Cave-dwelling
Guides
Aglossa pinguinalis
Large Tabby, Grease Moth
Aglossa pinguinalis is a pyralid moth commonly known as the large tabby or grease moth. The species exhibits a rare feeding strategy among Lepidoptera: coprophagy, with larvae consuming animal feces. Larval development spans approximately two years, and the species shows strong habitat affinity for caves and sheltered environments. It is native to the Palearctic region but has been introduced to North America and New Zealand.
Amblypygi
whip spiders, tailless whip scorpions, amblypygids
Amblypygi is an ancient order of arachnids comprising approximately 280 species across five families. These nocturnal predators are characterized by extremely elongated first pair of legs modified as sensory "whips" and raptorial pedipalps for capturing prey. They lack venom glands, silk production, and the tail-like flagellum present in their relatives the whip scorpions (Uropygi). Several species exhibit unusual social behaviors including maternal care and kin recognition.
Anapistula
Anapistula is a genus of minute araneomorph spiders in the family Symphytognathidae, established by Gertsch in 1941. The genus comprises 28 described species distributed across tropical and subtropical regions of Asia, South America, and Europe. Members exhibit diverse habitat preferences: approximately 18 species occur in leaf litter, soil, and mosses, while seven are obligate cave-dwellers (troglobites) and one inhabits cave entrances. The genus includes some of the smallest spiders known, with A. ataecina from Portugal representing both the first European record of the family and one of the smallest described spider species.
Androniscus
Androniscus is a genus of small woodlice in the family Trichoniscidae, established by Verhoeff in 1908. The genus includes the well-known species Androniscus dentiger, commonly called the rosy or pink woodlouse. Members of this genus are characterized by their small size, distinctive coloration, and association with calcareous environments. Some populations show cave-adapted ecology, particularly in the southern part of their range.
Anthroleucosomatidae
Anthroleucosomatidae is a family of small to medium-sized millipedes in the order Chordeumatida, with approximately 19 genera and at least 40 described species. The family exhibits a core distribution centered on the Balkans and Caucasus region, extending eastward to Iran and central Siberia, with one highly disjunct North American species in Washington State. Many species are troglobionts (obligate cave-dwellers), including the world's deepest-occurring millipede, Heterocaucaseuma deprofundum, found below 2,000 meters in Krubera-Voronja Cave. The family shows exceptional diversity in the Caucasus, where 15 new genera and 36 new species were described in a single monograph.
Archoleptoneta
Archoleptoneta is a genus of small spiders in the family Archoleptonetidae, containing only two known species restricted to the United States. The genus was established by Gertsch in 1974 and is characterized by reduced eyes and troglobitic adaptations. These spiders are among the least studied North American arachnids, with very few observations recorded.
Arrhopalites
springtails
Arrhopalites is a genus of globular springtails (Collembola: Symphypleona) in the family Arrhopalitidae. The genus comprises approximately 40 valid species, with roughly half considered troglobionts—obligate cave-dwelling organisms. Species occur across diverse habitats including cave systems, forest leaf litter, and soil environments, with distribution spanning the Neotropical Region, Asia, Europe, and other regions. The genus is taxonomically significant for phylogenetic studies of Symphypleona, though many aspects of its biology remain understudied.
Arrhopalitidae
Arrhopalitidae is a family of springtails (Collembola) in the order Symphypleona, superfamily Katiannoidea. The family includes three genera: Arrhopalites, Pygmarrhopalites, and Troglopalites. Many species are obligate cave-dwellers (troglobionts) exhibiting pronounced troglomorphy, including reduced pigmentation, elongated appendages, and modified sensory structures. The family has a sister-group relationship with Katiannidae.
Artema
Giant Cellar Spiders
Artema is a genus of cellar spiders (family Pholcidae) first described by Charles Athanase Walckenaer in 1837. The genus includes some of the largest pholcid spiders and is primarily distributed from West Africa through Central Asia to Pakistan. One species, A. atlanta, has been introduced worldwide. The genus comprises twelve recognized species as of October 2025.
Bembidiini
Bembidiini is a large tribe of ground beetles (Carabidae) comprising over 120 genera and approximately 3,100 described species. Members are predominantly small to minute beetles, often found in riparian, intertidal, subterranean, and arboreal habitats. The tribe includes notable subtribes such as Xystosomina (arboreal tropical beetles) and Anillina (small litter-dwelling forms), as well as the subtribe Lovriciina containing highly specialized cave-dwelling species. Many species exhibit narrow habitat specificity, including intertidal gravel beaches and tropical forest canopy systems.
Blaniulidae
Thin Snake Millipedes
Blaniulidae is a family of julid millipedes characterized by extremely elongated, slender bodies with length-to-width ratios reaching 30:1. Members possess distinctive ozadenes (odiferous defensive glands) visible as spots on each segment, and may have eyes present or absent depending on species. The family exhibits diverse habitat preferences including leaf litter, soil, caves, and synanthropic environments. Several genera contain troglobitic species restricted to subterranean habitats, while others are widespread across Europe and have been introduced to other continents.
Blattellinae
wood cockroaches
Blattellinae is a subfamily of Ectobiidae containing approximately 70 genera. It includes the globally distributed German cockroach (Blattella germanica), a major household pest, as well as several endangered species such as those in Hololeptoblatta and Miriamrothschildia. The subfamily exhibits diverse ecological adaptations, with members occupying habitats ranging from wetlands and grasslands to caves and human dwellings.
Brachyceridae
Brachyceridae is a family of weevils (Coleoptera: Curculionoidea) containing at least 150 genera. The family has been treated historically as a subfamily of Curculionidae, but is now recognized as distinct. Some constituent genera, such as Absoloniella, contain blind, wingless species restricted to Mediterranean regions. The family belongs to the superfamily Curculionoidea, placing it among the true weevils.
Calicina mariposa
Calicina mariposa is a species of armoured harvestman in the family Phalangodidae. It was originally described as Sitalcina mariposa by Briggs in 1968 before being transferred to the genus Calicina. The species is known from North America. Very little published information exists regarding its biology or ecology.
Callipodida
Crested Millipedes
Callipodida is an order of millipedes comprising approximately 130 species across three extant suborders and seven families. Members are characterized by elongated bodies with 40–60 segments, reaching up to 100 mm in length, and often display distinctive dorsal crests or ridges. The order exhibits a disjunct distribution across the Northern Hemisphere, with populations in North America, Europe, western Asia, southern China, and Southeast Asia. Sexual maturity is achieved through teloanamorphosis, with males possessing a single pair of gonopods derived from the seventh leg pair.
Calymmaria emertoni
Calymmaria emertoni is a small true spider in the family Cybaeidae (formerly Hahniidae). It is one of approximately 31 Calymmaria species found north of Mexico, with most species restricted to the Pacific coast region. This species is notable for its distinctive basket-shaped web and nocturnal habits.
Camillidae
Camillidae is a small family of minute flies (2–3.5 mm) within the superfamily Ephydroidea. The family contains five genera, four extant and one fossil. Adults are slender, lustrous black flies with hyaline wings and distinctive bristle patterns on the head and thorax. Larval biology is poorly known, though they have been reared from bat guano and rock hyrax dung, suggesting a role in decomposing organic matter in sheltered environments.
Catops
Catops is a genus of small carrion beetles in the family Leiodidae, subfamily Cholevinae, established by Paykull in 1798. The genus comprises approximately 16 described species distributed across the Holarctic region. Several species are associated with cave habitats, including the Iranian endemic Catops farsicus.
Catoptrichus
Catoptrichus is a genus of small carrion beetles in the family Leiodidae, subfamily Cholevinae. Members of this genus are eyeless, subterranean-adapted beetles associated with cave and soil habitats. The genus was established by Andrew Murray in 1856 and is placed in the subtribe Catopina.
Causeyella
Causeyella is a genus of troglobitic (cave-dwelling) millipedes in the family Trichopetalidae, order Chordeumatida. The genus was established by Shear in 2003 and comprises three species: C. causeyae, C. dendropus, and C. youngsteadtorum. These millipedes are restricted to cave habitats in the southern Ozarks region of the United States.
Centromerus
dwarf spiders
Centromerus is a genus of dwarf spiders in the family Linyphiidae, containing approximately 88 species and two subspecies. The genus has a broad distribution spanning the Holarctic region, with species recorded across Europe, North America, Asia, and North Africa. Many species exhibit restricted geographic ranges, while a few such as C. sylvaticus have exceptionally wide distributions. The genus was established by Dahl in 1886.
Ceratolasmatidae
Ceratolasmatidae is a family of harvestmen (Opiliones) comprising eleven described species across four genera. The family's monophyly is questionable, with three potentially distinct lineages showing affinities to different harvestman families: Ceratolasma and Acuclavella related to Ischyropsalididae, Hesperonemastoma related to Sabaconidae, and Crosbycus possibly warranting separate family status. Body sizes range from less than 1 mm to 6 mm depending on genus.
Ceuthophilinae
cave crickets, camel crickets, spider crickets
Ceuthophilinae is a subfamily of Rhaphidophoridae comprising North American cave crickets. Members exhibit morphological adaptations to subterranean life including reduced eyes and depigmentation. They serve as hosts for parasitic larval mites in the group Parasitengona. The subfamily includes genera such as Ceuthophilus and Hadenoecus.
Ceuthophilus chiricahuae
Chiricahua cave cricket
Ceuthophilus chiricahuae is a species of camel cricket (family Rhaphidophoridae) endemic to North America. The species was described by Hubbell in 1936 and is named for the Chiricahua Mountains region. Like other members of the genus Ceuthophilus, it is wingless and adapted to cool, dark, moist habitats. Very few observations of this species have been recorded, with only six documented occurrences in biodiversity databases.
Ceuthophilus guttulosus guttulosus
camel cricket, cave cricket
Ceuthophilus guttulosus guttulosus is a subspecies of camel cricket in the family Rhaphidophoridae. Like other members of the genus Ceuthophilus, it is wingless and possesses exceptionally long antennae adapted for navigation in dark environments. The species exhibits the characteristic hump-backed appearance that gives camel crickets their common name. This subspecies has been recorded from several eastern and midwestern U.S. states.
Ceuthophilus guttulosus nigricans
Ceuthophilus guttulosus nigricans is a subspecies of camel cricket in the family Rhaphidophoridae. Like other members of the genus Ceuthophilus, it is wingless, nocturnal, and adapted to dark, humid environments. The subspecies designation indicates darker coloration compared to the nominate form. It has been recorded in the eastern United States from Indiana to North Carolina.
Ceuthophilus stygius
Kentucky cave cricket, cave camel cricket, Stygian Cave Cricket
Ceuthophilus stygius is a camel cricket in the family Rhaphidophoridae, commonly known as the Kentucky cave cricket or Stygian Cave Cricket. It is native to North America, with records from the southeastern United States and Indiana. As a member of the genus Ceuthophilus, it shares traits with other cave crickets including elongated antennae, powerful jumping legs, and adaptation to dark, damp environments. The species has been studied for its metabolic and water economy physiology in relation to body size and temperature.
Chilopoda
Centipedes
Chilopoda is a class of venomous, predatory arthropods comprising the centipedes. These elongated metameric animals possess one pair of legs per body segment, with leg counts ranging from 30 to 354 depending on species. Centipedes are distinguished from millipedes (class Diplopoda) by their single pair of legs per segment, flattened body profile, and venomous forcipules derived from modified first leg pairs. The class contains four extant orders: Scutigeromorpha (house centipedes), Lithobiomorpha (stone centipedes), Geophilomorpha (soil centipedes), and Scolopendromorpha (giant centipedes). All are obligate carnivores with predatory lifestyles.
Chionea obtusa
Chionea obtusa is a wingless crane fly in the family Limoniidae, one of sixteen North American species in the genus Chionea commonly known as "snow flies." Described by George Byers in 1983, this species inhabits the Nearctic region of western North America. Like other Chionea species, adults are active during winter months and are adapted to cold environments, though specific biological details for C. obtusa remain largely unstudied.
Chionea scita
snow crane fly, snow fly
Chionea scita is a wingless crane fly native to North America, notable for its winter activity and ability to walk on snow at temperatures as low as 14°F. Adults emerge between September and February, with peak activity in November and December. The species is commonly found in the northeastern United States and Canada, extending south to Georgia and Tennessee. Both sexes are yellow-brown, hairy, and spider-like in appearance, with body lengths of 5–8 mm.
Chionea stoneana
snow fly
Chionea stoneana is a wingless crane fly species in the family Limoniidae, commonly known as a 'snow fly.' Adults are active during winter months and are frequently observed crawling on snow surfaces. The species occurs in the central United States, ranging from Minnesota and Indiana south to Kansas, Oklahoma, and Missouri. Like other members of its genus, it possesses physiological adaptations for cold tolerance, including glycerol in its body fluids that acts as an antifreeze.
Chordeumatida
Sausage Millipedes, Spinning Millipedes
Chordeumatida is a large order of millipedes containing over 1,100 species, commonly known as sausage millipedes or spinning millipedes. Members of this order are distinguished by their teloanamorphic development—adding segments through molts until reaching a fixed adult number, after which molting ceases. They are notable for possessing spinnerets on their telsons that produce silk used to construct protective chambers for molting and egg-laying. The order exhibits considerable morphological diversity, with species ranging from 3.5 to 42 mm in length and displaying both cylindrical and flat-backed body forms.
Cicurina
cave meshweaver
Cicurina is a genus of small spiders commonly known as cave meshweavers, first described by Anton Menge in 1871. The genus has undergone multiple family reassignments, most recently placed in the family Cicurinidae in 2023. Species exhibit considerable size variation, with some measuring under 2 mm and others exceeding 13 mm. Many species are specialized cave dwellers (troglobites), including several federally endangered species in the United States.
Cirolanides texensis
Cirolanides texensis is a subterranean isopod species endemic to Texas. It belongs to the family Cirolanidae, a group of predominantly aquatic isopods that includes many cave-dwelling species. The species was described in 1896 by Benedict and is known from limited observations in karst groundwater systems.
Cladonychiidae
cladonychiid harvestmen
A small family of harvestmen (suborder Laniatores) comprising approximately 33 described species. Body length ranges from under two to about four millimeters. Members possess robust, spined pedipalps and relatively short legs, though the second pair may reach two centimeters. Coloration varies from reddish brown to dark brown in surface-dwelling species to pale yellow in cave-dwelling forms.
Coecobrya
Coecobrya is a genus of springtails (Collembola: Entomobryidae) characterized by pronounced troglomorphic adaptations in cave-dwelling species. The genus includes both surface and subterranean species, with Thai cave species exhibiting the most extreme troglomorphy known for Collembola in Southeast Asia. Many species display elongated antennae, legs and furca, reduced pigmentation, and eye loss. The genus has been recorded from caves in Thailand, South Africa, and other regions, with molecular studies supporting species discrimination using COI, 16S rDNA and 28S rDNA markers.
Coecobrya tenebricosa
Coecobrya tenebricosa is a cave-dwelling springtail species in the family Entomobryidae. It belongs to the tenebricosa-group, characterized by the absence of eyes, an unguis with one unpaired median tooth, and a manubrium lacking smooth chaetae. The species has been recorded from multiple disjunct regions including the Caribbean mainland, Europe, Hawaii, Macaronesia, and the Mediterranean.
Conotylidae
Conotylidae is a family of millipedes in the order Chordeumatida, containing approximately 19 genera and at least 60 described species. Adult members possess 30 body segments, counting the collum as the first and the telson as the last. The family exhibits its highest diversity in northwestern North America, where five of six subfamilies and ten of fourteen previously described genera occur. Several species are troglobiotic, restricted to cave environments, and some are considered climatic relicts from the Pleistocene.
Copepoda
copepods
Copepods are small aquatic crustaceans and one of the most abundant and diverse multicellular organisms on Earth. They occupy nearly every aquatic habitat, from marine plankton to deep ocean floors, freshwater lakes, groundwater systems, and even moist terrestrial environments such as leaf litter and bromeliad phytotelmata. The group includes free-living forms as well as highly modified parasites. Copepods are fundamental components of aquatic food webs, serving as critical prey for fish, whales, and other marine life, while also contributing to nutrient cycling and carbon sequestration through the biological pump.
Corydioidea
Sand and Cave Cockroaches
Corydioidea is a superfamily of cockroaches (order Blattodea) containing two extant families: Corydiidae (sand cockroaches) and Nocticolidae (cave cockroaches). Together these comprise approximately 50 genera and 250 species. The superfamily also includes two extinct families: Liberiblattinidae and Manipulatoridae from the Cretaceous. Molecular dating places the origin of crown Corydioidea at the Triassic–Jurassic boundary.
Crumomyia
Crumomyia is a genus of small flies in the family Sphaeroceridae, commonly known as lesser dung flies. Species in this genus have been documented from caves in Europe, including a troglophilous population in the Czech Republic that shows morphological adaptations to subterranean life. The genus is distributed across northern and central Europe, with records from Denmark, Norway, and Sweden.
Cyphophthalmi
Mite Harvestmen
Cyphophthalmi is a suborder of harvestmen (Opiliones) comprising six families grouped into three infraorders: Boreophthalmi, Scopulophthalmi, and Sternophthalmi. Adults range from 1 to 7 mm in length with relatively short legs compared to other harvestmen. The suborder is characterized by unique morphological features including elevated cone-shaped ozophores, a spermatopositor instead of a true penis in males, and a completely open gonopore lacking a genital operculum. Many species are eyeless and inhabit leaf litter or caves. Cyphophthalmi represents one of two major lineages of harvestmen, with the other lineage (Phalangida) containing Laniatores, Dyspnoi, and Eupnoi.
Derolathrus
Jacobson's beetles
Derolathrus is a genus of minute beetles in the family Jacobsoniidae, containing 12 described species. The genus includes both extant and extinct species, with fossil records from Cretaceous amber deposits in Myanmar and France. Several species exhibit troglomorphic adaptations, including reduced or absent eyes. The genus has a disjunct global distribution spanning tropical and subtropical regions.
Derolathrus cavernicolus
Jacobson's beetle
Derolathrus cavernicolus is a small beetle in the family Jacobsoniidae, first described from continental North America in 2010. The species is cavernicolous, inhabiting caves and forested habitats in Florida and the Caribbean. It represents the first record of the family Jacobsoniidae for North America. Some populations may have been introduced through horticultural trade.
Dichoxenus
Dichoxenus is a genus of broad-nosed weevils in the family Curculionidae. A 2024 taxonomic revision recognized 25 species, including 17 newly described from Mexico and the southern United States. The genus was expanded through synonymy of Anametis and transfer of several species from other genera. Most species are nocturnal and associated with vegetation, though some occur in leaf litter and three species are known or suspected to be cave-dwelling.
Diplura
Two-pronged Bristletails
Diplura is an order of small, wingless, eyeless hexapods within the class Entognatha, characterized by a distinctive pair of caudal appendages (cerci) that give them their common name "two-pronged bristletails." The group comprises approximately 800 described species worldwide, ranging from 2 to 50 mm in length, with some species reaching up to 8 cm. They inhabit moist soil, leaf litter, humus, and caves across all continents except Antarctica. Diplurans possess concealed mouthparts (entognathous), long bead-like antennae, and the ability to autotomize their cerci when threatened.
Dorypteryx
cave barklice
Dorypteryx is a small genus of cave barklice in the family Psyllipsocidae, containing at least four described species. The genus has a notably wide geographic distribution spanning five biogeographic realms: West Palaearctic, Afrotropical, Australian, Nearctic, and Neotropical, with a 2009 discovery extending its range to China and the Oriental realm for the first time. Species occupy cave and domestic habitats. The genus was taxonomically revised by Lienhard in 1977, who synonymized the monotypic genus Dolopteryx with Dorypteryx.
Dorypteryx domestica
cave barklouse
Dorypteryx domestica is a small psocid, commonly known as the cave barklouse, in the family Psyllipsocidae. Originally described as Dolopteryx domestica from Zimbabwe in 1958, it was later transferred to Dorypteryx by Lienhard in 1977. The species has a remarkably broad geographic distribution spanning Africa, Europe, Northern Asia, and North America, and has been recorded from domestic as well as cave habitats.
Eidmannella
Eidmannella is a genus of scaffold web spiders (family Nesticidae) established by Roewer in 1935. The genus contains eight described species, including the invasive Eidmannella pallida, which has demonstrated significant invasive potential and ecological concern. Species in this genus are associated with both surface and subterranean habitats.
Eidmannella pallida
Pallid cave spider
Eidmannella pallida is a scaffold-web spider in the family Nesticidae with native distribution in North America and established invasive populations across multiple global regions. The species has been introduced to Pacific Islands, the Galápagos Islands, Macaronesia, Spain, Japan, and Italy, where it demonstrates high invasiveness potential. It readily establishes permanent populations in anthropogenic habitats including vineyards, annual crop fields, and olive groves, and shows particular tendency to exploit vulnerable subterranean ecosystems such as caves.