Cave Ecology

Contents......Page 6
 1.1 On Caves, Cave Ecology, and Cave Inhabitants......Page 9
 1.3 What Can Bring a Multi-author Book on Cave Ecology?......Page 10
 References......Page 11
Part I: Into Karst, Caves and Cave Habitats......Page 13
  2.1 Introduction......Page 14
  2.2 Cave Patterns......Page 15
  2.3 Size Distribution of Cave Voids......Page 16
   2.4.1 Speleogenesis......Page 18
   2.4.2 Hydromorphic Zones in Karst......Page 19
   2.5.1 Cave Ventilation......Page 22
   2.5.2 Attenuation of Surface Temperature......Page 23
   2.5.3 The Chemistry of the Cave Environment......Page 24
  2.6 Non-karstic Caves......Page 26
  References......Page 27
  3.1 Introduction......Page 29
   3.2.1 Abiotic Factors Characteristic of Cave Communities......Page 33
   3.2.2 Environmental Zones......Page 34
  3.3 Aquatic Cave Habitats......Page 36
   3.3.1 The Aquatic Living Space......Page 37
   3.3.2 Groundwater Habitat Features......Page 40
  References......Page 41
Part II: There is Rich and Diverse Life in Caves......Page 44
  4.1 History of Classification Schemes......Page 45
  4.2 Overview of Common Features of Cave Inhabitants......Page 47
   4.3.1 Morphological Adaptations......Page 49
   4.3.2 Physiological Adaptations......Page 50
   4.3.3 Behavioral Adaptations......Page 53
   4.3.4 Special Adaptations......Page 55
   4.4.1 Morphological Adaptations of Stygobionts......Page 57
   4.4.2 Physiological Adaptations of Stygobionts......Page 60
   4.4.3 Behavioral Adaptations of Stygobionts......Page 61
   4.4.4 Special Adaptations......Page 63
  References......Page 65
  5.1 Introduction......Page 72
  5.3 Microbial Diversity in Caves: The Molecular Era (1997-2012)......Page 74
  5.4  Microbial Diversity in Caves: The Genomics Era (2013-Present)......Page 78
  5.5 Ecological Themes in Cave Microbial Communities......Page 81
   5.6.1 Archaea......Page 83
   5.6.2 Fungi......Page 85
  5.7 Toward a Better Understanding of Microbial Cave Diversity......Page 86
  References......Page 88
  6.1 Introduction......Page 94
  6.2 Zonation of Phototrophic Communities......Page 97
   6.3.1 Diversity......Page 98
   6.3.2 Environmental Conditions and Estimates for Colonization......Page 100
   6.3.3 Adaptations of Phototrophs for Light-Deprived Cave Environments......Page 101
   6.3.4 Lampenflora: A Problem for Show Caves......Page 102
  6.4 Phototrophs Impact Cave (Micro)Morphology......Page 103
  References......Page 105
  7.1 Introduction......Page 110
   7.1.1 Statistics, Patterns, Sampling......Page 111
   7.1.2 Original Features of Terrestrial Subterranean Diversity......Page 113
   7.1.4 Temperate Versus Tropical Diversity......Page 115
   7.1.5 Lifestyles and Life Forms......Page 116
   7.2.2 Gastropoda (Snails, Fig. 7.3b)......Page 119
  7.3 Arachnida......Page 121
   7.3.1 Opiliones (Fig. 7.3d-g)......Page 122
   7.3.2 Scorpiones (Fig. 7.3h)......Page 124
   7.3.4 Palpigradi......Page 125
   7.3.6 Acari (Mites) (Fig. 7.3j, k)......Page 126
   7.3.7 Araneae (spiders, Fig. 7.3l-n)......Page 127
   7.3.8 Amblypygi (Whip Spiders, Fig. 7.3o, p)......Page 128
  7.4 Myriapoda......Page 129
   7.4.2 Diplopoda (Millipedes, Fig. 7.4b-i)......Page 130
   7.5.1 Decapoda: Brachyura (Crabs, Fig. 7.4j)......Page 135
   7.5.3 Isopoda: Oniscidea (Woodlice; Fig. 7.4k-m)......Page 136
  7.6 Collembola (Springtails; Fig. 7.4n-t)......Page 137
  7.7 Diplura (Fig. 7.4u)......Page 140
  7.8 Zygentoma (Fig. 7.4v)......Page 141
   7.9.1 Orthoptera (Fig. 7.5a-c)......Page 142
   7.9.2 Grylloblattodea......Page 145
   7.9.5 Psocoptera......Page 146
   7.9.7 Hemiptera: Fulgoromorpha (Fig. 7.5h)......Page 147
   7.9.10 Diptera (Flies, Fig. 7.5i)......Page 148
  7.10 Insecta: Coleoptera (Beetles)......Page 149
   7.10.1 Adephaga: Carabidae (Groundbeetles; Figs. 7.5j-q and 7.6a-d)......Page 150
   7.10.2 Polyphaga......Page 156
  References......Page 161
  8.1 Introduction......Page 176
   8.2.1 Protista: Protists......Page 177
   8.3.3 Platyhelminthes: Flat Worms......Page 180
   8.3.6 Nematoda: Roundworms......Page 182
   8.3.10 Annelida: Segmented Worms......Page 183
   8.4.1 Crustacea......Page 185
   8.4.2 Hydrachnidia: Water Mites......Page 190
   8.5.1 Pisces: Fishes......Page 191
  References......Page 192
  9.1 Introduction......Page 198
   9.2.1 Spatial Scale and Availability of Data......Page 200
   9.2.4 Dealing with Sampling Difficulties......Page 201
   9.2.7 Selection of Relevant Variables......Page 203
   9.3.1 Global Distribution of Biodiversity Hotspots......Page 204
  9.4 Striking Features of Subterranean Biodiversity Patterns at Continental Scales......Page 205
   9.4.1 Scale Dependence of Species Richness Patterns......Page 207
   9.4.2 Range Size Follows a Rapoport Rule in the Palearctic......Page 208
   9.4.3 A High Spatial Turnover in Species Composition Over Short Distances......Page 209
  9.5 Subterranean Biodiversity Patterns at Regional Scales......Page 211
   9.5.1 Regional Species Accumulation Curves Rarely Reach Saturation......Page 212
   9.5.2 The Disproportionally High Contribution of Regional Diversity to Overall Richness......Page 214
   9.5.3 Hotspots Within Hotspots......Page 215
   9.6.1 Environmental Factors: Energy, Spatial Heterogeneity, and History......Page 216
   9.6.2 Multi-causality and Spatial Non-stationarity......Page 217
   9.6.3 The Importance of Historical Climate Variability......Page 218
   9.6.5 Explanations Vary Among Regions......Page 220
   9.7.1 Cryptic Species Do Not Challenge Biodiversity Determinants......Page 221
   9.7.2 Environmental DNA Can Speed Up Species Inventory......Page 223
   9.7.3 Making Use of Phylogeny to Foster Our Understanding of Biodiversity Patterns......Page 224
  9.8 Conclusions......Page 225
  References......Page 226
  10.1 Introductory Background......Page 231
  10.2 The Crucial Dilemma: No Dispersal but Multiple Origins, or Single Origin but Geographic Dispersion and Diversification......Page 236
  10.3 Once or Many? Single Versus Multiple Origin of the Subterranean Colonisation......Page 238
  10.4 Relation Between Age and Troglomorphism......Page 241
  10.5 Dispersal and Diversification......Page 242
  10.6 General Overview and Concluding Remarks......Page 246
  References......Page 250
Part III: About Communities, Populations and Food......Page 255
  11.1 Cave Communities......Page 256
  11.2 Diversity of Cave Communities......Page 258
  11.3 Functional Composition of the Community......Page 259
  11.4 Turnover in Space......Page 260
  11.5 Turnover in Time......Page 261
  11.6 Ecological Niche and Interspecific Competition......Page 263
  References......Page 266
   12.1.1 The Genetics of Cave Populations Reflects Evolutionary History......Page 269
   12.1.2 The Paradox of Large Subterranean Ranges......Page 270
   12.1.3 Exogenous and Endogenous Processes Determine the Structure of Subterranean Populations......Page 272
   12.2.1 Brief Historical Account......Page 273
   12.2.2 Genetic Structure Results from Historical Processes and Concurrent Brakes in Connectivity Between Cave Populations......Page 275
   12.3.1 The Sluggish Interstitial Highway Produces Strong Genetic Structuring and Isolation by Distance......Page 276
   12.3.2 Cave Populations Tend to Have an ``All-or-Nothing´´ Structure......Page 279
   12.3.3 Dispersal of Evolutionarily Old and Large-Bodied Cave Species Probably Proceeds via the Surface......Page 280
   12.3.4 Non-dispersal Has Different Meanings in Karstic and in Interstitial Groundwater......Page 282
   12.3.5 Terrestrial Troglobionts Are Less Studied but Seem to Share Basic Patterns of Genetic Structuring with Aquatic Ones......Page 283
   12.4.1 Conflicting Theoretical Predictions and a Paradox......Page 285
   12.4.2 Exogenous and Endogenous Factors Affect Genetic Variability in Different Ways......Page 287
   12.4.3 Exogenous Propagule Pressure Appears to Shape the Genetics of Young Cave Populations More than Bottlenecks......Page 288
  12.5 Conclusions and Perspectives......Page 290
  References......Page 292
  13.1 Introduction......Page 296
  13.2 Food Variability and Communities of Biota......Page 298
  13.3 Microorganisms in Oligotrophic Caves......Page 299
  13.4 Faunal Communities in Oligotrophic Caves......Page 300
  13.5 Morphological and Biological Adaptations of the Cave Fauna......Page 301
  References......Page 304
  14.1 Introduction......Page 307
  14.2 Detritus-Based Cave Ecosystems......Page 308
   14.2.1 Dissolved Organic Matter......Page 309
   14.2.2 Particulate Organic Matter......Page 310
   14.2.3 The Gatekeeper: The Role of Microbes in Detritus-Based Cave Food Webs......Page 311
   14.2.4 Detritus Quantity......Page 312
   14.2.5 Surface Connectivity, Detritus Surpluses, and Spatial Distribution of Detritus......Page 313
   14.2.6 Nutrients and Detrital Quality......Page 314
   14.2.7 Beyond Detritus......Page 316
   14.3.1 Chemolithoautotrophic Primary Production......Page 317
   14.3.2 A Complex Story: The Role of Chemolithoautotrophic Primary Production in Structuring Cave Communities......Page 318
   14.3.3 The Linked Conceptual Model: Chemolithoautotrophically Based and Detritus-Based Ecosystems......Page 319
  References......Page 320
Part IV: Particular Cave Ecosystems......Page 327
  15.1 Introduction......Page 328
  15.2 Ice Caves as Ecosystems......Page 332
  15.3 Ice Caves and Palaeoclimate Reconstruction......Page 337
  15.5 Examples of Ice Caves Ecosystems......Page 338
  15.6 Ice Caves and Global Change......Page 342
  References......Page 343
  16.1 Introduction to Sulphide-Based Ecosystems......Page 347
  16.2 Hypogenic Caves......Page 348
   16.2.1 Trophic Webs in Hypogenic Caves......Page 349
   16.3.1 Movile Cave: The First Terrestrial Chemosynthesis-Based Ecosystem Ever Discovered......Page 350
   16.3.2 Cueva de Villa Luz (Mexico): A Cave Where Speciation Occurs in Its Multiple Habitats......Page 358
  References......Page 362
  17.1 Introduction......Page 365
  17.2 Geological Aspects......Page 366
  17.3 Ecological Features......Page 368
  17.4 Animal Communities......Page 370
  References......Page 376
  18.1 What Are Anchialine Caves?......Page 378
  18.2 Exploration and Investigation of Anchialine Caves......Page 380
  18.3 Diversity of Anchialine Fauna......Page 381
  18.4 Anchialine Cave Food Webs and Functioning......Page 387
  18.5 Future Studies......Page 388
  References......Page 389
  19.1 Introduction......Page 393
  19.2 Anchialine System in Lanzarote......Page 394
  19.3 Biodiversity and Ecological Studies in La Corona Lava Tube: A Hotspot of Diversity for Anchialine Subterranean Fauna......Page 397
  19.4 Biodiversity Studies in Anchialine Pools and Wells in Lanzarote......Page 402
  19.5 Evolutionary Studies in Lanzarote: Origin of the Anchialine Fauna......Page 403
  19.7 Conservation of Anchialine Habitats......Page 404
  References......Page 406
  20.1 Introduction......Page 409
  20.2 Subterranean Habitat Other than Caves......Page 411
  20.3 Subterranean Fauna Outside Caves......Page 414
   20.3.1 Stygofauna......Page 416
   20.3.2 Troglofauna......Page 418
  20.4 Species Distributions......Page 422
  References......Page 424
  21.1 Introduction......Page 429
  21.2 Subterranean Environments in Iron Ore Landscapes......Page 430
  21.3 Troglobitic Species......Page 432
  21.4 Iron Ore Formations in Brazil and Australia......Page 435
  21.5 Ecology......Page 436
  21.6 Threats......Page 438
  References......Page 439
Part V: Conservation and Protection of Cave Habitats and Cave Fauna......Page 442
  22.1 Introduction......Page 443
  22.2 International Legislation......Page 446
  22.3 Conservation Organizations......Page 454
  22.4 National Legislation......Page 458
   22.4.1 European Legislation......Page 459
   22.4.2 North, Central, and South American Legislation......Page 461
   22.4.3 Regional and Local Conservation Organizations......Page 465
  22.5 Conclusions......Page 466
  References......Page 467
  23.1 Introduction......Page 471
  23.2 Complexities of Subterranean Fauna Assessment......Page 473
   23.2.1 Sampling Framework......Page 475
   23.2.2 Sampling Efficiency......Page 477
   23.2.3 Inferring Distributions......Page 478
   23.2.4 Identifying Species......Page 480
   23.2.5 Determining Impacts......Page 481
  23.4 Awareness of Subterranean Fauna......Page 482
  References......Page 483
Part VI: Cave Ecology for the Twenty-First Century......Page 487
  24.1 Introduction......Page 488
  24.3 Natural History: Basic Biology......Page 489
  24.4 Enumerating Numbers and Movements of the Fauna......Page 490
  24.5 Sampling of Aquatic and Terrestrial Animals......Page 492
  24.6 Sampling of Fauna in Anchialine Systems......Page 493
  24.8 Diversity of Subterranean Fauna......Page 494
  24.9 Metagenomics and Environmental DNA (eDNA)......Page 497
  24.10 Determining Predator-Prey Relationships......Page 499
  24.11 Are Parasites and Commensals of Subterranean Fauna So Rare?......Page 500
  24.12 Ecotone and the Transition to Subterranean Life......Page 501
  24.14 Is Carbon Dioxide Concentration Significant to Cave Ecology?......Page 502
  24.15 Understanding the Dynamics of Cave Climate......Page 503
  24.16 Cave Faunas as Indicators of Climate Change......Page 504
  24.17 Biofilms, the Drivers of Cave Ecology?......Page 505
  24.18 Biogeochemical Processes as Drivers of Cave Ecology......Page 507
  24.19 Ecosystem Functions (Services)......Page 508
  24.20 Age of Subterranean Lineages as Indicators of Past Conditions......Page 509
  24.21 Conservation Ecology......Page 510
  24.22 Be Alert for Opportunistic Experiments......Page 511
  References......Page 512
Glossary......Page 524
Index......Page 532