This paper serves as an introduction to a multi-part monograph on the systematics of the subterranean amphipod genus Stygobromus, originally initiated by the late Dr. John R. Holsinger. The manuscript, left incomplete at the time of his death in 2018, was substantially completed and prepared for publication by the author, Julian J. Lewis, as a tribute to Holsinger’s lifelong contributions to subterranean biology. This first installment focuses on Stygobromus species from the Ozark, Arbuckle, and Ouachita mountains, extending into the glaciated plains of the central United States. The paper outlines the historical context of the work, the methodological challenges in preparing a decades-old manuscript for modern publication, and the taxonomic scope of the species treated. Special attention is given to preserving Holsinger’s original taxonomic intent and style, while acknowledging the limitations of pre-genetic systematics and the potential for cryptic species. This work represents a significant step toward completing Holsinger’s vision of a comprehensive treatment of Stygobromus systematics across North America.
An inventory of the paleontological resources in Virginia caves was initiated by the authors in 1996. Invertebrate and vertebrate fossil occurrences in Virginia were examined to understand the range and scope of fossils present in caves to aid in determination as to what should constitute cave significance in the documentation of paleontological resources for Virginia’s Significant Cave List. This cave study reviewed previously known vertebrate fossil sites and new vertebrate and invertebrate fossil sites before focusing on vertebrate fossil occurrences and the evidence of how they came to be deposited. As part of the inventory, to address the question of the percentage of Virginia caves that are paleontologically significant, 229 Virginia caves were examined to identify 100 caves with exposed non-human bones and/or teeth. Thirteen of the 100 caves yielded extinct or significant extirpated taxa, including one determined to have been excavated by E.D. Cope in 1867. Thirteen percent of the 100 bone caves examined or 5.7 % of the 229 study caves were documented as paleontologically significant. A selection of vertebrate cave fossils were radiocarbon dated and ranged in age from 11,986 +/- 76 to 39,300 +/- 1100 YBP. Uranium-thorium dating at one site suggests the fossil were older than 300,000 years and may be older than 450,000 YBP, indicating the age of this bone deposit is Pleistocene (Irvingtonian). A total of 43 cave sites were documented as paleontologically significant in the inventory and reported in Appendix C. PRIOVAC was recognized as an NSS Project from 2001 through 2006 and progress talks were presented at NSS Conventions in 1995, 1997, 1998, 2000, 2001, 2002, 2006, 2007, and a version of this summary in 2023 with abstracts published in the NSS Journal of Cave and Karst Studies and talks at the UIS Congresses in 1997 (Switzerland), 2001 (Brazil), and 2009 (US) published as papers in the UIS Proceedings. This paper is a summary report on PRIOVAC and the first publication of some of the bone dates and the U-Th dates.
Fossils accumulate in caves in multiple ways. Each introduces biases that must be understood before interpreting the assemblage. The ontogenetic age distributions of individual specimens of field mice, Peromyscus spp., based on tooth wear, provides insight into taphonomic pathways for two different talus deposits in Parker’s Pit, Black Hills, South Dakota. Main Cone accumulated under the modern 12 m vertical pit entrance throughout the Pleistocene and Holocene. The sink hole prohibits escape of randomly selected taxa. Red Cone formed as a talus slope in an old, now closed, entrance with a ramp-like slope that allowed animals to enter and exit the cave. Differences in ontogenetic age distributions between these two populations are statistically significant. Main Cone individuals show a wider and generally older age distribution than those from Red Cone. The older age distribution results from individuals surviving the vertical fall and living into old age in an essentially predator-free environment. These individuals probably subsisted on organic debris washed into the cave and resident invertebrates as observed in a living population of Peromyscus pectorralis in Longhorn Cavern, central Texas. The Red Cone population is ontogenetically younger, as exhibited by less dental wear, than the Main Cone population but slightly older than living populations studied in Washington, USA, and British Columbia, Canada. Bone damage indicative of predation by small carnivores, especially weasels (e.g., Mustela frenata) in the Red Cone sample is not apparent in the Main Cone population, reinforcing different taphonomic pathways of the two populations. The Red Cone sample is biased by predator selection. The Main Cone fossil assemblage likely represents a random, local sample of the biota around the cave that facilitates paleoecological analyses. The predator biased Red Cone sample is not amenable to reconstruction of communities but does provide insights into predator-prey interactions and selected species in the vicinity of the cave.
Stanton’s Cave is a large solutional cavern in the eastern region of Grand Canyon National Park, Arizona. The cave is best known to Quaternary paleontologists for its abundant Late Pleistocene and early Holocene fossil remains of mammals, birds, and Archaic artifacts. Re-examination of the fossil vertebrate collection, housed at the Grand Canyon Museum Collections, has resulted in re-identification of two fox dentaries (GRCA 76272) that were originally published as Urocyon cinereoargenteus (gray fox). Morphometric and morphological comparisons presented here find the dentaries diagnostic to Vulpes vulpes (red fox) and not the gray fox. The remains are the first record of Vulpes vulpes from the Grand Canyon and may be the first fossil evidence of the species from Arizona.
The first published inventory of paleontological resources associated with National Park Service (NPS) caves was issued in 2001 (Santucci et al. 2001). This report documented occurrences of fossil resources within caves in 35 NPS units and the challenges of managing and protecting these paleontological resources. Many significant NPS cave paleontology discoveries have been made in the following two decades. Highlights include: the first paleontological inventory of Carlsbad Caverns National Park, which documented both Permian fossils in cavern bedrock, and bones of Quaternary megafauna; reassessments of Quaternary fossils, particularly avifauna, from rock shelters and caves of Channel Islands NP; the discovery of thousands of Pleistocene-Holocene mummified bats from a cave in Grand Canyon NP, and the identification of an American cheetah skull from Rampart Cave; documentation of extinct or locally extirpated taxa from Great Basin NP; discoveries of abundant Mississippian fossils, particularly sharks, in cave bedrock of Mammoth Cave NP and Natchez Trace Parkway; the relocation of the Port Kennedy Bone Cave site in Valley Forge National Historical Park; and an inventory of Neotoma middens from across the NPS. The rise of photogrammetry as an investigative technique has been key to facilitating study of in situ specimens in these challenging settings. The possibility of paleontological resources in caves is now addressed during any paleontological inventory work in NPS units. We can expect that many more cave paleontology discoveries from NPS caves will be made in the decades to come.
Bats are the group of mammals most closely associated with caves. More than half of the 45 living species of bats from temperate North America (NA) are cave-dwellers in the families Vespertilionidae, Molossidae, Phyllostomidae, and Mormoopidae. Bats that inhabit caves also die in caves, leaving their skeletons to become preserved in fossil deposits on the cave floor. During the latter half of the Pleistocene epoch (-10 ka to 1 Ma), bats have an excellent fossil record in NA caves, primarily vespertilionids of extant species. We recognize only two extinct species of vespertilionids, Myotis rectidentis (late Pleistocene) and Corynorhinus alleganiensis (middle Pleistocene). Two other extinct bats are known from late Pleistocene cave deposits in the US: a large vampire, Desmodus stocki (Phyllostomidae), in 8 caves in the southern US, 4 in Mexico, and 6 karst deposits in Florida, and the extinct mustached bat, Pteronotus pristinus (Mormoopidae) of Cuba, but is also known from a karst deposit in Florida. The record of bats from late Pleistocene cave deposits in Mesoamerica consists of 11 caves in Mexico and 1 in Belize. The only other extinct Pleistocene bat in Mesoamerica besides Desmodus stocki is the giant vampire, D. draculae, known from Lolt & uacute;n Cave in Mexico and Cebada Cave in Belize. A diverse fauna of Quaternary bats is known from several hundred cave deposits across the West Indies, including more extinct species (eight) than are found in Pleistocene deposits in continental NA: six extinct species from Cuba, Pteronotus pristinus and Mormoops magna (Mormoopidae; also known from Hispaniola) and Artibeus anthonyi, Cubanycteris silvai, Phyllops silvai, and Phyllops vetus (Phyllostomidae); Tonatia saurophila (Phyllostomidae) known only from Jamaica; and Phyllonycteris major (Phyllostomidae) from Puerto Rico, Antigua, and Marie Galante. There are also 18 living species of bats that underwent local extinction on one or more Antillean islands during the Holocene, including the extant vampire bat Desmodus rotundus in Cuba. The oldest NA cave deposits containing fossil bats are early Pleistocene in age (-0.8-1.0 Ma) in West Virginia, Texas, Colorado, and New Mexico. NA bat fossil sites olderthan early Pleistocene are primarily paleokarst deposits (sinkholes, fissures, solution cavities) in peninsular Florida. The richest of these are: Inglis 1A (early Pleistocene,-2 Ma), with 7 species, including the oldest NA record of Desmodus and the only eastern NA record of Antrozous; Thomas Farm (early Miocene,-18 Ma), a sediment-filled sinkhole/cave complex containing 9 species including Primonatalus prattae, the oldest named species in the strictly cave-dwelling family Natalidae, Floridopteryx poyeri in the Emballonuridae; and 3 extinct genera of vespertilionids, Karstala, Miomyotis, and Suaptenos; and two Oligocene (26-30 Ma) fissure fills, I-75 and Brooksville 2, with 7 species of bats, including Koopmanycteris palaeomormoops, the oldest member of Mormoopidae; 2 species of Speonycteris representing an extinct family (Speonycteridae) with Neotropical affinities; and Oligopteryx floridanus and O. hamaxitos, the earliest New World emballonurids. Although karstic deposits are widely distributed in NA, they mostly lack fossil bats, except in Florida. We suspect undiscovered Tertiary karst deposits with bats exist elsewhere in NA.
We report on the fossils of the North American badger, Taxidea taxus (Carnivora) recovered from Snake Creek Burial Cave (SCBC), White Pine County, Nevada. The natural trap cave with its large funnel-shaped entrance at the base of the southern Snake Range, east-central Great Basin, contains deposits with radiocarbon ages that span from a median 8,709 cal yr BP to 48,028 cal yr BP. Skeletal remains of Taxidea are not overly abundant in the deposit compared to other more common carnivorans such as the canids (e.g., foxes). Measurements of the cranial and postcranial remains fall somewhere in the medium range relative to extant badger specimens. Thus, this badger appears to have been of the typical size of extant members and not a large variety emblematic of many Late Pleistocene mammals. Macrobotanical remains from packrat middens in the east-central Great Basin suggest that subalpine and montane conifers grew on the lower mountain slopes and valley bottoms above high lake stands of Lake Bonneville. Given the right substrate, select species of subalpine and montane conifers, along with sagebrush and other shrubs, likely formed a continuous or near-continuous belt from the Wasatch Front of Utah to the Sierra Nevada of California/Nevada. Continued work on the faunas and floras from caves and packrat middens in the Snake Range (e.g., Smith Creek, Ladder, Combustion, Cathedral, Arches Caves) and low in the valley (e.g., SCBC, Garrison Cave) will help in the reconstruction of the Late Pleistocene biotic communities above the high stands of Lake Bonneville and how the biotic communities, including the badger, adapted to climate change.
The most northerly excavated partial skeleton of a fossil jaguar (Panthera onca /augusta) from the USA (Oregon Caves National Monument, or ORCA) is described, illustrated and identified. Most of the individual bones preserved as fossils are compared with those found in living puma (Puma concolor) and jaguar (Panthera onca) skeletons to demonstrate the identification characters for each element and to establish their identification as those of a fossil jaguar. The carbon date on this specimen dates to 38,620 +/- 440 years BP, making it the oldest dated jaguar specimen to date, and within the Rancholabrean land mammal age. Its dentition is about average in size for a Rancholabrean jaguar, but its canine teeth are larger, suggesting a male. Eight localities (including ORCA) containing specimens of fossil jaguar are identified from the western USA., whereas six others containing felid specimens purported to be jaguar are identified as other felids. Given that there was no apparent bias against the preservation of felids in the fossil record in the western USA, it is suggested that there was a relative lack of the preferred prey species in the west (peccaries, armadillos or species associated with water such as turtles, caimans, capybaras or tapirs) compared to the southeast USA, where fossil jaguars are more frequently found.
We examine the Pleistocene record of fossil pronghorn (Mammalia: Antilocapridae) bones found primarily in caves throughout the Intermountain West of the United States and Mexico, but also in Florida, noting the different species present and their distribution in time and space. We briefly review the development of a framework for investigating the taphonomy of vertebrate fossils in caves. We review previously published explanations for pronghorn fossils found in caves and karst features. We then examine population structures, taphonomic factors, preservation biases, and abundance to identify patterns in the record that can resolve those different explanations. We focus on Pleistocene (Ice Age) sites known from Arizona, Colorado, Florida, Nevada, New Mexico, Texas, Wyoming, and in Mexico for the broader perspective, and on 4 species of pronghorn from late Pleistocene (Rancholabrean) cave sites for a narrower perspective. The taxa include: the extinct Stock’s pronghorn Stockoceros conklingi from Papago Springs Cave in southern Arizona, Shelter Cave and Muskox Caves in New Mexico and San Josecito Cave in Nuevo Leon, Mexico; the dwarf pronghorn, Capromeryx furcifer from multiple sites; an older species of dwarf pronghorn, Capromeryx arizonensis, from the early Pleistocene Inglis 1A site in Florida; and the extant pronghorn Antilocapra americana from Natural Trap Cave in Wyoming. We conclude that pitfall deaths account for those caves with large numbers of pronghorn fossils, while caves with smaller numbers of specimens are likely the result of carnivore activity, scavenging activity by rodents, or sporadic habitation by pronghorn.
Dietary remains recovered from Bassariscus (ringtail) midden deposits of middle Holocene age in the Weeping Cliffs, lower Grand Canyon, Arizona, indicate a diverse ringtail diet of plants and small vertebrates. Remains of fruits and seeds predominated among the plant parts, especially of hackberry, several cacti, and groundcherry. Highly fragmented remains of small vertebrates included anurans, lizards, snakes, bats, and rodents. Only one recovered vertebrate specimen exceeded 5 mm in length, necessitating meticulous morphological comparisons for taxonomic identifications, which were usually limited to generic level. Two radioisotopic dates on the middens range from ca. 7300 to 7900 cal yr BP. Vertebrate remains from the middens provide the first Quaternary records of amphibians (including Anaxyrus and Hyla/ Dryophytes) and a bat (Nyctinomops macrotis) from the Grand Canyon. Rare, fragmentary specimens provide one of only two fossil records of Dipsosaurus in Arizona.
La Tetera is a cave formed in the mid-late Paleozoic limestones of the Rincon Mountains of southern Arizona, USA. The cave was sealed since the late Pleistocene and preserves a small vertebrate fauna reflecting the Rancholabrean NALMA including two extinct large mammals, horse (Equus conversidens) and camel (Camelops hesternus), as well as an extinct vampire bat (Desmodus stocki). Additional recovered biotic remains include hackberry endocarps (Celtis), charcoal, and a harmonious assemblage of vertebrates including toad (Anaxyrus), tortoise (Gopherus), squamates (Heloderma, Dipsosaurus, Phrynosoma, Uta or Urosaurus, Aspidoscelis, Sonora, Rhinocheilus, Masticophis or Coluber, and Crotalus), roadrunner (Geococcyx), wren (Salpinctes), owl (cf. Athene), heteromyid and cricetid rodents, rabbits, Myotis sp., and shrew. Further investigation will likely reveal additional biotic remains. Preservation of bone is relatively poor, probably due to as-yet-undetermined, corrosive geochemical processes. Fossils occur as isolated skeletal elements scattered sparsely in several areas of the small cave or those recovered by screening of unconsolidated cave floor sediments (in which Desmodus is the second most commonly recovered taxon, after toads). A large, stratified paleoguano deposit in one room provides the potential to recover ancient environmental DNA from the bats, their dietary sources, and autochthonous and allochthonous microorganisms. A sample of the guano deposit gave a calibrated radiocarbon age of 23.7 ka, confirming a late Pleistocene age for the deposit and placing it within late Wisconsinan full glacial time and within Marine Isotope Stage 2. The radioisotopic age(s) of the vertebrate fossils are unknown. Several of the fossil vertebrates reflect a desertscrub fauna similar to that of the region today, and may reflect a southwestern lowland refugium for arid-adapted biota during the late Pleistocene.
The dolomitic limestone Pahasapa Formation exposed in the Black Hills of South Dakota is well known to contain a multitude of caves and other karst features. In order to raise greater awareness of the Black Hills caves and karst fissures as a source of paleontologic data, we summarize the major Quaternary and Pliocene cave and karst localities which contain fossils. The Richmond Hill quarry provides at least eight fissures deposits which contain a diverse range of vertebrate and invertebrate fossils of the Pliocene. Nine caves (Don’s Gooseberry Pit, Graveyard Cave, Jewel Cave, Parker’s Pit, Persistence Cave, Rushmore Cave, Salamander Cave, Stagebarn Cave, and Wind Cave) contain late Quaternary age fossils. Many of these faunas are part of on-going study. A record of climate and environmental change is emerging from these caves which illustrates that the Black Hills have been part of the dynamic changes throughout the Quaternary.
Exploration of both dry and submerged caves in the Yucatan Peninsula by cave divers has revealed a high diversity of Pleistocene giant ground sloths, representing all four families. Researchers working in the caves of Quintana Roo, Mexico, have identified up to five genera of sloths; a sixth genus is known from Belize. Three of the sloths, Nohochichak xibalbahka, Xibalbaonyx oviceps, and X. exenferis (if valid) are endemic to the peninsula. Preliminary dating indicates the genera were not all sympatric, with Nothrotheriops and a mylodontid occurring in the north during the last glacial maximum in what was likely an arid habitat, and with Eremotherium in forested central Belize. The northern part of the peninsula supported at least two megalonychids in earlier millennia in what was likely a more forested habitat. Better dating of known specimens is needed, along with stable isotope analyses, to clarify the history and ecology of sloths in the Yucatan.
Over thousands of years, a pit within Cueva del Tesoro in Queretaro, Mexico, has functioned as a death trap for mammals. This has resulted in the accumulation of mammalian fossils. At the bottom of this pit there is an almost complete and articulated skeleton of a ground sloth (Xenarthra, Folivora, Nothrotheriidae), Nothrotheriops shastensis. Remains of other animals include partial remains of at least a second ground sloth, a bovid, and bats. Given the extreme fragility and brittle nature of the remains, no attempt was made to extract or excavate them. Here we report our results mainly through photographic documentation in situ, with particular emphasis on dentary elements.
This study presents the results of monitoring air temperature, atmospheric pressure, relative humidity, air radon (222Rn) concentration and CO2 concentration in a huge cave, Torca del Carlista (-353 m). The cave ends in one of the largest halls in Europe, the Grupo de Espeleolog & iacute;a Vizcaino Hall, at-158 m. This hall has a volume of2.14 Mm3 and is connected to the outside by a narrow vertical tube 56 m long and 2 m wide. The objective was to determine the processes that control the airflow of this huge hall during a short time period (two weeks) with marked changes in the meteorological conditions (rapid shifts from cold and rainy to hot and dry days). The results show that the 12.5 h, 23.5 h and 23.6 h oscillations of CO2 and 222Rn are a response to Earth tidal forcing. We suggest that K1-O1 and M2-S2 Earth tides are detectable in the radon and CO2 concentrations of a natural cave. The poroelastic deformation under the tidal forcing could explain this signal. In consequence, the airflow in Torca del Carlista is a process that depends on thermo-barometric constraints, thermo-hygrometric conditions and Earth tides.
Here we describe a shallow-water karst ecosystem observed along the coast of Taiwan's Orchid Island, terrestrially-derived groundwater discharge into the coastal marine environment. Within these pools, dead coral formations dominated at land-adjacent locations, whereas mid- and sea-adjacent locations housed assemblages of corals, algae, and sponges. Several caves were found to house a rich variety of sponges, including 17 species that are potentially new to science. The sponge fauna included sclerosponges, which are hard, coral-like, atypical sponges that combine calcareous and siliceous skeletal parts. Other notable species included the recently described Vetulina incrustans and potentially new species belonging to the lithistid family, Theonellidae. Additionally, the caves housed red macroalgae and ahermatypic corals. The singular ecosystem currently receives no formal protection, and parts of it are adjacent to a nuclear storage waste facility. Overall, our study revealed that the cave and pool system houses a complex and diverse marine community of uncommon organisms, including a large number of potentially new species, highlighting its ecological significance.
The phreatic speleothems of hydrothermal speleogenesis are mainly mammillaries and spar calcite. In this study, seven samples from four caves in three areas of Greece are investigated. Morphological evidence from these caves along with the presence of the phreatic calcite indicate their hydrothermal origin. The analyzed calcite deposits contain layers of metallic oxides that were investigated with scanning electron microscopy aided by energy dispersive spectrometer semi-quantitative measurements. Their chemical composition is dominated by manganese. Rarely, double-terminated phantom calcite crystals display two repetitions of these manganese oxide layers along the growth surfaces, but most commonly mammillaries exhibit one to four manganese oxide layers. In one case, the manganese layer was formed along with partial dissolution of the calcite that resulted in small pits on the growth surfaces. We interpret these layers in the phantom spar crystals and cave mammillaries to changes in the fluid conditions and fluctuations that are common in phantom crystals result in the deposition of mangese oxides along with or without contemporary dissolution. In order to mark the changes in conditions of their formation, these layers are described as "manganese events." Because of the low deposition rate of the mammillaries, the repetitions of manganese events are plausibly independent of exogenous factors, such as seeping water. This event can be found in the secondary minerals of both shallow and deep phreatic settings of hydrothermal speleogenesis. The available data set shows that the manganese event is part of a repetitive depositional pattern that appears in at least 17 % of the carbonate speleothems of hydrothermal speleogenesis.
This article presents hydrological, hydrogeochemical, and mineralogical features of karstic lakes located in the basin of the disappearing, karstic Shulgan River (southern Ural, Russia). The lakes are currently isolated from karstic channels by bottom sediments; for several decades, their levels were relatively stable. We document a case of a "breakthrough" drainage event (occurring approximately once every 50 years), which occurred in 2021-2022 and the subsequent recovery of one of the lakes, Lake Ygyshma. Thanks to hydrological monitoring at the Goluboe Ozero spring and in the Shulgan-Tash Cave, it was possible to record the time of the breakthrough and the time of lake water transit, confirming the hydrogeological connection between lakes and the underground Shulgan River. This paper proposes a conceptual model of the evolution of isolated karstic lakes and the formation of the unconventional (for epigenic carbonate karst) hydrogeochemical characteristics in them (enrichment with metals, gases, etc.).
This paper presents the results of a study to identify the species and distribution of diatoms growing in various habitats and on different substrates in Hall's Cave, a limestone cave located on the Edward's Plateau in western Kerr County, central Texas, USA. A diverse diatom assemblage that includes at least seventeen taxa was found growing in low very low light conditions in different parts of the cave interior and on the walls of the cave entrance. Many other species were found in low numbers and may have been transported into the cave. One diatom species, Nitzschia palmida Carter is dominant in many samples, and overall, is the most abundant diatom observed in this investigation. This is the first published occurrence of N. palmida in the Western Hemisphere. These diatoms, all found typically in wet or damp settings, have adapted to growing in the cave environment. Nine species were common in only one sample, a scraping of wet guano-rich sediment on the floor at the back of the cave. These diatoms either survived passage through the digestive system of bats or were transported to the back of the cave by other means. The abundance and diversity diatoms in Hall's Cave suggests that diatoms may have been living in the cave for as long as there has been adequate moisture, nutrients and light, making them potential proxies for paleoenvironmental studies.
