The field of cave and karst science is served by a literature that is dispersed across far-flung topical journals, government publications, and club newsletters. As part of an inter-institutional project to globalize karst information (KIP, the Karst Information Portal), the USF Library undertook a structured battery of literature searches to map the domain of karst literature. The study used 4,300 individual searches and four literature databases: GeoRef, BIOSIS, Anthropology Plus, and GPO Access. The searches were based on a list of 632 terms including 321 karst-related keywords culled from three leading encyclopedias and glossaries of cave and karst science. An examination of yearly changes in publication rate indicates that for the last 45 years, the number of cave and karst publications has increased steadily, as has the number of journals in which they appear. In particular, the past ten years cover a period of rapid growth where karst-specific journals achieved peer-review status, and individual journals accepted more cave and karst papers for publication.
Air-filled caves surveyed in the Brooksville Ridge of west-central Florida provide insight into the organization of karstic permeability within the unconfined portions of the Upper Floridan Aquifer. The morphology of the passages that compose these caves in geologically young, high-permeability limestones is strikingly different from caves found in ancient carbonates far from the influence of the coast. Cave passages in west-central Florida are laterally extensive and tiered. Principal horizons of cave development occur between +3 m and +5 m, +12 m and +15 m, and +20 m and +22 m above modern sea level. The primary guide of cave passage orientations within these cave levels is widespread fractures oriented approximately NE-SW and NW-SE. Cave passages of human dimensions form at the intersection of the laterally extensive cavities and fractures and often acquire a characteristic plus-sign shape. The walls of cave passages in west-central Florida are porous and complex, with small-scale solution features such as pockets and tafoni structures extending into the host bedrock. Additionally, these cave passages often end in blind pockets, ever-narrowing fissures, sediment fills, and collapses. The passages do not appear to represent an integrated system of conduits between aquifer inputs and outputs.
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 (allozyrne electrophoresis and mitochondria1 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 panrnictic 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 lo6 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 Wester,,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 fi-om understanding and integration with abiotic groundwater system processes, especially hydrogeologic and geomorphic processes.
Carbon dioxide concentrations in Ballynamintra Cave, S. Ireland, generally increase with distance from the entrance, but this trend is non-linear because physical constrictions and slope changes compartmentalize the cave into zones with distinct PCO2 signatures. In this cave, CO2 originates from the soil and enters the cave by degassing from dripwater and by seeping through fractures, and is then transported throughout the cave by advection. Elevated concentrations in roof fissures, joints, and adjacent to walls suggest that these locations shelter CO2 gas from advection and permit local accumulation. CO2 enrichment was noted over a sediment accumulation, suggesting that microbial oxidation of organic compounds in the sediment provided an additional CO2 source distinct from the soil zone above the cave. Advection driven by external barometric pressure variations caused ventilation, which is the principal CO2 sink. The data presented here underscore the need for high resolution data to adequately characterize cave air PCO2 variability.
The Kentucky GIS coverage of sinkholes, completed in 2003, consists of 101,176 polygons representing the uppermost closed contour of every karst sinkhole identified using USGS 1:24,000 scale topographic maps. This resource is a useful tool for delineating karst landscapes in Kentucky because karstified limestones underlie 55% of the areal surface of the state. For hydrologic studies, alignments of sinkholes commonly indicate preferential flowpaths for groundwater; and this information aids in large-scale planning and zoning. In this paper, I demonstrate the effectiveness of using this sinkhole coverage as a tool for delimiting structural features of Kentucky.
The Bonne Femme watershed, located in central Missouri, is a karst watershed in a rapidly urbanizing area. This study was undertaken to characterize the hydrology of two karst aquifers within this watershed before significant increases in impervious surfaces take place. The specific objectives of this study were to: 1) use dye tracing to delineate the recharge area for Hunters Cave (HC); 2) quantify and summarize annual and monthly stream discharge at the resurgence of HC and Devils Icebox (DI) caves; and 3) characterize the chemical and physical status of the cave streams relative to temperature, pH, specific conductance, dissolved oxygen, and turbidity. The quantity and quality of the water at the resurgence of both cave streams was monitored from April 1999 to March 2002. Both recharge areas were determined to be of similar size (33.3 km² for HC and 34.0 km² for DI) and were formed in the same geologic strata. Average annual discharge was 55,900 m³ km-² at DI and 35,200 m³ km-² at HC. Relative discharge, as a percent of annual precipitation, averaged 6.1% at DI and 3.8% at HC. Average monthly discharge was 2,930 m³ km-² at HC and 4,650 m³ km-² at DI; however, median instantaneous discharge over the three years was about 18% higher at HC (74 m³ h-¹) compared to DI (63 m³ h-¹). Turbidity and pH showed the largest differences between sites over the three years. The higher turbidity and lower pH at DI reflected the greater magnitude and duration of runoff events for this system. The physical characteristics of the two recharge areas explained the observed differences in discharge. The HC recharge area is characterized by limited sub-surface conduit development, small conduits, short flow paths from surface to resurgence, and predominantly allogenic recharge. The DI recharge area is characterized by extensive sub-surface conduit development, large conduits, long flow paths to the resurgence, and autogenic and allogenic recharge.
This paper presents newly discovered candidate cave entrances into
Martian near-surface lava tubes, volcano-tectonic fracture systems, and
pit craters and describes their characteristics and exploration
possibilities. These candidates are all collapse features that occur
either intermittently along laterally continuous trench-like depressions
or in the floors of sheer-walled atypical pit craters. As viewed from
orbit, locations of most candidates are visibly consistent with known
terrestrial features such as tube-fed lava flows, volcano-tectonic
fractures, and pit craters, each of which forms by mechanisms that can
produce caves. Although we cannot determine subsurface extents of the
Martian features discussed here, some may continue unimpeded for many
kilometers if terrestrial examples are indeed analogous. The features
presented here were identified in images acquired by the Mars Odyssey's
Thermal Emission Imaging System visiblewavelength camera, and by the Mars
Reconnaissance Orbiter's Context Camera. Select candidates have since
been targeted by the High-Resolution Imaging Science Experiment. Martian
caves are promising potential sites for future human habitation and
astrobiology investigations; understanding their characteristics is
critical for long-term mission planning and for developing the necessary
exploration technologies.
Cosmogenic burial dating of quartzose cave sediments deposited in multilevel caves beneath the western margin of the Cumberland Plateau dates ∼5.7 Ma of cave development in step with episodic incision of the Upper Cumberland River. These particular cave systems are characterized by hydrologically abandoned, low-gradient passages concentrated at common levels above the modern water table. Previous studies recognized morphometric differences between the majority of smaller hydrologically active "plateau-margin" caves and large, abandoned "fossil" or "Cumberland-style" caves. This study links the origin of multilevel caves on the western margin to a prolonged period of Late Tertiary water table stability, and the development of levels to distinct episodes of Plio-Pleistocene river incision. In this study, clastic sediments in multilevel cave passages are dated using cosmogenic 26Al and 10Be, and are shown to correspond with 1) deposition of upland ("Lafayette-type") gravels between ∼3.5 Ma and ∼5 Ma; 2) initial incision of the Cumberland River into the Highland Rim after ∼3.5 Ma; 3) development of the Parker strath between ∼3.5 Ma and ∼2 Ma; 4) incision of the Parker strath at ∼2 Ma; 5) shorter cycles of incision after ∼1.3 Ma associated with terraces above the modern flood plain; and 6) regional aggradation at ∼0.8 Ma. Burial ages of cave sediments record more than five million years of incision history within the unglaciated Appalachian Plateaus and constrain the developmental history of multilevel caves associated with the Upper Cumberland River.
Serpents Cave, located in the French Alps, contains a sulfidic-thermal (41 degrees C) karst spring. Degassing of the sulfidic vapor produces diverse sulfate minerals. The reaction with the limestone host-rock produces gypsum, anhydrite, sulfur, and magnesium calcite. The reaction with an artificial material (aluminum door) produces alunogen, tschermigite, and jurbanite. Microbial activity is suspected in the genesis of sulfur and tschermigite. Aluminum sulfates have usually been reported in mines, in volcanic settings, and in rock-shelters in phyllites. Some of these alum minerals such as tschermigite are rarely observed in eaves, and jurbanite is identified here for the first time in a cave. Serpents Cave is therefore an important site for sulfate minerals in caves, even if the aluminum sulfates should be considered border minerals because they originate from sulfur vapor reaction with artificial media.
Like most other branches of speleology, cave archaeology in the U.S. grew and developed significantly during the mid to late twentieth century. Originally viewed as marginal to mainstream Americanist archaeology, pursuit of prehistoric and historic archaeology underground is now widely accepted as making valuable contributions to knowledge of human past. The National Speleological Society played a central role in that development and continues to do so. We outline the establishment and growth of cave archaeology in North America, with special emphasis on relations between the NSS and archaeology performed in dark zone, deep cave interiors.
A new species, Plutomurus birsteini sp. n., from Georgian caves is described and illustrated. It is similar to Plutomurus baschkiricus (Skorikow, 1899). Differences between the species are discussed. A key to the genus Plutomurus species found in the Caucasus is provided.
Since 2006, a white fungus named Geomyces destructans has been observed on the muzzles, noses, ears, and (or) wings of bats in the eastern United States, and bat colonies that are infected with this fungus have experienced dramatic incidences of mortality. Although it is not exactly certain how and why these bats are dying, this condition has been named white-nose syndrome (WNS). WNS appears to have spread from an initial infection site at a cave that is connected to a commercial cave in New York, and by the end of August 2009 was identified in at least 74 other sites in the eastern United States. Although detailed geographical and geological data are limited, a review of the available data shows that sites infected with WNS before September 2009 include both natural caves and mines. These infected sites extend from New Hampshire to Virginia, and known site elevations range from 84 to 2693 feet above sea level. In terms of geological setting, the infected sites include sedimentary, metamorphic, and igneous rocks of ages ranging from Precambrian to Jurassic. However, by the end of August 2009, no infected sites had been identified in strata of Mississippian, Cretaceous, or Triassic age. Meteorological data are sparse, but most of the recorded air temperatures in the known WNS-infected caves and mines range from 0 to 13.9 degrees C, and humidity measurements range from 68 to 100 percent. Although it is not certain which environmental parameters are important for WNS, it is hoped that the geographical and geological information presented in this paper will inform and clarify some of the debate about WNS, lead to greater understanding of the environmental parameters associated with WNS, and highlight the paucity of scientific data from caves in the eastern United States.
Karst is a specific type of terrain (or landscapes) with characteristic suites of well-known surface and subsurface dissolutional features. The latter result from integrated subsurface drainage. A variety of nondissolutional processes forms terrains analogous to certain types of karst; these are termed pseudokarst. Before 1906, these generally were believed to be karst somehow formed in poorly soluble rocks. They share a considerable range of features, resources and values with karst, commonly (but not invariably) including eaves, and the two are linked across a wide spectrum of processes and features (e.g., between dissolutional and piping caves). Unlike karst, integrated subsurface drainage may not be present. Isolated caves define neither karst nor pseudokarst. Multiprocess terrains and landscapes are not uncommon. Based largely on conclusions of a working session of the 1997 International Congress of Speleology, eight types of pseudokarst are identified, with notably different implications for extraterrestrial habitats: rheogenic pseudokarst, glacier pseudokarst, badlands and piping pseudokarst, permafrost pseudokarst, talus pseudokarst, crevice pseudokarst, compaction pseudokarst and consequent pseudokarst. Some appear to exist on Mars. Speleologists expert in their differentiation should serve as consultants to planetary geologists.
Organic compounds derived from plants are found in many cave formations, which are collectively termed speleothems. Both the carbon in the organic compounds and the carbon in the speleothem CaCO3 have distinct ratios of the stable isotopes of carbon (12C and 13C that are expressed as δ13C values. Values of δ13C in the organic compounds are lower than δ13C values of speleothem calcium carbonate and could affect the δ13C values of speleothems with high organic matter concentrations, if the organic compounds were not removed. Four treatments conventionally used to destroy organic matter in carbonates prior to geochemical analysis were evaluated in this study. The treatments were oven roasting at 340° C, soaking in bleach, soaking in 30% H2O2, and soaking in distilled deionized water. There is no statistically significant difference between results from untreated and treated samples. These results suggest that the treatments do not affect the δ13C value of speleothems' calcium carbonate. The treatments might be helpful in removing organic matter in speleothems that have high concentrations of organic matter. However, most speleothems have low organic carbon concentrations that do not affect the δ13C value of the speleothem, even if left untreated. Ultimately these treatments only need to be applied to speleothems with unusually high concentrations of organic matter.
Abaco Island, Bahamas, contains a large number of conical hills that strongly resemble cone karst, a dissolutional landform associated with karst processes in tropical carbonate localities around the world. Field investigation demonstrated that the conical hills are roughly symmetrical in shape and consist of mid to late Pleistocene eolian calcarenites (carbonate sand dunes) partly mantled by talus. The original hummocky depositional topography of the eolian ridges has been dissected by a variety of processes including dissolution pit formation that enhances slope failure, vegetative disruption of the rock surface, and fire-induced exfoliation of the bedrock. The original asymmetrical shape of continuous dune ridges has thus been modified into a series of hills with a symmetrical cone shape, in which the steeper leeward depositional slopes of the dunes have been masked by talus to create a lesser slope that approximates the dip of the exposed bedrock slope of the more gentle windward depositional faces of the dunes. The conical hills are primarily constructional in nature, modified by mass-wasting slope processes only partly influenced by dissolution; therefore these conical hills are not true cone karst, but pseudokarst, despite their cone shape and their development in soluble carbonate rock. Abaco is the only island in the Bahamian Archipelago with both high eolian relief and a significant positive water budget; that water budget supports the degree of dissolution and forest growth necessary to create the root wedging and fire-induced exfoliation that most modify the steep leeward slopes of the dunes. As the eolian calcarenites were deposited only during the brief glacioeustatic sea-level highstands of the last few hundred thousand years, the development of cone-shaped hills in these eogenetic rocks has been geologically very rapid.
Observations and hypotheses on the possible influence of unidentified calcifying bacteria on moonmilk speleothem formation in the Grotta Nera are reported for the first time. The Majella Massif hosts a complex karst system of several caves; the accessible Grotta Nera is the most interesting one. Despite its name, the cave is characterized by particularly abundant ivory-white deposits of moonmilk. Two samples of moonmilk were analyzed to determine the geochemistry, fabric, depositional setting, and extent of biogenicity. For this, we combined geochemical, scanning electron microscopic, microbiological, and in vitro precipitation studies. X-ray diffraction of the moonmilk deposits gave clear evidence for the presence of calcite. Scanning electron microscopy showed that moonmilk in the Grotta Nera consists of a network of calcite fibers oriented in all directions, resembling a felted mat. The cultivation on specific medium of moonmilk and drip-water samples showed the presence of fungi, actinomycetes, and other bacteria, but the dominant cultivable microorganisms were bacteria, which produced significant crystallization. Examination of Gram-stained smears taken from the fifteen different colony types showed that the majority (66.7%) of the bacterial isolates were Gram-negative. Single small rods and rod chains were the most common bacteria isolated from the Grotta Nera. None of the molds isolated from the Grotta Nera samples were able to precipitate CaCO3 crystals, suggesting a major bacterial contribution to moonmilk deposition in the cave. Bacteria were capable of precipitating CaCO3 on B-4 solid medium at 15 (cave temperature), 22, and 32 degrees C. The calcifying bacteria isolated from the Grotta Nera showed a greater capability to solubilize CaCO3 than those associated with consolidated stalactites sampled from previously studied caves. The electron microscopy and microbiological evidences, together with the geochemistry and environmental data, allowed us to postulate the biogenic nature of the moonmilk in the Grotta Nera Cave.
We compare micro-charcoal abundances in laminated cave-stream sediments to the presences of Native Americans and later settlers in the same watershed. Samples were obtained from a core taken from a 2.5 m high point bar located 1 km inside of Buckeye Creek Cave, West Virginia. Thirty-three subsamples were treated with hydrogen peroxide to bleach or whiten non-charcoal organic matter. In the absence of opaque mineral grains, this technique creates a large visual contrast between dark charcoal grains and other substances. The subsamples were photographed using a microscope-mounted camera, and pixels darker than 99/255 (grayscale) were used to calculate charcoal concentrations. The record spans the last 6,000 years, and four of the five highest charcoal concentrations are from the last 2,000 years. The highest concentration is from AD 1093, and the second-highest concentration is from the nineteenth century. Post-Colonial settlers began making extensive use of the watershed sometime in the eighteenth century and may, therefore, be responsible for the second-highest charcoal concentration. However, archaeologists independently concluded that Native Americans made peak use of the watershed between AD 1000 and 1200, which coincides with the highest charcoal concentration in the record. Native Americans are known to have extensively used fire, so there is good circumstantial evidence tying high concentrations in the last 2,000 years to human activities. Our method is suitable for use elsewhere, and we present a detailed statistical analysis of our data as a guide toward interpreting charcoal concentrations in karst and non-karst deposits.
A new genus and two new species of ereynetid mites, one edaphobitic, the other troglobitic, are described from three European karst areas. Free-living species of the Riccardoella-complex exhibit ramified barbules in guard setae associated with tarsal solenidia, whereas parasitic species lack these characters. Ramified barbules in guard setae are thus considered specific adaptations to soil habitat. Free-living species of the Riccardoella-complex are seemingly restricted to karstic and other calcareous-rich areas while parasitic species live exclusively on slugs and snails. The relationship between calcium and Riccardoella-complex mites is discussed. A key is provided for the genera of Ereynetinae.
Anelpistina specusprofundi, n. sp., is described and separated from other species of the subfamily Cubacubaninae (Nicoletiidae: Zygentoma: Insecta). The specimens were collected in So  tano de San Agustõ Ân and in Nita Ka (Huautla system) in Oaxaca, Me Âxico. This cave system is currently the tenth deepest in the world. It is likely that A. specusprofundi is the sister species of A. asymmetrica from nearby caves in Sierra Negra, Puebla. The new species of nicoletiid described here may be the key link that allows for a deep underground food chain with specialized, troglobitic, and comparatively large predators such as the tarantula spider Schizopelma grieta and the 70 mm long scorpion Alacran tartarus that inhabit the bottom of Huautla system.
Previous reports of reticulated filaments, an unknown microbe, document that they are ubiquitous in subsurface environments, including limestone caves, lava tubes, and even granite tunnels. Although initial reports of fossil reticulated filaments described preserved organic matter, additional instances involve replacement by calcite, Mn-oxides, silica, or copper silicates. We report on living reticulated filaments found in the limestone Herbstlabyrinth-Adventhöhle Cave System, Hesse, Germany. Samples from soft pool-fingers, pool-bottom clays, and clay-coated rocks along the flow path of incoming water all contain living reticulated filaments associated with abundant biofilm. Most of the reticulated filaments are approximately 0.5 mm in diameter, reach lengths between 150 and 200 mm, and have irregular chambers with spines, a newly identified morphological variant. EDX of these filaments confirms an organic composition not replaced by minerals. They are the dominant visible microbial form in these biofilms, providing hope that they can be isolated and identified.
White-nose syndrome is an emerging infectious disease that has led to a dramatic decline in cave-hibernating bat species. White-nose syndrome is caused by the newly described fungal pathogen Geomyces destructans, which infects the ear, muzzle, and wing membranes of bat's. Although the exact mechanism by which the fungus causes death is not yet understood, G. destructans leads to a high mortality rate in infected animals. While the primary mechanism of infection appears to be bat-to-bat transfer, it is still unclear what role human activity may play in the spread of this pathogen. Here we evaluate the effectiveness of decontamination protocols that can be utilized by speleologists to reduce the likelihood of spreading this dangerous pathogen to naive bats or uninfected hibernacula. Our results show that pre-cleaning to remove muds and/or sediments followed by the use of commercially available disinfectants can effectively remove G. destructans from caving fabrics. Alternatively, immersion in water above 50 degrees C for at least 20 minutes effectively destroys the fungal spores. These results have allowed the development of a decontamination protocol (http://www.fws.gov/WhiteNoseSyndrome/cavers.html) that, when appropriately followed, can greatly reduce the likelihood of the human mediated transfer of G. destructans from an infected to uninfected site.
A new Agraphorura species is described from North American caves (Idaho, USA) and seven additional taxa are referred to this genus. The new species can be distinguished from congeners by the unusual structure of antennal III sense organ. An identification key for all of the known species of Agraphorura is provided.
