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Groundwater is a significant component of the world’s water balance and accounts for >90 % of usable freshwater. Around the world groundwater is an important source of water for major cities, towns, industries, agriculture and forestry. Groundwater plays a role in the ecological processes and ‘health’ of many surface ecosystems, and is the critical habitat for subterranean aquatic animals (stygofauna). Over-abstraction or contamination of groundwater resources may imperil the survival of stygofauna and other groundwater-dependent ecosystems (GDEs). In two karst areas in Western Australia (Yanchep and Leeuwin-Naturaliste Ridge), rich stygofauna communities occur in cave waters containing submerged tree roots. These aquatic root mat communities were listed as critically endangered because of declining groundwater levels, presumably caused by lower rainfall, groundwater abstraction, and/or forest plantations. Investigation of the hydrology and ecology of the cave systems was considered essential for the conservation and recovery of these threatened ecological communities (TECs). This thesis investigated the hydrology and ecology of one of the TECs, located in the Jewel Cave karst system in the Leeuwin-Naturaliste Ridge. A multi-disciplinary approach was used to explore aspects pertinent to the hydrology and ecology of the groundwater system.
Thermoluminescence dating of the limestone suggested that development of the karst system dates from the Early Pleistocene and that caves have been available for colonisation by groundwater fauna since that time. Speleogenesis of the watertable maze caves occurred in a flank margin setting during earlier periods of wetter climate and/or elevated base levels. Field mapping and leveling were used to determine hydrologic relationships between caves and the boundaries of the karst aquifer. Monitoring of groundwater levels was undertaken to characterise the conditions of recharge, storage, flow and discharge. A hydrogeologic model of the karst system was developed.
The groundwater hydrograph for the last 50 years was reconstructed from old photographs and records whilst radiometric dating and leveling of stratigraphic horizons enabled reconstruction of a history of watertable fluctuations spanning the Holocene to Late Pleistocene. The watertable fluctuations over the previous 50 years did not exceed the range of fluctuations experienced in the Quaternary history, including a period 11,000 to 13,000 years ago when the watertable was lower than the present level.
The recent groundwater decline in Jewel Cave was not reflected in the annual rainfall trend, which was above average during the period (1976 to 1988) when the major drop in water levels occurred. Groundwater abstraction and tree plantations in nearby catchments have not contributed to the groundwater decline as previously suggested. The period of major watertable decline coincided with a substantial reduction in fire frequency within the karst catchment. The resultant increase in understorey vegetation and ground litter may have contributed to a reduction in groundwater recharge, through increased evapotranspiration and interception of rainfall. To better understand the relationships between rainfall, vegetation and fire and their effects on groundwater recharge, an experiment is proposed that involves a prescribed burn of the cave catchment with before-after monitoring of rainfall, leaf-area, ground litter, soil moisture, vadose infiltration and groundwater levels.
Molecular genetic techniques (allozyme electrophoresis and mitochondrial DNA) were used to assess the species and population boundaries of two genera and species of cave dwelling Amphipoda. Populations of both species were largely panmictic which was consistent with the hydrogeologic model. The molecular data supported the conclusion that both species of amphipod have survived lower watertable levels experienced in the caves during the Late Pleistocene. A mechanism for the colonization and isolation of populations in caves is proposed.
Multi Dimensional Scaling was used to investigate patterns in groundwater biodiversity including species diversity, species assemblages, habitat associations and biogeography. Faunal patterns were related to abiotic environmental parameters. Investigation of hydrochemistry and water quality characterized the ecological water requirements (EWR) of the TEC and established a baseline against which to evaluate potential impacts such as groundwater pollution.
The conservation status of the listed TEC was significantly improved by increasing the number of known occurrences and distribution range of the community (from 10 m2 to > 2 x 106 m2), and by showing that earlier perceived threatening processes (rainfall decline, groundwater pumping, tree plantations) were either ameliorated or inoperative within this catchment. The GDE in the Jewel Cave karst system may not have been endangered by the major phase of watertable decline experienced 1975-1987, or by the relatively stable level experienced up until 2000. However, if the present trend of declining rainfall in southwest Western Australia continues, and the cave watertable declines > 0.5 m below the present level, then the GDE may become more vulnerable to extinction.
The occurrence and distribution of aquatic root mat communities and related groundwater fauna in other karst catchments in the Leeuwin-Naturaliste Ridge is substantially greater than previously thought, however some of these are predicted to be threatened by groundwater pumping and pollution associated with increasing urban and rural developments. The taxonomy of most stygofauna taxa and the distribution of root mat communities is too poorly known to enable proper assessment of their conservation requirements. A regional-scale survey of stygofauna in southwest Western Australia is required to address this problem. In the interim, conservation actions for the listed TECs need to be focused at the most appropriate spatial scale, which is the karst drainage system and catchment area. Conservation of GDEs in Western Australia will benefit from understanding and integration with abiotic groundwater system processes, especially hydrogeologic and geomorphic processes.
This thesis investigates the hydrology and ecology of a threatened aquatic root mat community in the Jewel Cave karst system in the Leeuwin-Naturaliste Ridge, Western Australia. Development of the karst system dates from the Early Pleistocene and the caves have been available for colonisation by groundwater fauna since that time. Speleogenesis of the watertable maze caves occurred in a flank margin setting during earlier periods of wetter climate and/or elevated base levels. Watertable fluctuations over the last 50 years did not exceed the range experienced in the Quaternary history. The recent groundwater decline in Jewel Cave was not related to rainfall, nor groundwater abstraction nor nearby tree plantations. However, it did coincide with a reduction in fire frequency within the karst catchment. The resultant increase in understorey vegetation and ground litter may have reduced groundwater recharge through increased evapotranspiration and interception of rainfall. The populations of two genera and species of cave dwelling Amphipoda are largely panmictic. Both species have survived lower watertable levels during the Late Pleistocene. A mechanism for the colonization and isolation of populations in caves is proposed. Faunal patterns (including species diversity, species assemblages, habitat associations and biogeography) were related to abiotic environmental parameters. The ecological water requirements of the community were determined as a baseline for evaluation of impacts such as groundwater pollution. If rainfall continues to decline, and the cave watertable declines > 0.5 m below the present level, then the groundwater ecosystem may become more vulnerable to extinction. The taxonomy and distribution of root mat communities is poorly known and a regional-scale survey is required to properly assess their conservation requirements. Meanwhile, conservation actions for the communities need to be focused at the scale of the karst drainage system and catchment area.
The last two decades were particularly prolific in historical biogeography because of new information introduced from other sciences, such as paleogeography, by the development of quantitative methods and by molecular phylogeny. Subterranean beetles represent an excellent object of study for historical biogeography because they are the group with the best representation in the subterranean domain. In addition, species have reduced mobility, display different degrees of adaptations to life in caves and many specialists work on this group. Three processes have shaped the present distribution of the tribe Leptodirini (Coleoptera Cholevinae) in the world: dispersal, vicariance, and extinction. Therefore, three successive stages can be established in the space-time evolution of Leptodirini: (1) dispersal from a center of origin in the present area(s); (2) dispersal, extinction and vicariance among the present area(s); and (3) colonization and speciation in the subterranean domain. The Romanian Leptodirini, especially those from Western Carpathians is examined with respect to these processes. Their pattern of distribution in different massifs and at different altitudes is discussed, with possible explanations from a historical biogeographic point of view.
Geographic distribution of stygobionts is often used to estimate age of a group by assuming vicariant speciation with little or no subsequent dispersal. We investigated the utility of using distributional data for Slovenian stygobiotic copepods by assuming that dispersal is a way to measure age of a species. We list some species of Copepoda that, on the basis of their range and frequency of occupancy within their range, should be older. Body size is not predictor either of range or frequency of occupancy.
The applicability and effectiveness of different kinds of evidence used to estimate the age of lineages – morphological, molecular, phylogenetic, biogeographical, geological – are discussed. Examples from the Brazilian subterranean fauna are presented, using mainly fishes, one of the best studied groups, as a model. Only three taxa including troglobites are object of molecular studies, all in progress. Therefore, molecular clocks cannot be applied yet, and indirect evidence is used. Few phylogenies are available, e.g. for the catfish families Heptapteridae and Trichomycteridae. Theoretically, basal troglobitic clades are older than apical ones, but the possible existence of extinct epigean taxa belonging to such clades hampers the comparison. As well, the limitations of the use degrees of troglomorphism to estimate phylogenetic ages are analyzed with focus on the complexity of the mechanisms underlying morphological differentiation. Paleoclimatic reconstructions based on dating of speleothems from caves in northeastern and southeastern Brazil are available, but limited up to the last 200,000 years, thus useful for relatively recent lineages. Topographic isolation, probable for some fish groups from Central Brazil, is also within the time range of 105 years. Older dated events (in the order of 106 years or more) that may represent vicariant events affecting aquatic lineages with subterranean derivatives are related to the establishment of the modern South American main river basins. In view of the paucity of data useful for estimating the age of Brazilian troglobitic lineages, combined evidence, including morphology, systematics and biogeography, seems to be the best approach at the moment.
The copepod crustacean fauna collected from subterranean habitats, including caves, wells, and the hyporheos of streams in and near the Interior Low Plateaus of the United States is dominated by Cyclopoida, with 39 species, followed by Harpacticoida with 9, and Calanoida with 2. Nearly all of the harpacticoid and calanoid species are widespread, primarily surface-dwelling generalists. Fourteen of the cyclopoids, members of the genera Diacyclops, Itocyclops, Megacyclops, and Rheocyclops, are apparently obligate stygobionts or hyporheic. Several of the species that are more strongly modified for subterranean existence occur only in the more southern, unglaciated areas. Our sampling data support the hypothesis that the more specialized, groundwater-interstitial species have been unable to disperse into previously glaciated regions; whereas some, less-specialized species may have invaded groundwaters from surface habitats as the glaciers receded.
New data fail to support the prevailing theory that meandering bedrock valleys inherit their sinuosity from ancient alluvial rivers. In the Ozarks, observations indicate that bedrock meanders emerge during incision as a result of erosion by emergent groundwater and surface flow. Crustal tilting pressurizes deep aquifers that feed a huge base flow to large springs. Because of their large size and persistence in time, these artesian conduits have the potential to create new base levels of erosion. Transverse speleogenesis causes groundwater flow lines and surface streams to converge toward the springs, thereby further increasing the rate of landscape lowering and creating bedrock meanders. Groundwater outflow accelerates stream piracy, creates asymmetric drainage patterns and cuts channels across structural upwarps. By contrast, the antecedent meander theory favors long-term drainage stability that cannot explain the incredible diversity of the freshwater fauna found in the Central Highlands. Widely disjunct species of highland fish that thrive only in clear, high-gradient streams indicate that the Ouachitas, the Ozarks and the Eastern Highlands were once a continuous upland connected by a “land bridge” in southern Illinois. This connection allowed ancestral species to become widespread enough to be affected by a vicariant event, usually attributed to onset of glaciation. However, a 400-km eastward shift in Gulf of Mexico sedimentation indicates this vicariant event may have occurred in the middle Pleistocene, when it is proposed that the Mississippi River dissected the Central Highlands, separating the Interior Highlands from the Eastern Highlands.
Till recently information about bone accumulations in caves of the Crimean Mountains was comparatively modest. Now several dozens of caves are known on high plateaus, which hold a rich potential regarding Late Pleistocene and Holocene fauna of the Crimean Mountains. The Chatyrdag Plateau is the best studied area with its nine bone-bearing caves documented and two caves (Marble Cave and Emine-Bair-Khosar Cave) where detailed investigations have been conducted. Paleontological potential of the Emine-Bair-Khosar Cave is tremendous. Fauna complexes belonging to the LGM (the complex of cold steps, also confirmed by paleopalinological data) and interstadials and the postglacial (predominantly related to meadows and forested landscapes of warmer climate) are identified in this cave. Enormous amount of bones, their well preserved condition, tafocenosis and species diversity give a special importance to this cave. The ongoing study of the Emine-Bair-Khosar Cave site will contribute to resolution of many issues about paleobiogeography, paleoecology and paleoclimate of this region. In other caves of Crimea important information has been also obtained.
This research is a contribution to the study of subterranean biodiversity in karst areas of Portugal, towards its conservation.
The relative inaccessibility of the subterranean environment is a challenge for the study of its fauna, often accessible only in caves but more widely distributed. The subterranean animals are among the most rare, threatened and worldwide underprotected, often by the simple fact of being unknown.
Karst areas of Portugal occupy a considerable part of the territory and harbor more than 2000 caves. The complex biogeographical history of the Iberian Peninsula allowed the survival of several relict arthropod refugees in the subterranean environment.
Subterranean invertebrates have been ignored, as for as the protection of karst systems are concerned in Portugal, largely because knowledge was scarce and disorganized. Reviewing all the bibliographic sources about subterranean fauna from Portugal and listing troglobiont and stygobiont species and locations, was essential to understand the state of knowledge of species richness and the biogeography and conservation status for the studied areas.
In order to understand subterranean biodiversity patterns in karst areas from Portugal, one year of intense fieldwork was performed in more than 40 caves from 14 karst units. Several new species for science were discovered and 7 taxa comprising 2 new genera and 5 new species were described.
Bearing in mind that spatial distribution of subterranean species is crucial to ecological research and conservation, the distribution of hypogean species, from Portuguese karst areas, was mapped using geographic information systems. Also, its subterranean richness was compared with other areas of the world and missing species were estimated on a regional scale. The subterranean biodiversity patterns were analyzed, and several factors were tested to explain richness patterns. Evapotranspiration and the consequent high productivity on the surface may be determinant in the species richness in the different karst units of Portugal, but the depth of the caves and the unique geological features of every massif seemed to play a more important role.
In order to evaluate the tolerance of organisms to groundwater contamination, the acute toxicity of two substances were tested on stygobiont crustaceans with different degrees of troglomorphism. Our study showed that the high levels of endemism contribute to remarkably different toxicological responses within the same genus.
The major problems related to conservation of subterranean habitats were associated to direct destruction and their contamination. These ecosystems lack of specific protection, implying an adequate management of surface habitats and the establishment of priority areas. Integrating all the previous information, this study establishes a ranking of sites for conservation of subterranean fauna in karst areas of Portugal.This research is a contribution to the study of subterranean biodiversity in karst areas of Portugal, towards its conservation.
The relative inaccessibility of the subterranean environment is a challenge for the study of its fauna, often accessible only in caves but more widely distributed. The subterranean animals are among the most rare, threatened and worldwide underprotected, often by the simple fact of being unknown.
Karst areas of Portugal occupy a considerable part of the territory and harbor more than 2000 caves. The complex biogeographical history of the Iberian Peninsula allowed the survival of several relict arthropod refugees in the subterranean environment.
Subterranean invertebrates have been ignored, as for as the protection of karst systems are concerned in Portugal, largely because knowledge was scarce and disorganized. Reviewing all the bibliographic sources about subterranean fauna from Portugal and listing troglobiont and stygobiont species and locations, was essential to understand the state of knowledge of species richness and the biogeography and conservation status for the studied areas.
In order to understand subterranean biodiversity patterns in karst areas from Portugal, one year of intense fieldwork was performed in more than 40 caves from 14 karst units. Several new species for science were discovered and 7 taxa comprising 2 new genera and 5 new species were described.
Bearing in mind that spatial distribution of subterranean species is crucial to ecological research and conservation, the distribution of hypogean species, from Portuguese karst areas, was mapped using geographic information systems. Also, its subterranean richness was compared with other areas of the world and missing species were estimated on a regional scale. The subterranean biodiversity patterns were analyzed, and several factors were tested to explain richness patterns. Evapotranspiration and the consequent high productivity on the surface may be determinant in the species richness in the different karst units of Portugal, but the depth of the caves and the unique geological features of every massif seemed to play a more important role.
In order to evaluate the tolerance of organisms to groundwater contamination, the acute toxicity of two substances were tested on stygobiont crustaceans with different degrees of troglomorphism. Our study showed that the high levels of endemism contribute to remarkably different toxicological responses within the same genus.
The major problems related to conservation of subterranean habitats were associated to direct destruction and their contamination. These ecosystems lack of specific protection, implying an adequate management of surface habitats and the establishment of priority areas. Integrating all the previous information, this study establishes a ranking of sites for conservation of subterranean fauna in karst areas of Portugal.
This research is a contribution to the study of subterranean biodiversity in karst areas of Portugal, towards its conservation.
The relative inaccessibility of the subterranean environment is a challenge for the study of its fauna, often accessible only in caves but more widely distributed. The subterranean animals are among the most rare, threatened and worldwide underprotected, often by the simple fact of being unknown.
Karst areas of Portugal occupy a considerable part of the territory and harbor more than 2000 caves. The complex biogeographical history of the Iberian Peninsula allowed the survival of several relict arthropod refugees in the subterranean environment.
Subterranean invertebrates have been ignored, as for as the protection of karst systems are concerned in Portugal, largely because knowledge was scarce and disorganized. Reviewing all the bibliographic sources about subterranean fauna from Portugal and listing troglobiont and stygobiont species and locations, was essential to understand the state of knowledge of species richness and the biogeography and conservation status for the studied areas.
In order to understand subterranean biodiversity patterns in karst areas from Portugal, one year of intense fieldwork was performed in more than 40 caves from 14 karst units. Several new species for science were discovered and 7 taxa comprising 2 new genera and 5 new species were described.
Bearing in mind that spatial distribution of subterranean species is crucial to ecological research and conservation, the distribution of hypogean species, from Portuguese karst areas, was mapped using geographic information systems. Also, its subterranean richness was compared with other areas of the world and missing species were estimated on a regional scale. The subterranean biodiversity patterns were analyzed, and several factors were tested to explain richness patterns. Evapotranspiration and the consequent high productivity on the surface may be determinant in the species richness in the different karst units of Portugal, but the depth of the caves and the unique geological features of every massif seemed to play a more important role.
In order to evaluate the tolerance of organisms to groundwater contamination, the acute toxicity of two substances were tested on stygobiont crustaceans with different degrees of troglomorphism. Our study showed that the high levels of endemism contribute to remarkably different toxicological responses within the same genus.
The major problems related to conservation of subterranean habitats were associated to direct destruction and their contamination. These ecosystems lack of specific protection, implying an adequate management of surface habitats and the establishment of priority areas. Integrating all the previous information, this study establishes a ranking of sites for conservation of subterranean fauna in karst areas of Portugal.This research is a contribution to the study of subterranean biodiversity in karst areas of Portugal, towards its conservation.
The relative inaccessibility of the subterranean environment is a challenge for the study of its fauna, often accessible only in caves but more widely distributed. The subterranean animals are among the most rare, threatened and worldwide underprotected, often by the simple fact of being unknown.
Karst areas of Portugal occupy a considerable part of the territory and harbor more than 2000 caves. The complex biogeographical history of the Iberian Peninsula allowed the survival of several relict arthropod refugees in the subterranean environment.
Subterranean invertebrates have been ignored, as for as the protection of karst systems are concerned in Portugal, largely because knowledge was scarce and disorganized. Reviewing all the bibliographic sources about subterranean fauna from Portugal and listing troglobiont and stygobiont species and locations, was essential to understand the state of knowledge of species richness and the biogeography and conservation status for the studied areas.
In order to understand subterranean biodiversity patterns in karst areas from Portugal, one year of intense fieldwork was performed in more than 40 caves from 14 karst units. Several new species for science were discovered and 7 taxa comprising 2 new genera and 5 new species were described.
Bearing in mind that spatial distribution of subterranean species is crucial to ecological research and conservation, the distribution of hypogean species, from Portuguese karst areas, was mapped using geographic information systems. Also, its subterranean richness was compared with other areas of the world and missing species were estimated on a regional scale. The subterranean biodiversity patterns were analyzed, and several factors were tested to explain richness patterns. Evapotranspiration and the consequent high productivity on the surface may be determinant in the species richness in the different karst units of Portugal, but the depth of the caves and the unique geological features of every massif seemed to play a more important role.
In order to evaluate the tolerance of organisms to groundwater contamination, the acute toxicity of two substances were tested on stygobiont crustaceans with different degrees of troglomorphism. Our study showed that the high levels of endemism contribute to remarkably different toxicological responses within the same genus.
The major problems related to conservation of subterranean habitats were associated to direct destruction and their contamination. These ecosystems lack of specific protection, implying an adequate management of surface habitats and the establishment of priority areas. Integrating all the previous information, this study establishes a ranking of sites for conservation of subterranean fauna in karst areas of Portugal.
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