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matter processing (e.g. trophic structure of communities) may be broadly similar across geographically diverse areas. Introduction Cave streams are quintessential donor-controlled ecosystems representing one endpoint along a heterotrophic spectrum (Polis & Strong, 1996; Simon, Benfield & Macko, 2003). Lacking photoautotrophs, the productivity of most cave ecosystems (except those based on chemolithoautotrophy; Sarbu, Kane & Kinkle, 1996, Sarbu, 2001; Engel et al., 2004) depends on organic matter from the surface (Poulson & Lavoie, 2001). The amount and type of organic matter imported into a cave is largely determined by its connectivity to the surface (Poulson & Lavoie, 2001). Systems weakly connected to the surface typically receive fine particulate and dissolved organic matter, while systems that are well connected via cave entrances and sink holes may receive relatively high inputs of dissolved, fine and coarse particulate organic matter (e.g. leaves and woody debris, Poulson & Lavoie, 2001). Regardless of connectivity, the quantity and quality of these inputs are often low compared to surface streams due to a lack of direct riparian inputs and prior biological processing in surface and soil habitats, resulting in the characterization of cave ecosystems as being energy-limited (Culver, 1982; Culver, Kane & Fong, 1995; Poulson & Lavoie, 2001; Graening & Brown, 2003). As a consequence of differences in cave-surface connectivity, however, gradients in resource availability exist that may influence the biomass and diversity of species found within caves. These differences in community structure may ultimately affect cave ecosystem processes. A few studies have attempted to examine the links among organic matter inputs, community structure and ecosystem processes in cave systems. Schneider, Christman & Fagan (2011) showed that organic matter quality (leaf litter vs. rat carcasses) influenced terrestrial 7
community structure in cave “pits” (i.e. vertical cave passages with direct connections to the surface). Huntsman et al. (2011b) found that growth rates and production of the obligate cave salamander Gyrinophilus palleucus (McCrady) were positively related to amount of organic matter in cave streams, presumably because of higher prey availability. In a study of groundwater systems, Datry, Malard & Gibert (2005) reported that aquifers recharged by storm water received higher inputs of dissolved organic matter and had higher densities and diversities of invertebrates, suggesting that organic matter supply may influence groundwater community structure. Similar patterns have been observed in both phreatic aquifers and cave streams affected by organic pollution (Sinton, 1984; Smith et al., 1986; Madsen, Sinclair & Ghiorse, 1991; Notenboom, Plénet & Turquin, 1994; Simon & Buikema, 1997; Sket, 1999). Fewer studies have examined how organic matter abundance affects ecosystem processes in cave streams. Cooney & Simon (2009) used a microcosm experiment to show that amendment of dissolved organic matter increased microbial productivity and enzyme activity on cave-stream rocks. The processing of particulate organic matter in caves has also received some attention. Simon & Benfield (2001) found that cave streams with direct upstream connections to the surface (e.g. cave entrances and sinkholes) had high organic matter inputs, communities dominated by epigean (surface) and stygophilic (species that reproduce in both cave and surface habitats) taxa and faster decomposition of white oak (Quercus alba L.) litter. In contrast, streams without large upstream openings to the surface had lower organic matter standing stocks, were dominated by hypogean (cave specialist) taxa and had slower rates of litter breakdown. Differences in litter breakdown rates among cave streams were attributed to the degree of surface connection, which controlled both organic matter inputs and the biomass of surface taxa. Relationships between organic matter abundance and litter breakdown are not simple, however. 8
Page 1 and 2: THE INFLUENCE OF ENERGY AVAILABILIT
Page 3 and 4: ABSTRACT Detritus from surface envi
Page 5 and 6: LIST OF ABBREVIATIONS AND SYMBOLS m
Page 7 and 8: NC IL ON U.K. VIAT VIE Fig. North C
Page 9 and 10: AIC K-S Akaike information criterio
Page 11 and 12: laboratory work and was an excellen
Page 13 and 14: 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: CHAPTER 2 EFFECTS OF ORGANIC MATTER
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 and 66: crop organic matter significantly i
Page 67 and 68: macroinvertebrate biomass to levels
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
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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
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Huryn A.D. & Wallace J.B. (1987) Pr
Page 109 and 110:
Whitmore N. & Huryn A.D. (1999) Lif
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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
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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
Page 139 and 140:
Whitmore N. & Huryn A. (1999) Life
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Table 2. Assimilation efficiencies
Page 143 and 144:
Table 3. Continued Tanypodinae 0.03
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Table 5. Estimates of mean wood and
Page 147 and 148:
Figure 1. (A) Box and whisker plot
Page 149 and 150:
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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