou
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Crustacean Biology, 4, 35-54. Momot
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Whitmore N. & Huryn A. (1999) Life
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Table 2. Assimilation efficiencies
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Table 3. Continued Tanypodinae 0.03
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Table 5. Estimates of mean wood and
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Figure 1. (A) Box and whisker plot
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CHAPTER 6 OVERALL CONCLUSIONS Due t
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characteristics (e.g., higher growt
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characteristics, highly efficient p
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