Spider taxonomy
Spider taxonomy encompasses the scientific classification, naming, and phylogenetic study of spiders, which constitute the order Araneae within the class Arachnida of the phylum Arthropoda.[1] As the largest and most diverse order of arachnids, Araneae includes approximately 53,547 validly described species organized into 4,492 genera and 139 families as of November 16, 2025.[2]
The taxonomic structure of Araneae is hierarchically organized, beginning with two primary suborders: Mesothelae, a relict group of primitively segmented spiders represented by a single family (Liphistiidae) comprising 88 species primarily found in Southeast Asia; and Opisthothelae, which encompasses the vast majority of spider diversity and is further divided into the infraorders Mygalomorphae (18 families, including tarantulas and trapdoor spiders) and Araneomorphae (120 families, including familiar groups like orb-weavers, wolf spiders, and jumping spiders).[3][4] This division is based on key morphological traits, such as cheliceral fang orientation and spinneret placement, which reflect evolutionary adaptations for predation and silk production.[4]
Modern spider taxonomy has been profoundly shaped by molecular phylogenetics, with comprehensive studies analyzing genetic data from hundreds of species across nearly all families to resolve long-standing debates on relationships among major clades.[4] For instance, analyses confirm the monophyly of groups like Entelegynae (a large subclade of Araneomorphae characterized by complex genital structures) and reveal non-monophyletic assemblages among cribellate orb-weavers, prompting revisions to family boundaries.[4] The World Spider Catalog, maintained by the Natural History Museum Bern, serves as the authoritative global reference, continuously updated with new discoveries—at a rate of about 900 species annually—and integrating nomenclatural changes since the order's establishment in 1758.[1] This dynamic framework not only aids in biodiversity assessment but also informs research on spider evolution, ecology, and biomedical applications, such as venom-derived therapeutics.[5]
History of Spider Taxonomy
Early Classifications
Early classifications of spiders were rooted in ancient and medieval natural history observations, which often grouped them loosely with insects due to superficial similarities in form and behavior. Aristotle, in his History of Animals (circa 350 BCE), described spiders' silk production and web-building as key traits, noting their egg-laying in silken webs and classifying them among insects (entoma) alongside scorpions and other small arthropods, emphasizing their aerial and predatory habits. This perspective persisted into medieval texts, where Isidore of Seville in his Etymologies (circa 636 CE) categorized spiders (aranea) as vermin generated spontaneously from the air, reinforcing their association with insects while highlighting their web-spinning as a means of aerial nourishment.[6] These early accounts focused on descriptive natural history rather than systematic taxonomy, viewing spiders through a lens of wonder and utility in folklore and medicine, without distinct ordinal status.
The advent of formal taxonomy in the 18th century marked a shift toward structured classification, though spiders remained embedded within insect groupings. In the 10th edition of Systema Naturae (1758), Carl Linnaeus placed spiders in the class Insecta under the order Aptera (wingless insects), establishing the order Araneae with the primary genus Aranea encompassing numerous species based on shared morphological features like eight legs and chelicerae, while distinguishing them from other apterous forms such as harvestmen (Phalangium).[7] Linnaeus's system relied on binomial nomenclature and observable traits like body segmentation and silk glands, but it did not separate spiders ordinally from insects, treating Araneae as one of seven orders within Aptera and listing around 70 spider species under Aranea. This framework provided a foundational catalog for European spiders, prioritizing regional descriptions over evolutionary relationships.
By the early 19th century, naturalists began advocating for spiders' separation from insects, introducing basic familial divisions grounded in cheliceral and fang morphology. Georges Cuvier, in Le Règne Animal (1817), elevated arachnids including spiders to a distinct class Arachnida within the phylum Articulata, differentiating them from Insecta by their lack of antennae, fused body segments, and four pairs of walking legs, while Pierre-André Latreille contributed detailed subsections on arachnids in the same work, emphasizing fang orientation as a key trait for subdivision.[8] Building on this, Tord Thorell in On European Spiders (1869) proposed the suborder Mesothelae for primitive forms like Liphistius with median (paraxial) cheliceral fangs, contrasting them with the transverse (diaxial) fangs of more derived Opisthothelae, thus laying groundwork for recognizing fang position as a fundamental classificatory criterion without phylogenetic intent.[9]
Regional monographs exemplified these descriptive approaches, focusing on morphological details for identification. John Blackwall's 1841 work in the Annals and Magazine of Natural History provided extensive descriptions of British spiders, cataloging species like Araneus diadematus based on eye arrangement, leg spination, and genital structures, proposing tribal divisions by eye number (e.g., six-eyed vs. eight-eyed) to aid regional surveys, though without broader systematic phylogeny.
During the late 19th and early 20th centuries, French arachnologist Eugène Simon advanced spider taxonomy through extensive descriptive work, naming over 1,000 new species between 1892 and 1903 in his multivolume Histoire Naturelle des Araignées. Simon's classifications emphasized morphological traits such as chelicerae structure and spinneret arrangement, which he used to delineate families including Salticidae (jumping spiders) and Theridiidae (cobweb spiders), establishing foundational groupings that integrated global collections and refined earlier regional efforts.[10][11]
In the early 20th century, Alexander Petrunkevitch contributed to systematic organization with his 1923 paper "On Families of Spiders" and subsequent catalogs, culminating in the incomplete Catalogue of American Spiders (1939), which began cataloging the suborder Mygalomorphae with 62 genera and 278 species while planning a comprehensive overview of American spider diversity based on anatomical features such as respiratory organs and genital structures.[12][13] Although planned as a multi-part work, only the first part was published. Petrunkevitch's work extended to global syntheses, incorporating internal anatomy to propose natural classifications that bridged regional faunas and highlighted evolutionary patterns in spider diversity.[14]
Mid-20th-century refinements built on these foundations, with B.J. Kaston's 1948 publication How to Know the Spiders providing detailed morphological keys that formalized suborders Mesothelae and Opisthothelae, distinguishing them by the position of book lungs and silk glands—ventral in Mesothelae and posterior in Opisthothelae—to clarify primitive versus derived forms.[15] Kaston's approach, echoed by contemporaries, prioritized external and internal morphology for identification, aiding in the delineation of families and genera amid growing collections.[16]
Catalogs by Carl Friedrich Roewer and Pierre Bonnet marked a peak in morphological compilation during the mid-20th century; Roewer's Katalog der Araneae (1942–1955) and Bonnet's Bibliographia Araneorum (1945–1959) together documented approximately 30,000 described species by 1955, organizing them into families and genera based on chelicerae, leg spination, and abdominal features.[1][10]
From the 1940s to the 1970s, taxonomists engaged in debates over the divisions between infraorders Mygalomorphae and Araneomorphae within Opisthothelae, questioning their monophyly and relationships through morphological comparisons of chelicerae orientation, tarsal claws, and respiratory systems, which led to refinements in superfamily boundaries and family assignments.[14]
Molecular Phylogenetics Era
The molecular phylogenetics era in spider taxonomy marked a paradigm shift from morphological classifications to DNA-based approaches starting in the late 20th century, enabling more robust inferences of evolutionary relationships across Araneae. Initial efforts focused on nuclear ribosomal markers like 18S rRNA and mitochondrial genes such as cytochrome c oxidase subunit I (COI), which provided the first molecular evidence for deep phylogenetic structure. For instance, Wheeler et al. (1993) analyzed 18S rRNA sequences in a combined morphological-molecular framework for arthropods, confirming Mesothelae as the basal lineage of spiders and supporting the monophyly of Opisthothelae.[17] These markers offered advantages over anatomy alone by revealing cryptic divergences and resolving longstanding ambiguities in higher-level groupings.
Advancements accelerated in the 2000s and 2010s with the advent of high-throughput sequencing, leading to phylogenomic studies that sampled thousands of loci. Bond et al. (2014) pioneered transcriptome analysis in spiders, sequencing over 1,000 genes from 41 species to reconstruct a backbone phylogeny; their results robustly supported Mygalomorphae monophyly while challenging prior assumptions about orb-web evolution. Building on this, Fernández et al. (2018) expanded to phylogenomics with approximately 2,500 nuclear protein-coding genes from 159 species, resolving key Araneomorphae clades such as the RTA (retrolateral tibial apophysis) group and contributing to taxonomic stability; by 2025, these insights underpin the recognition of 139 families in the World Spider Catalog.[18][19]
Recent phylogenomic innovations, particularly ultraconserved elements (UCEs), have further refined spider systematics as of 2025. Hedin et al. (2025) used UCE data from 200+ loci across North American taxa to affirm Cybaeidae monophyly, delineating five major lineages and describing new species in the Klamath Mountains region. Similarly, phylogenomic analyses of Idiopidae revealed intercontinental divergences within Idiopinae, prompting subfamily revisions and highlighting biogeographic contrasts between African and Australian clades.[20][21] These studies illustrate a broader trend where molecular evidence has driven 5-10% of family-level taxonomic changes since 2010, often by identifying hidden diversity.
The impacts of this era are profound, including the resolution of polyphyletic assemblages that confounded earlier classifications. A key 2025 example is the reevaluation of Pisauridae, where Yu et al. demonstrated non-monophyly through multi-locus phylogenetics, leading to the elevation of Dolomedidae as a distinct family and reassigning several genera. Such revisions have enhanced the estimated total of described spider species to 53,546 as of November 15, 2025, underscoring the ongoing discovery of biodiversity informed by genomic tools.[22]
Phylogenetic Overview
Evolutionary Origins
Spiders (order Araneae) represent a monophyletic lineage within the class Arachnida, characterized by basal traits such as paired book lungs for respiration and forward-projecting chelicerae adapted for piercing and liquefying prey.[23] Phylogenetic analyses place Araneae as part of the Tetrapulmonata clade, which also encompasses the orders Uropygi (whip scorpions), Amblypygi (whip spiders), and Schizomida (short-tailed whip scorpions), with this group forming a broader Arachnopulmonata clade sister to Scorpiones.[23] Cladograms derived from phylogenomic data illustrate Araneae branching from this Tetrapulmonata stem, highlighting shared pulmonary and appendage innovations that distinguish them from other arachnids like mites or harvestmen.[24]
The fossil record provides key insights into spider origins, with the earliest evidence of silk production appearing in the arachnid fossil Attercopus fimbriunguis from Middle Devonian deposits (~386 million years ago, Ma) in New York, USA.[25] Although Attercopus possessed spigots for silk extrusion on ventral plates, it lacked true spinnerets and other defining spider features, leading to its classification in the extinct order Uraraneida rather than as a crown-group spider.[25] True spiders, exhibiting spinnerets and book lungs, first appear in the Late Carboniferous (~310–305 Ma), as seen in fossils like Idmonarachne brasieri from France, marking the emergence of silk glands specialized for web-building and other functions around 300 Ma.[26]
Bayesian divergence dating, incorporating molecular clock analyses and fossil calibrations, estimates the crown-group origin of Araneae between 334 and 397 Ma, aligning with the Late Devonian to Early Carboniferous transition and reflecting terrestrialization events within Arachnida.[24] Basal spider ancestors resembled modern mygalomorphs, with two pairs of book lungs and downward-striking chelicerae; key divergences led to the Opisthothelae suborder, where araneomorph innovations—such as parallel chelicerae and advanced silk use—evolved, enabling diverse web architectures by the Late Triassic to Jurassic (~247–191 Ma) for orb-webs.[24] This transition underscores silk's role as a pivotal adaptation, facilitating predation and dispersal in terrestrial ecosystems.[25]
Key Relationships and Clades
The phylogenetic relationships among spiders (Araneae) are structured around three primary lineages: Mesothelae, which serves as the sister group to Opisthothelae, with the latter comprising Mygalomorphae as the basal clade to the more diverse Araneomorphae.[27][28] This topology is robustly supported by recent phylogenomic analyses integrating hundreds of loci from anchored hybrid enrichment and mitogenomic data, confirming the monophyly of these major clades.[29] Fossil evidence from the Carboniferous period indicates that both Mesothelae and Opisthothelae coexisted by approximately 300 million years ago, aligning with molecular divergence estimates.[30]
Key evolutionary transitions within this framework include the reduction and loss of the anterior median spinnerets in Opisthothelae relative to Mesothelae, a synapomorphy marking the divergence of the former suborder.[31] Within Araneomorphae, the anterior median spinnerets are further modified, often fusing to form the cribellum—a sieve-like structure for producing dry, adhesive silk—in the ancestral condition; this feature defines the Cribellata, while multiple independent losses gave rise to the Ecribellata, which rely on wet, viscid silk instead.[28] These transitions facilitated diverse web architectures and foraging strategies, with the Mygalomorphae-Araneomorphae split dated to approximately 270–310 million years ago based on fossil-calibrated phylogenomic models.[27][28]
Several clades exhibit strong monophyly in contemporary analyses. For instance, Atypidae (within Mygalomorphae) forms a cohesive group supported by shared morphological and molecular synapomorphies, such as specialized cheliceral structures.[32] Similarly, Theridiidae (an araneomorph family) is monophyletic, characterized by reduced abdominal sclerites and colulus remnants, as confirmed across mitogenomic and multi-locus datasets.[33] Recent phylogenomic revisions have unified the RTA-clade—defined by the retrolateral tibial apophysis on the male pedipalp—within the broader Entelegynae, resolving longstanding ambiguities in araneomorph interfamilial relationships through integrated morphological and genomic evidence.[34][35] Key araneomorph clades include the basal Haplogynae, characterized by simple genital structures, and the more derived Entelegynae, which encompasses the majority of araneomorph diversity including the RTA-clade and both cribellate and ecribellate lineages.
A simplified cladogram of spider phylogeny can be visualized as follows, with branch lengths roughly proportional to divergence times (in millions of years ago, Mya) from recent estimates:
Araneae
├── [Mesothelae](/page/Mesothelae) (sister; ~300 Mya split)
└── [Opisthothelae](/page/Opisthothelae)
├── Mygalomorphae (basal; ~280 Mya split from [Araneomorphae](/page/Araneomorphae))
│ └── e.g., Atypidae (monophyletic)
└── [Araneomorphae](/page/Araneomorphae)
├── Haplogynae (basal clade)
└── [Entelegynae](/page/Entelegynae) (derived; includes RTA-clade and cribellate lineages like Deinopoidea, with multiple [cribellum](/page/Cribellum) losses)
└── e.g., [Theridiidae](/page/Theridiidae) (monophyletic)
Araneae
├── [Mesothelae](/page/Mesothelae) (sister; ~300 Mya split)
└── [Opisthothelae](/page/Opisthothelae)
├── Mygalomorphae (basal; ~280 Mya split from [Araneomorphae](/page/Araneomorphae))
│ └── e.g., Atypidae (monophyletic)
└── [Araneomorphae](/page/Araneomorphae)
├── Haplogynae (basal clade)
└── [Entelegynae](/page/Entelegynae) (derived; includes RTA-clade and cribellate lineages like Deinopoidea, with multiple [cribellum](/page/Cribellum) losses)
└── e.g., [Theridiidae](/page/Theridiidae) (monophyletic)
This structure highlights the basal position of Mygalomorphae and the radiation of Araneomorphae, encompassing over 90% of spider diversity.[27][28]
Suborder Mesothelae
Morphological Traits
Mesothelae spiders exhibit a suite of primitive morphological traits that reflect their basal position in spider phylogeny, serving as the sister group to all other extant Araneae. These features include a segmented abdomen with visible tergites across multiple segments, orthognathous chelicerae, and respiratory structures retaining ancestral configurations.[36]
A defining characteristic is the arrangement of spinnerets, typically numbering seven or eight and positioned ventrally in the mid-abdomen on segments 3 and 4, rather than posteriorly as in more derived spiders. In the genus Liphistius, exactly eight spinnerets are consistently present, arranged in two transverse rows. The chelicerae are vertically oriented (orthognathous) and feature biserial dentition, with rows of teeth on both the promargin (anterior margin, often 2–5 teeth) and retromargin (posterior margin, often 2–3 teeth) of the cheliceral furrow, a plesiomorphic condition shared with some Paleozoic fossils. Additionally, these spiders possess a distinct transverse fovea on the carapace and lack fused opercula over the book lungs; instead, two pairs of book lungs are covered by separate, unfused sclerites integrated into the ventral opisthosomal sclerotization, allowing direct exposure to the environment.[36][30][25]
Silk production in Mesothelae involves a primitive glandular system akin to that of mygalomorph spiders, with numerous pyriform, tubuliform, and aciniform glands associated with the spinnerets, but distinguished by multi-segmented, appendage-like spinnerets bearing a high density of spigots for diverse silk types used in burrow lining and web construction. These spinnerets support the production of silk for structural purposes, reflecting an early evolutionary stage in arachnid silk utilization.[36]
Behaviorally, Mesothelae are obligate burrow-dwellers, constructing silk-lined tubular burrows capped with trapdoor-like lids composed of silk, soil, and debris, which they use for ambush predation; this sedentary lifestyle contributes to their limited dispersal capabilities. Their distribution is highly restricted to humid forest floors in Southeast Asia, including southern China, Japan, Vietnam, Thailand, and Myanmar, with the genus Liphistius exemplifying endemism to regions like the Malay Peninsula.[36]
Taxonomic Composition
The suborder Mesothelae comprises a single extant family, Liphistiidae, which has been recognized as the sole family within this basal spider lineage since its establishment by Tamerlan Thorell in 1869. This family encompasses all known living mesotheline spiders, characterized by their primitive abdominal segmentation and trapdoor burrowing habits. The type genus, Liphistius Schiødte, 1849, serves as the nomenclatural foundation for the family, with the name derived from Greek roots reflecting unique aspects of its morphology, though interpretations vary in the literature.
As of November 2025, Liphistiidae includes eight genera: Ganthela Xu & Kuntner, 2015; Heptathela Kishida, 1923; Liphistius Schiødte, 1849; Luthela Xu & Li, 2022; Qiongthela Xu & Kuntner, 2015; Ryuthela Haupt, 1983; Songthela Ono, 2000; and Vinathela Ono, 2000.[37] These genera collectively account for 196 described species, representing a modest diversity compared to other spider suborders, with all species endemic to humid forest habitats in Japan, China, and Southeast Asia, including regions like Thailand, Myanmar, Vietnam, and the Malay Peninsula.[2] Representative genera such as Liphistius, the most species-rich with over 70 valid species, are primarily distributed in Southeast Asia, while Ryuthela is restricted to Japan and Songthela to southern China.
Taxonomic revisions since the 2010s have included genus-level rearrangements, species descriptions, and a brief higher-level change with the elevation of Heptathelinae to family rank as Heptathelidae in 2022, which was synonymized in 2024 following molecular phylogenetic analyses supporting the monophyly of Liphistiidae.[38][39] Recent additions, such as two new Songthela species described in 2025 from Hunan Province, China, highlight ongoing discoveries driven by integrative morphological and molecular approaches, but these have not altered the familial structure.[40] Overall, the taxonomy remains stable, emphasizing the ancient, relict nature of this suborder with limited diversification in modern times.
Suborder Opisthothelae
Defining Features
The suborder Opisthothelae encompasses the vast majority of extant spider diversity and is distinguished from the more primitive Mesothelae primarily by derived morphological adaptations in the abdomen and respiratory system, reflecting a key divergence in arachnid evolution during the Mesozoic era.[28] These spiders exhibit spinnerets clustered at the posterior end of the opisthosoma, a result of the expansion of the third abdominal segment and reduction of posterior segments, which repositions silk-producing organs away from the more ventral, plesiomorphic arrangement seen in Mesothelae.[41] Additionally, Opisthothelae possess a longitudinal cardiac fovea on the carapace, a linear depression marking the position of the heart, contrasting with the transverse fovea of Mesothelae; in some taxa, the anterior book-lung opercula are fused into a single structure, streamlining ventral morphology. Genital structures are notably advanced, with males featuring complex palpal bulbs for sperm transfer and females displaying sclerotized epigynes or bursae, enabling intricate mating behaviors absent in the simpler gonopores of Mesothelae.[42]
Respiratory evolution within Opisthothelae shows a progression from the ancestral condition, with mygalomorphs retaining two pairs of book lungs—lamellate structures for gas exchange—while araneomorphs exhibit reduction to a single pair or complete loss, supplemented by a tracheal system that extends anteriorly into the prosoma for enhanced oxygenation in active lifestyles.[43] This shift correlates with diverse ecological niches, as book lungs provide efficient diffusion in burrowing or sedentary species, whereas tracheae support higher metabolic demands in agile hunters.[28]
Silk production in Opisthothelae has diversified beyond the basic spinneret output of ancestors, with some lineages evolving a cribellum—a sieve-like plate on the abdomen that produces dry, adhesive cribellate silk fibrils, enabling the construction of capture threads that entangle prey without viscous glue.[44] This innovation, plesiomorphic in certain araneomorph clades, facilitates varied web architectures and hunting strategies, such as orb webs or sheet traps, contributing to the suborder's adaptive radiation.[41]
Opisthothelae dominate global spider biodiversity, comprising approximately 99% of all described species, with approximately 53,400 species documented as of November 2025, distributed across nearly every terrestrial habitat from tropical rainforests to arid deserts.[2]
Infraorder Mygalomorphae
The infraorder Mygalomorphae represents the basal lineage within the suborder Opisthothelae, characterized by several plesiomorphic traits shared with more primitive spiders, such as the positioning of the anterior median spinnerets near the base of the abdomen.[45] This group encompasses 2,933 described species distributed across 16 families, with notable diversity in tropical and subtropical regions worldwide.[2]
Morphologically, Mygalomorphae are distinguished by their paraxial chelicerae, which feature parallel-hinged fangs that move vertically in a mastication-like motion, contrasting with the more derived diagonal orientation in other spiders.[46] Their respiratory system retains two pairs of book lungs located on abdominal segments VII and VIII, enabling gas exchange through stacked lamellae filled with hemolymph, while lacking any tracheal system for supplemental oxygenation.[47] Many species exhibit large body sizes, particularly in the family Theraphosidae (tarantulas), which includes over 1,100 species with body lengths often exceeding 5 cm and leg spans up to 30 cm.[48] Other families, such as Atypidae, are smaller and more secretive, with purse-web spiders constructing silken tubes on the soil surface.[29]
Ecologically, Mygalomorphae are predominantly ambush predators that rely on cryptic waiting strategies rather than active hunting or web-building for prey capture.[49] A significant portion are burrow-dwellers, excavating silk-lined tubes in soil or under bark for protection and ambushing passing insects or small vertebrates; for instance, members of the family Ctenizidae construct burrows capped with a camouflaged, hinged trapdoor made of soil and silk to launch sudden attacks.[50] Their venom, delivered via the paraxial chelicerae, is potent for subduing prey and can cause significant local pain, edema, and systemic effects in humans, though fatalities are rare due to available medical interventions and the low incidence of severe envenomations.[51]
Mygalomorphae constitute an ancient lineage, with the earliest fossils, such as Rosamygale grauvogeli, dating to the Middle Triassic around 245 million years ago, indicating their persistence since the Mesozoic era.[52] Recent mitogenomic analyses of over 250 spider species, incorporating complete mitochondrial genomes, have robustly confirmed the monophyly of Mygalomorphae with high support (bootstrap value = 99), resolving internal relationships into major clades like Atypoidea and Avicularioidea that diverged around 244 million years ago in the Middle Triassic.[27] This molecular evidence aligns with fossil-calibrated divergence estimates placing the infraorder's origin in the late Permian to early Triassic, underscoring their evolutionary stability.[52]
Infraorder Araneomorphae
The infraorder Araneomorphae constitutes the most species-rich lineage within spiders, comprising approximately 122 families and over 50,000 described species, representing more than 90% of all extant Araneae.[2] This group forms the derived sister infraorder to Mygalomorphae within the suborder Opisthothelae, having diverged during the Mesozoic era and undergone extensive radiation into diverse ecological niches.[53] Araneomorphs are distinguished by their orthogonal chelicerae, featuring transversely oriented fangs that articulate horizontally in a scissor-like pinching motion, enabling precise prey capture and venom injection.[53]
Key respiratory innovations include the reduction or transformation of the posterior book lungs into tracheal spiracles, which improve oxygen uptake and support higher metabolic rates associated with agile foraging behaviors.[53] In many male araneomorphs, particularly within the RTA clade, a retrolateral tibial apophysis on the pedipalp tibia serves as a stabilizing structure during copulation, reflecting adaptations in reproductive morphology.[53] These traits, combined with versatile silk production from spinnerets—including a primitive cribellum in basal lineages—have facilitated complex web architectures and hunting strategies.
Evolutionary dynamics within Araneomorphae feature notable bursts, such as the repeated loss of the cribellum in the Ecribellata clade, which decoupled silk production from substrate-bound constraints and promoted innovations like viscid capture threads in aerial webs.[44] This loss correlates with diversification into web-builders, such as orb-weavers in Araneidae, and active hunters, like jumping spiders in Salticidae.[53] Recent phylogenomic analyses in 2025 have refined resolutions of basal araneomorph groups, including the placement of Austrochilidae within early-diverging lineages like Austrochiloidea.[20] Globally, araneomorphs prevail across all terrestrial habitats, from humid forests to arid deserts, exerting significant predatory influence on arthropod communities.[54]
Classification of Araneomorphae
Haplogynae Clade
The Haplogynae represent a major clade within the Araneomorphae suborder of spiders, distinguished primarily by the haplogyne condition in female genitalia, which features a single, unpaired pair of spermathecae for sperm storage and lacks a hardened epigyne plate typical of more derived araneomorphs.[55] This simplified reproductive anatomy contrasts with the complex, paired structures in related groups and is associated with direct sperm transfer via the male embolus into the female's genital opening during mating.[56] Many haplogyne species are active hunters or ground-dwellers, often lacking elaborate orb webs and relying on ambulatory foraging in leaf litter, soil, or under bark, with a global distribution but highest diversity in tropical regions.[57]
Phylogenetically, the Haplogynae are positioned as basal araneomorphs, forming the sister group to the more diverse Entelegynae clade, a relationship supported by molecular and morphological analyses including genitalic simplicity and respiratory system traits. Recent phylogenomic studies, such as those reconstructing araneomorph relationships using transcriptome data, affirm the monophyly of the core Haplogynae, excluding certain peripheral lineages like Filistatidae, which nest outside as sisters to other basal groups. Araneomorph innovations, such as the retrolateral tibial apophysis (RTA) on the male pedipalp for improved grasping during courtship, are evident in many haplogyne taxa. Comprising approximately 18 families, the clade encompasses around 10% of all described spider species, with notable diversity in small to medium-sized forms adapted to humid, vegetated habitats.[22]
Key families include the Dysderidae, comprising about 660 species of robust, ground-active hunters with powerful chelicerae for piercing soft-bodied prey; and Oonopidae, the goblin spiders, with nearly 2,000 tiny, six-eyed species often found in tropical leaf litter.[58][59] These families exemplify the clade's ecological breadth, from web-building specialists to cursorial oonopids. Haplogyne diversity peaks in the tropics, where unique behavioral adaptations such as seismic communication—via substrate vibrations produced by stridulatory organs—facilitate mate location in certain species.[60]
Entelegynae Clade
The Entelegynae constitute the most species-rich clade within the Araneomorphae infraorder, accounting for approximately 90% of all described spider species and encompassing around 80 families.[34] This clade is defined by sophisticated female genital morphology, including paired spermathecae that store sperm and an overlying epigyne sclerite—a hardened plate that directs sperm flow during insemination, enabling a more controlled and efficient fertilization process compared to simpler systems. These structures reflect an evolutionary refinement in reproductive biology, supporting diverse mating strategies across the group. The Entelegynae also encompass the RTA-clade, named for the retrolateral tibial apophysis on the male pedipalp, which unites major lineages such as orb-weavers and jumping spiders, highlighting the clade's morphological and behavioral diversity.[61]
Prominent superfamilies within the Entelegynae include Araneoidea, which features families like Araneidae with over 3,000 species of orb-weaving spiders that construct geometrically precise capture webs, and Lycosoidea, home to wolf spiders (Lycosidae) that pursue prey through active ground hunting without webs.[62] Representative families illustrate the clade's ecological breadth: Pholcidae, known as daddy longlegs spiders, with over 2,000 species characterized by long, slender legs and fragile sheet webs in caves or under debris; Theridiidae, or cobweb spiders, produce irregular, sticky three-dimensional tangle webs to ensnare flying insects, while Salticidae, the jumping spiders, rely on acute vision and precise leaps for predation, forgoing webs entirely.[63][64] These groups underscore the Entelegynae's adaptability, with the clade's ~80 families spanning habitats from forests to urban environments worldwide.
In terms of evolution, the Entelegynae have driven much of spider diversification since their divergence, with molecular phylogenies indicating rapid radiations and recent taxonomic revisions, such as the 2025 delineation of species boundaries in the huntsman spider genus Heteropoda through multi-locus analyses.[65] Web architectures within the clade show multiple independent origins, with 2025 mitogenomic studies estimating 4 to 9 evolutionary events for various web types, from orbicular to sheet-like forms, facilitating niche exploitation and contributing to the group's dominance in arthropod predation.[27]
Families and Higher Taxa
Superfamilies Overview
The spider suborder Opisthothelae encompasses two infraorders, Mygalomorphae and Araneomorphae, each organized into distinct superfamilies that reflect evolutionary divergences based on key morphological traits. In Mygalomorphae, approximately five main superfamilies are recognized, including Avicularioidea (containing the diverse tarantula family Theraphosidae), Ctenizoidea (featuring trapdoor spiders like those in Ctenizidae), Atypoidea (purseweb spiders), Dipluroidea (funnel-web relatives), and Theraphosoidea (some primitive mygalomorphs). These groupings, updated in the World Spider Catalog version 26 as of 2025, highlight the relatively low diversity of mygalomorphs compared to their araneomorph counterparts, with around 3,000 species total across 17 families.[22][66]
Araneomorphae, comprising over 90% of all spider species, are divided into roughly 30 superfamilies, reflecting greater phylogenetic complexity. Notable examples include Dictynoidea, which groups cribellate sheet-weaving spiders in families like Dictynidae, and Thomisoidea, encompassing relatives of jumping spiders such as the crab spider family Thomisidae. Other significant superfamilies encompass orb-weavers (Araneoidea), wolf spiders (Lycosoidea), and huntsman spiders (Sparassoidea), with ongoing revisions incorporating molecular data to refine boundaries. The 2025 World Spider Catalog updates reflect new transfers and additions, such as reclassifications in Thomisoidea based on recent phylogenetic analyses.[22][67]
Classification of these superfamilies relies on homologies in genital structures, spinneret arrangements, and cheliceral fang orientations, with mygalomorph superfamilies distinguished by paraxial cheliceral movement and araneomorphs by more versatile, diagonal fang actions. Some groupings, like the revised Lycosoidea, exhibit non-monophyly due to paraphyletic elements resolved through molecular phylogenetics, prompting ongoing taxonomic adjustments. Transitional superfamilies, such as Gradunguloidea, bridge the infraorders by retaining primitive traits like multiple lung books while exhibiting early araneomorph genitalic features, underscoring evolutionary links between mygalomorph and araneomorph lineages.[22][68]
Family Diversity and List
Spider taxonomy encompasses 139 families and 53,547 described species as of November 2025, reflecting ongoing discoveries and taxonomic revisions.[2] Of these, the infraorder Mygalomorphae accounts for 17 families and 2,957 species, primarily characterized by primitive traits such as two pairs of book lungs and downward-pointing chelicerae.[2] In contrast, the infraorder Araneomorphae includes 121 families and 50,417 species, featuring more advanced features like forward-pointing chelicerae and diverse web-building behaviors.[2]
Family diversity varies widely in geographic distribution and species richness, with cosmopolitan families dominating global spider counts. The Salticidae, known for their keen vision and hunting prowess without webs, are nearly ubiquitous across all continents except Antarctica and boast over 7,000 species.[2] Similarly, the Linyphiidae, small sheetweb weavers often called money spiders, are most abundant in temperate regions and represent the second-largest family with 4,949 species.[69] Other notable patterns include high endemism in tropical families like Theraphosidae (tarantulas) and regional concentrations in Australasia for trapdoor spider families.
Recent taxonomic updates in 2025 have added three new Araneomorphae families: Ancylometidae (1 genus, 1 species; orb-weavers from South America), Lathyidae (specialized araneomorphs with unique spinneret morphology), and Melloinidae (mygalomorph-like araneomorphs).[3] These changes highlight ongoing refinements in superfamily assignments, such as within the Dictynoidea.
The tables below summarize all 139 families by infraorder, including valid species counts, brief key traits, and notes on recent changes where applicable. Data are drawn from the World Spider Catalog.[2]
Mygalomorphae (17 families, 2,957 species)
| Family | Species Count | Key Traits (Brief) | Recent Changes (2025) |
|---|
| Actinopodidae | 128 | Wishbone spiders; robust hunters | None |
| Atracidae | 39 | Mouse spiders; highly venomous | None |
| Barychelidae | 294 | Dwarf tarantulas; ground dwellers | None |
| Ctenizidae | 6 | Trapdoor spiders; burrowing | None |
| Cyrtaucheniidae | 109 | Primitive trapdoor spiders | None |
| Dipluridae | 154 | Funnelweb mygalomorphs | None |
| Euctenizidae | 79 | Trapdoor spiders; North American | None |
| Hexathelidae | 45 | Funnel-web spiders; primitive | None |
| Idiopidae | 452 | Trapdoor spiders; arid specialists | None |
| Macrothelidae | 56 | Burrowing mygalomorphs | None |
| Mecicobothriidae | 2 | Dwarf tube spiders | None |
| Microstigmatidae | 44 | Small mygalomorphs; South American | None |
| Migidae | 108 | Tree trapdoor spiders | None |
| Nemesiidae | 196 | Funnel trapdoor spiders | None |
| Paratropididae | 37 | Burrowing mygalomorphs | None |
| Porrhothelidae | 5 | Primitive mygalomorphs | None |
| Theraphosidae | 1,180 | Tarantulas; diverse habitats | None |
Araneomorphae (121 families, 50,417 species)
| Family | Species Count | Key Traits (Brief) | Recent Changes (2025) |
|---|
| Agelenidae | 1,464 | Funnel weavers; fast runners | None |
| Amaurobiidae | 285 | Hacklebanded hackled orbweavers | None |
| Ammoxenidae | 31 | Tarantula-like hunters | None |
| Anapidae | 266 | Cobweb weavers; tiny globose | None |
| Ancylometidae | 1 | Orb-weavers; South American | New family |
| Anyphaenidae | 483 | Lynx-like hunters | None |
| Araneidae | 3,159 | Orb-weavers; classic wheel webs | None |
| Archaeidae | 90 | Pelican spiders; ant mimics | None |
| Arkyidae | 25 | Ray spiders; leaf mimics | None |
| Austrochilidae | 8 | Long-legged hunters | None |
| Baryphymidae | 19 | Bark-dwelling crab spiders | None |
| Clubionidae | 618 | Sac spiders; foliage hunters | None |
| Corinnidae | 1,116 | Ground hunters; ant mimics | None |
| Ctenidae | 582 | Wandering spiders; large hunters | None |
| Cyatholipidae | 1 | Basal araneomorphs; tiny | None |
| Cybaeidae | 173 | Water spiders; sheet webs | New subfamilies |
| Desidae | 317 | Door-builder spiders | None |
| Dictynidae | 652 | Meshweb weavers; irregular webs | None |
| Dionycha (various) | Varies | One-claw families; diverse | None |
| ... (abbreviated for brevity; full list includes 105 additional families such as Eresidae, Gnaphosidae, Hersiliidae, Linyphiidae [4,949 spp.; sheetweb weavers], Lycosidae [2,509 spp.; wolf spiders], Oonopidae [1,981 spp.; goblin spiders], Pholcidae [2,055 spp.; cellar spiders], Salticidae [over 7,000 spp.; jumping spiders], Sparassidae [1,233 spp.; huntsman spiders], Theridiidae [2,605 spp.; comb-footed spiders], Thomisidae [2,195 spp.; crab spiders], and new 2025 families Lathyidae and Melloinidae) | - | - | - |