Cave-dwelling
Guides
Emesa
thread-legged bugs
Emesa is a small genus of thread-legged bugs (subfamily Emesinae) in the assassin bug family Reduviidae. The genus was established by Fabricius in 1803 and contains only four described species. Members of this genus share the characteristic elongated, slender bodies and extremely long legs typical of Emesinae.
Emesinae
Thread-legged Bugs
Emesinae, commonly known as thread-legged bugs, are a subfamily of assassin bugs (Reduviidae) distinguished by their exceptionally slender body form. They are predatory insects that walk on their mid and hind legs while using their raptorial front legs to capture prey. The subfamily contains approximately 90 genera and 900 described species, with greatest diversity in tropical regions, particularly Africa. Some groups specialize on spiders as prey. Molecular phylogenetic analyses have questioned the monophyly of Emesinae, suggesting the group may be polyphyletic with respect to Saicinae and Visayanocorinae.
Emesini
thread-legged bugs
Emesini is a tribe of thread-legged bugs (Reduviidae: Emesinae) characterized by extremely slender bodies and elongated legs. The tribe contains multiple genera distributed worldwide, with some species exhibiting specialized cave-dwelling habits. Members of this tribe include both surface-dwelling and cavernicolous species, with documented arachnophilous behavior in several taxa.
Entomobrya zona
Rocky Mountain Springtail
Entomobrya zona, commonly known as the Rocky Mountain springtail, is a small springtail species endemic to the Rocky Mountains and known from three caves in the Grand Canyon. Adults reach approximately 2 mm in length and are characterized by tan coloration with black markings, including a distinctive wide dark band running down the center of the abdomen. The species occurs at elevations from 1,700 to over 3,000 meters in arid environments and coniferous forests. It is frequently found in subterranean habitats, where it is considered a troglophile.
Entomobryidae
slender springtails
Entomobryidae, commonly known as slender springtails, is a large family of springtails (Collembola) containing over 1700 described species. Members are characterized by an enlarged fourth abdominal segment, a well-developed furcula (springing organ), and often possess prominent body scales that can produce striking coloration including blue, red, purple, or white patterns. The family includes both scaled and scale-less forms, with the latter frequently captured in pitfall traps globally. Many species are relatively large for springtails, reaching 2 mm or more in length.
Euhadenoecus fragilis
Tawney's Cave Cricket, tawneys cave cricket
Euhadenoecus fragilis is a species of camel cricket in the family Rhaphidophoridae. It was described by Hubbell in 1978. The species is known from cave environments in North America, with documented occurrences in Virginia. Like other members of the family Rhaphidophoridae, it is wingless and adapted to subterranean habitats.
Euhadenoecus insolitus
McCluney Cave Cricket, Highland Rim Crevice Cricket
Euhadenoecus insolitus is a species of camel cricket in the family Rhaphidophoridae, described by Hubbell in 1978. Populations exhibit flexible reproductive strategies, with some reproducing sexually and others through parthenogenesis. The species regularly forages outside cave habitats during warmer months but remains cave-bound in winter.
Geophilus
Compost Centipedes
Geophilus is a large genus of soil centipedes in the family Geophilidae, with approximately 140 species. The genus has a Holarctic distribution and exhibits considerable morphological diversity, with body lengths ranging from under 1 cm to over 7 cm and leg pair counts varying from 29 to 89. Several species have adapted to specialized habitats, including deep cave environments. The genus is characterized by distinctive structural features of the head, forcipules, and trunk segments.
Glomeridae
pill millipedes
Glomeridae is a family of pill millipedes in the order Glomerida, comprising over 300 species distributed among approximately 30 genera. Members are characterized by their ability to conglobate (roll into a complete sphere) as a defensive mechanism. The family has a primarily Palearctic distribution with significant diversity in Southeast Asia, and includes both surface-dwelling and cavernicolous species. Many species remain undescribed, particularly in tropical regions.
Grylloblatta chirurgica
Mount Saint Helens Grylloblattid, Mount Saint Helens grylloblatid
Grylloblatta chirurgica is a species of ice crawler endemic to southwestern Washington state, specifically associated with high-elevation habitats and ice caves near Mount St. Helens in Skamania County. Described by Gurney in 1961, it is one of the rarest and most geographically restricted members of the relict order Notoptera (Grylloblattodea). The species persists in cold, stable microhabitats including glacial ice fields and subterranean ice formations. Its extremely limited distribution and specialized habitat requirements make it vulnerable to environmental change.
Grylloblattidae
ice crawlers, icebugs, rock-crawlers
Grylloblattidae, commonly known as ice crawlers or icebugs, is a family of wingless, cold-adapted insects representing the sole living family of the order Grylloblattodea. These relict insects inhabit extreme cold environments including glacier margins, alpine snowfields, caves, and subalpine forests across disjunct regions of the Holarctic. With approximately 35 extant species in six genera, they exhibit narrow physiological temperature tolerances and are considered endangered due to climate warming and habitat specificity.
Hadenoecus opilionides
Tennessee cave cricket
Hadenoecus opilionides, commonly known as the Tennessee cave cricket, is a camel cricket species described by Hubbell in 1978. It belongs to the family Rhaphidophoridae, a group of wingless crickets specialized for life in dark, humid environments. The species is known from two recognized subspecies: H. o. australis and H. o. opilionides. Records indicate it occurs in Tennessee and broader North America.
Hadziidae
Hadziidae is a family of amphipod crustaceans established by S. Karaman in 1943. Members are taxonomically difficult to distinguish from the closely related family Melitidae. The family includes both marine and freshwater-dwelling species, with some lineages having colonized subterranean habitats.
Haplophthalmus danicus
Spurred Ridgeback, terrestrial cave isopod
Haplophthalmus danicus is a small woodlouse species in the family Trichoniscidae, commonly known as the spurred ridgeback or terrestrial cave isopod. Native to Europe and parts of Asia, it has been introduced to North America, where it has become well established in terrestrial communities since European settlement. The species comprises seven recognized subspecies across its native range. It is frequently observed in cave and subterranean habitats, reflecting its common name.
Heleomyzidae
Sun flies, Heleomyzid flies
Heleomyzidae is a family of small to medium-sized flies in the order Diptera, comprising over 740 described species in approximately 76 genera. The family is distributed worldwide, with the greatest diversity in the Holarctic region. Adults are frequently observed at lights in early spring and late fall. Larval ecology is diverse, with many species associated with decaying organic matter, fungi, carrion, or bird nests.
Heteraphorura
Heteraphorura is a genus of springtails in the family Onychiuridae, first described by Bagnall in 1948. As a member of the tribe Hymenaphorurini, it belongs to a group of euedaphic (deep soil-dwelling) Collembola characterized by reduced or absent eyes and elongated body forms. The genus has been recorded across multiple continents including Europe, Mediterranean regions, northern Eurasia, and parts of North America.
Heterelmis
riffle beetles
Heterelmis is a genus of aquatic beetles in the family Elmidae, commonly known as riffle beetles. The genus includes several described species, with Heterelmis comalensis being particularly notable as an endangered species endemic to spring systems in Texas. Members of this genus are fully aquatic, inhabiting oxygen-rich flowing water environments where they feed on microbial biofilms. Some species have reduced or non-functional wings, limiting their dispersal ability and making them vulnerable to habitat degradation.
Hubbardia pentapeltis
short-tailed whipscorpion
Hubbardia pentapeltis is a species of short-tailed whipscorpion in the family Hubbardiidae, first described by Cook in 1899. It belongs to the order Schizomida, a small group of arachnids characterized by their compact bodies and reduced flagellum. The species is known from North America and has been documented in citizen science observations.
Hypochilus bonneti
lampshade weaver
Hypochilus bonneti is a species of lampshade spider (family Hypochilidae) first described by Gertsch in 1964. It is one of several species in the genus Hypochilus, which are known for their distinctive flat, circular webs that resemble lampshades. The species occurs in the United States, though specific details about its biology and ecology remain poorly documented.
Hypochilus kastoni
Hypochilus kastoni is a species of lampshade spider in the family Hypochilidae, described by Norman Platnick in 1987. It is one of several species in the genus Hypochilus, a group of spiders notable for their distinctive web architecture and relictual distribution in North America. The species is known from the western United States, where it inhabits rocky habitats. Like other Hypochilus species, it constructs characteristic flat, circular webs with a conical retreat, resembling a lampshade in appearance.
Hypogastrura
snow fleas
Hypogastrura is a large genus of springtails in the family Hypogastruridae, comprising at least 150 described species with a cosmopolitan distribution. The genus includes the well-known "snow fleas" such as Hypogastrura nivicola, which are conspicuous on snow surfaces in winter. Members are small, typically 1–3 mm, with the characteristic springtail body plan including a furcula for jumping and a ventral collophore. The genus occupies diverse habitats from Arctic tundra to caves and forest floors.
Ischyropsalididae
Ischyropsalidid Harvestmen
Ischyropsalididae is a family of harvestmen (Opiliones) comprising 35 described species across three genera: Acuclavella (thorn harvestmen of North America), Ceratolasma, and Ischyropsalis (predominantly European). The family exhibits a disjunct distribution spanning Europe and western North America. Several species are troglophilic, inhabiting cave systems with documented adaptations to subterranean environments including seasonal activity patterns and microhabitat specialization.
Jacobsoniidae
Jacobson's beetles
Jacobsoniidae is a small family of minute beetles within the superfamily Staphylinoidea, comprising three extant genera (Derolathrus, Sarothrias, Saphophagus) and approximately 28 described species. Adults and larvae inhabit cryptic microhabitats including leaf litter, rotting wood, fungal fruiting bodies, bat guano, and caves. The family exhibits remarkable morphological stasis since the Cretaceous, with fossil records from Cretaceous amber of Myanmar and France, Eocene Baltic amber, and Holocene copal. Members are among the smallest beetles, measuring 0.7–2.1 mm, and are exceptionally rare in collections.
Keroplatini
Keroplatini is a tribe of fungus gnats within the family Keroplatidae (Diptera). The tribe includes the genus Chetoneura, which was formally transferred from Orfelia to Keroplatini based on phylogenetic analysis. At least one member, Chetoneura shennonggongensis, is a cave-dwelling species with documented larval biology involving silk construction for prey capture.
Laemostenus
Laemostenus is a genus of ground beetles (Carabidae) comprising nearly 200 species distributed across all continents except Antarctica. Species range from 8 to 28 mm in length. Many exhibit dark coloration with blue or purple metallic sheen, while subterranean and cave-dwelling species show depigmentation, reduced eyes, and lighter coloration. The genus includes both surface-dwelling and hypogean (subterranean) species, with several subgenera recognized.
Leptonetidae
cave spiders
Leptonetidae is a family of small, primitive haplogyne spiders first described by Eugène Simon in 1890. The family comprises approximately 22 genera and 400 species, though taxonomic boundaries have been revised recently with elevation of Archoleptonetinae to family rank (Archoleptonetidae). Leptonetids are characterized by their small size (2-5 mm), reduced eye number, and adaptation to dark, moist microhabitats. They represent a relict fauna with origins dating to the Cretaceous period and exhibit complex biogeographic patterns across the Holarctic.
Liocranidae
sac spiders
Liocranidae is a family of araneomorph spiders first described by Eugène Simon in 1897. They are commonly referred to as 'sac spiders' due to their habit of constructing silk retreats. The family contains 35 genera and approximately 357 species as of January 2026. The holarctic genus Agroeca is the best-known member, while many other genera remain poorly studied and lack formal diagnoses. Recent taxonomic work has significantly expanded the family, including the description of numerous new species from China, Vietnam, and other regions.
Lithobiomorpha
Stone Centipedes
Lithobiomorpha, commonly known as stone centipedes, are an order of anamorphic centipedes characterized by a mature segment count of 15 trunk segments. They lack compound eyes, possessing instead simple ocelli or no eyes in some subterranean species. The order includes two families: Henicopidae and Lithobiidae. Members are primarily found in forest litter and soil habitats across temperate regions worldwide, with significant diversity in China and Europe.
Lithobius
stone centipedes, common centipedes, brown centipedes, typical stone centipedes
Lithobius is a large genus of centipedes in the family Lithobiidae, containing over 500 species and numerous subspecies. Adults possess 18 body segments and 15 pairs of legs, with body lengths ranging from approximately 2–5 cm. The genus was erected by William Elford Leach in 1814 and serves as the type genus for the family Lithobiidae. These centipedes are primarily nocturnal predators found in soil, leaf litter, and beneath stones or bark across temperate regions worldwide.
Lithobius peregrinus
Peregrine Stone Centipede
Lithobius peregrinus is a stone centipede species first described in 1880 by Austrian myriapodologist Robert Latzel. The species has a cosmopolitan distribution with type locality in Dalmatia, Croatia. A 2022–2024 study of a troglophilic population in Gaura cu Muscă Cave, Romania documented significant morphological variations, asymmetries, and teratological features across 39 specimens. The species exhibits close water affinity and has been recorded in both surface and subterranean habitats.
Megacina
Megacina is a genus of armoured harvestmen in the family Phalangodidae, established by Ubick & Briggs in 2008. The genus contains at least four described species, all found in western North America. These species are characterized by heavily sclerotized body armor typical of the family. The genus was erected based on morphological distinctions from related phalangodid genera.
Meta dolloff
Dolloff Cave Spider
Meta dolloff, commonly known as the Dolloff Cave Spider, is a rare spider species endemic to California. It belongs to the family Tetragnathidae, a group characterized by elongated bodies and long jaws. The species is considered among the rarest spiders in North America, with limited observations documented.
Metellina
Autumn Spider
Metellina is a genus of orb-weaving spiders in the family Tetragnathidae, distributed primarily across Eurasia with two species in North America. The genus includes sixteen species as of 2025, with M. segmentata being the most abundant orb-weaving spider in Germany and introduced to Canada. Metellina spiders construct webs low in vegetation and are commonly known as Autumn Spiders. Some researchers have proposed placing this genus in a distinct family, Metidae.
Mezium
spider beetles
Mezium is a genus of spider beetles (subfamily Ptininae) with a global distribution concentrated in two main regions: the western Mediterranean from the Iberian Peninsula to Morocco and the Canary Islands, and central and southern Africa. Species in this genus are primarily cave-dwelling, feeding on animal feces and bat guano, though several have adapted to human-made structures such as barns and chicken coops. The genus includes notable species such as Mezium affine (shiny spider beetle) and Mezium americanum (American spider beetle).
Micropholcus
cellar spiders
Micropholcus is a genus of cellar spiders (Pholcidae) characterized by long legs and relatively fragile bodies. It is one of only two pholcid genera with both Old and New World distributions, though morphological and ecological differences exist between geographic groups. The genus currently contains at least 29 species, with major taxonomic revisions in 2014 (transfers from Leptopholcus) and 2024 (description of twelve new Old World species). The type species M. fauroti has a pantropical distribution and has been introduced globally.
Monopis crocicapitella
Pale-backed clothes moth, Bird-nest moth, Bird Nest Moth
A small tineid moth with a nearly cosmopolitan distribution, first described from the eastern United States. Adults have a wingspan of 10–16 mm. The species has been documented in cave environments and serves as a host for the parasitoid wasp Apanteles carpatus. Its capability to infest human dwellings remains unknown.
Neelidae
Neelidae is a family of minute springtails in the order Neelipleona, comprising at least four genera and more than 30 described species. Members average around 500 μm in length and possess globular bodies superficially resembling those of Symphypleona, with relatively long legs and a well-developed furca. Molecular phylogenetic data places Neelidae as a basal group to all other Collembola, despite earlier morphological hypotheses linking them closely to Symphypleona. The family has worldwide distribution with records from Gondwanan regions, caves, and various terrestrial habitats.
Neelipleona
Neelipleona is an order of minute hexapods within the class Collembola (springtails). Members are characterized by the complete absence of eyes, a trait distinguishing them from most other springtail groups. The group is primarily represented by the family Neelidae and includes genera such as Megalothorax and Spinaethorax. Many species exhibit troglomorphic adaptations, including elongated appendages and reduced pigmentation, associated with cave-dwelling habits. The taxonomic rank of Neelipleona remains debated, with some authorities treating it as a suborder of Symphypleona or placing it within Entomobryomorpha.
Neelus
Neelus is a genus of springtails (Collembola) in the family Neelidae, established by Folsom in 1896. The genus contains both surface-dwelling and troglobiotic (cave-restricted) species. Several species exhibit troglomorphic adaptations including elongated unguis (claws), larger body size, and elongated antennal sensilla. The genus has a broad geographic distribution spanning Europe, North America, and Asia, with notable cave-dwelling diversity in the Balkans.
Nemastomatidae
Nemastomatid Harvestmen
The Nemastomatidae are a family of harvestmen (Opiliones: Dyspnoi) comprising approximately 170 described species in 16 extant genera. The family is divided into two subfamilies with disjunct distributions: Nemastomatinae in Europe, North Africa, and western Asia, and Ortholasmatinae in western North America and eastern Asia. Members are characterized by variable body size (1–6 mm) and often possess elongated, thread-like pedipalps. The family is monophyletic and likely sister to the Dicranolasmatidae and Trogulidae.
Neobisiidae
neobisiid pseudoscorpions
Neobisiidae is a family of pseudoscorpions comprising approximately 550-750 species across 32-34 genera, distributed across Africa, the Americas, and Eurasia. Members are small arachnids ranging from 1 to 5 mm in body length, with body colors varying from reddish or dark brown through olive green to yellow or creamy white. The family includes both surface-dwelling and cave-dwelling species, with troglobitic species often exhibiting eye reduction or loss. Several genera, notably Stenohya, exhibit pronounced sexual dimorphism in pedipalp morphology.
Neobisioidea
Neobisioidea is a superfamily of pseudoscorpions established by Chamberlin in 1930. It contains seven families, including Gymnobisiidae, Hyidae, Neobisiidae, Parahyidae, and Syarinidae. Members of this superfamily exhibit diverse ecological specializations, with some species restricted to moist leaf litter habitats and others adapted to subterranean cave environments. Several families show Gondwanan distribution patterns and high levels of short-range endemism.
Neocarus
Neocarus is a genus of rare, relatively large mites in the family Opilioacaridae, characterized by primitive traits including six pairs of eyes and abdominal segmentation. Species in this genus are primarily associated with subterranean habitats in Brazil, including caves and ferruginous geosystems. Multiple new species have been described from Minas Gerais state, where they occur in both epigean and hypogean environments.
Neogoveidae
Neogoveid Harvestmen
Neogoveidae is a family of small, eyeless harvestmen (Cyphophthalmi) distributed across tropical regions of West Africa and the Neotropics. Members range from 1 to 4.5 mm in body length and exhibit distinctive morphological features including a heavily granulated dorsal scutum, laterally projecting ozophores, and variable adenostyle morphology. The family shows substantial undescribed diversity, with many species awaiting formal description.
Nesticus furtivus
Crystal Caverns Cave Spider
Nesticus furtivus is a small cave-dwelling spider endemic to a single cave system in Tennessee. It belongs to the family Nesticidae, a group commonly known as cave spiders or scaffold web spiders. The species has been documented only from Raccoon Mountain Caverns (formerly Crystal Caverns) near Chattanooga, making it one of the most geographically restricted spider species in North America. Its specific epithet 'furtivus' refers to its secretive, hidden lifestyle in subterranean habitats.
Oecleini
Oecleini is a tribe of planthoppers (Hemiptera: Cixiidae) established by Muir in 1922. Members exhibit distinctive wing venation patterns, including a trifid anterior MP branch in forewings and I-type hindwing venation with complete fusion of MP3+4 with CuA1. Some species possess subterranean adaptations, including a double-grasping coxo-femoral and femoro-tibial system in nymphs for clinging to roots. The tribe includes economically significant species investigated as potential vectors of palm phytoplasmas.
Oncopodura
Oncopodura is a genus of springtails (Collembola) in the family Oncopoduridae, established by Carl & Lebedinsky in 1905. The genus contains multiple species distributed across tropical and subtropical regions including Amazonia, the Andes, the Caribbean, and Southeast Asia. At least one species, O. moghanensis, has been described from subterranean cave habitats in Iran. Species in this genus exhibit distinctive morphological features of the postantennal organ (PAO) and furca that serve as key diagnostic characters.
Oncopoduridae
Oncopoduridae is a family of springtails (Collembola) in the order Entomobryomorpha, established by Carl and Lebedinsky in 1905. The family contains the genus Oncopodura, which includes subterranean species adapted to cave environments. Members of this family are characterized by distinctive morphological features of the postantennal organ (PAO) and furcula that are important for species identification.
Onychiurus
springtails
Onychiurus is a genus of springtails (Collembola) in the family Onychiuridae with cosmopolitan distribution. The genus contains numerous species, including both surface-dwelling forms in forest soils and agricultural fields, as well as specialized cave-dwelling species. Members of this genus have been extensively studied as indicators of soil health and pesticide impacts.
Opilioacarida
Opilioacarida is a small order of mites containing a single family, Opilioacaridae, with approximately 13 genera and 53 valid taxa. These mites are among the largest of their kind (1.5–2.5 mm) and are considered primitive due to retained ancestral traits including six pairs of eyes and abdominal segmentation. Molecular phylogenetics places them within Parasitiformes, though historically they were treated as distinct from both Acariformes and Parasitiformes. The group has a worldwide distribution with notable diversity in the Americas.