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macroinvertebrate biomass was not a significant predictor of breakdown rates of either litter type in coarse- or fine-mesh bags (F 1,2 = 0.06 to 8.10, r 2 = 3 to 80%, P = 0.1 to 0.8). Discussion A large range (~0-850 g AFDM m -2 ) in the abundance of organic matter among the cave streams was found in this study. A few previous studies have reported the abundance of organic matter in cave streams from c. 30 to 130 g AFDM m -2 (Simon & Benfield, 2001, 2002; Simon, Pipan & Culver, 2007). The large range in organic matter abundance shown in this study illustrates how resource supply can both vary greatly among caves within a limited geographic area and in some cases can be similar to that in many surface stream types (e.g. arid/semi-arid, deciduous and boreal streams; Jones, 1997). Factors that contribute to this variability in organic matter abundance among cave ecosystems are unknown, but are likely to be a combination of landscape features (e.g. topology and plant community structure), cave structure (e.g. depth of cave and size of voids and fractures in the surrounding bedrock) and climate (e.g. precipitation and hydrology). For example, the large quantities of organic matter in Tony Sinks Cave were probably the result of strong surface connections. This cave contained several entrances, both vertical and horizontal, that intersected intermittent stream channels. Additionally, large ceiling fissures were present in portions of the cave, below which large deposits of organic matter were observed. In surface streams, the quantity of organic matter is known to influence macroinvertebrate biomass (Dobson & Hildrew, 1992; Wallace et al., 1999; Negishi & Richardson, 2003). In this study, however, no significant correlation was found between ambient organic matter biomass and macroinvertebrate biomass in litter bags. Nevertheless, Tony Sinks Cave had both the largest amount of organic matter and the highest macroinvertebrate biomass 19
per litter bag. Similarly, Huntsman et al. (2011b) reported that benthic macroinvertebrate biomass was higher in cave streams with higher quantities of organic matter. Together, these studies show that organic matter abundance can influence macroinvertebrate biomass in cave streams. Epigean taxa dominated the communities in all caves in this study, while potentially shredding taxa contributed little to total biomass. Other litter breakdown studies in cave streams have also reported a relatively high biomass of epigean taxa, with shredding taxa composing a small percentage of the overall community (Galas et al., 1996; Brussock et al., 1988). The absence of shredding taxa in cave systems is probably due to the combined limitation in both habitat access and resource availability (e.g. coarse particulate organic matter). Active colonization by shredding insect taxa via oviposition is reduced in cave systems because of the limited number of openings to surface habitats. Additionally, perennial aquatic surface habitats are not always present in recharge areas, thereby limiting the possibility of cave streams being actively (e.g. via drift) or passively (e.g. displacement due to floods) colonized by both insect and crustacean (e.g. amphipods) shredding taxa. Simon & Benfield’s (2001) study illustrates how connectivity and resource availability interact to structure cave stream communities. Cave streams with direct upstream surface connections received higher inputs of coarse particulate organic matter and had higher shredder abundances (epigean and stygophilic taxa) than disconnected cave streams that only received fine particulate organic matter. Similar to Simon & Benfield (2001), the highest shredder biomass was found in Tony Sinks Cave, which had both the highest organic matter abundance and strongest apparent surface connection. The breakdown of organic matter in cave and surface streams is mediated by the same processes, including leaching, microbial processing, consumption, and physical fragmentation 20
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 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: Figs. 5a, b). The breakdown rate of
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
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
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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
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
Page 139 and 140:
Whitmore N. & Huryn A. (1999) Life
Page 141 and 142:
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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