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Spitting spider

Spitting spiders, belonging to the family Scytodidae, are araneomorph spiders renowned for their distinctive hunting strategy of projecting a high-velocity stream of sticky silk combined with venom from their chelicerae to ensnare and immobilize prey, such as insects and other small arthropods.^[1] These spiders typically measure 3–6 mm in body length, feature a pale yellow to light brown coloration often marked with dark patterns, and possess six eyes arranged in two triads rather than the eight eyes common in most spiders.^[2] Their cephalothorax is notably bulbous and elongated, housing enlarged venom glands that produce both the toxic and adhesive components of their projectile, while their long, slender legs—equipped with sensory setae—aid in detecting vibrations from potential prey despite their limited eyesight.^[1]^[3] The family encompasses approximately 250 species across four genera, with Scytodes being the largest and most widespread, comprising over 220 species.^[4]^[5] These spiders are predominantly distributed in tropical and subtropical regions worldwide, including the Neotropics, Nearctic, Palearctic, and Indo-Malayan areas, though many have become synanthropic—thriving in human-modified environments such as homes, caves, cellars, and under rocks or bark.^[1]^[3] Nocturnal by nature, they remain hidden in dark crevices during the day and actively hunt at night, relying on stealth and their specialized silk ejection rather than web-building.^[2] The spitting mechanism itself is remarkably efficient, propelling the silk at speeds up to 30 meters per second in a zigzag pattern that covers prey in under 1/700th of a second, allowing the spider to then approach, bite, and consume the immobilized victim after wrapping it in additional silk.^[1]^[2] Ecologically, spitting spiders play a beneficial role as predators of household pests, contributing to natural pest control without posing significant threats to humans, as their venom is mild and their fangs are too small to penetrate skin effectively.^[1] Reproduction involves females producing an egg sac containing 20–35 eggs, which they carry attached to their spinnerets for several weeks before the spiderlings undergo their first molt under maternal protection.^[1] Recent research has highlighted the potential biomedical applications of their venom peptides, which are under investigation for treating conditions like pain due to their unique molecular properties.^[2] Overall, the Scytodidae exemplify adaptive innovation in arachnid predation, with their global presence and urban adaptability underscoring their evolutionary success.^[4]

Taxonomy

Classification history

The family Scytodidae was first described by John Blackwall in 1864, based on specimens from Great Britain and Ireland, with Scytodes as the type genus.^[6] Early classifications placed several genera within Scytodidae, including Loxosceles, which was initially grouped there due to shared morphological traits such as modified chelicerae adapted for liquid prey capture.^[7] However, subsequent revisions in the 20th century separated Loxosceles into its own family, Loxoscelidae, in 1949, before it was briefly returned to Scytodidae in 1967; by the late 20th century, molecular and morphological evidence confirmed its placement in the distinct family Sicariidae, erected by Eugène Simon in 1893 for Sicarius and Loxosceles, reflecting phylogenetic divergence within the superfamily Scytodoidea. Taxonomic boundaries continued to evolve through the late 20th and early 21st centuries, driven by detailed morphological studies of genitalic structures and silk glands, which highlighted inconsistencies in family-level groupings.^[7] Molecular phylogenies, incorporating genes like 18S rRNA and 28S rRNA, further refined these classifications by revealing monophyletic clades within Scytodoidea and supporting the exclusion of genera like those now in Drymusidae. A significant shift occurred in 2012 with the revision of the genus Stedocys, which emphasized morphological autapomorphies such as unique tibial apophyses, prompting reevaluation of subfamily divisions.^[7] In 2022, a comprehensive taxonomic revision proposed the division of Scytodidae into two subfamilies based on integrated morphological (e.g., cheliceral gland morphology) and molecular evidence (e.g., COI and 16S rRNA sequences): Scytodinae Blackwall, 1864, encompassing Scytodes and Dictis, and the newly erected Scyloxinae Zamani, Stockmann, Magalhães, Baze, Wunderlich, Schembri & Marusik, 2022, including Scyloxes and Stedocys. This restructuring addressed prior paraphyly in Scytodinae and aligned the family more closely with phylogenetic hypotheses from total-evidence analyses. As of 2025, the World Spider Catalog recognizes Scytodidae as comprising 4 genera and 253 valid species, reflecting ongoing updates from these evidence-based revisions.^[8]

Current genera and species

The family Scytodidae currently includes four valid genera and a total of 253 valid species, as documented in the World Spider Catalog updated in October 2025.^[5] These genera are classified into two subfamilies: Scytodinae Blackwall, 1864, encompassing Scytodes Latreille, 1804 and Dictis L. Koch, 1872; and Scyloxinae Zamani, Stockmann, Magalhães, Baze, Wunderlich, Schembri & Marusik, 2022, containing Scyloxes Dunin, 1992 and Stedocys Ono, 1995. The genus Scytodes, the most species-rich in the family, comprises 226 valid species and has a cosmopolitan distribution, with the majority occurring in tropical and subtropical regions across all continents.^[9] Dictis contains 9 valid species, primarily distributed in Asia (from the Middle East to Southeast Asia and the Pacific) and Australia, though some, like Dictis striatipes L. Koch, 1872, have been introduced to the Americas.^[10] Scyloxes is represented by 3 valid species, all endemic to Asia, recorded from Central Asia (Tajikistan) to Southeast Asia (Malaysia and Thailand).^[11] Stedocys includes 15 valid species, confined to Asia, with concentrations in China, Japan, Malaysia, and Thailand.^[12] Among the most notable species is Scytodes thoracica Latreille, 1802, a cosmopolitan and synanthropic taxon commonly found in human habitations worldwide, from Europe and Africa to the Americas and Oceania. Another prominent example is Scytodes fusca C. L. Koch, 1845, native to tropical regions but forming an invasive species complex with records spanning the Neotropics, Asia, and introduced populations elsewhere. Taxonomic research remains active, contributing to the family's growing diversity.

Physical characteristics

Morphology

Spitting spiders, belonging to the family Scytodidae, exhibit a distinctive body plan characterized by an elongated, narrow cephalothorax and abdomen that gives them a hobo-like appearance, with the cephalothorax often described as oversized and sloping upward posteriorly, while the circular abdomen slopes downward and connects via a thin pedicel.^[1]^[13] This structure is adapted for their ambulatory lifestyle and unique predation strategy, emphasizing mobility over web construction. Like other members of the Haplogynae clade, spitting spiders possess six eyes arranged in two triads rather than the typical eight found in most araneomorph spiders, which reflects their reliance on other sensory modalities for navigation and prey detection.^[14]^[1] The chelicerae of spitting spiders are notably modified, featuring fused fangs that are immovable independently and serve as conduits for ejection; these are supported by enlarged poison glands housed in the cephalothorax, divided into an anterior compartment for venom production and a larger posterior compartment for a mucilaginous substance.^[1]^[15] These glands enable the spider's characteristic spitting mechanism, with fang-like projections facilitating the directed release of the glandular contents. Their legs are long and thin, typically glabrous except for scattered short sensory setae, and lack heavy spines, promoting a slow-walking gait suited to stalking; the front pair is often the longest, aiding in prey detection through trichobothria and chemoreceptors.^[1]^[13] Unlike web-building spiders, they possess reduced spinnerets that do not support cribellate silk production or elaborate web structures.^[16] Coloration in spitting spiders is generally pale yellow to brown, accented by darker speckled or banded patterns on the cephalothorax and legs that provide camouflage against bark, walls, and foliage in their typical habitats.^[1]^[17] Sexual dimorphism is minimal in Scytodidae, though males typically exhibit slightly more slender builds and enlarged pedipalps relative to females, which are adapted for reproductive functions.^[1]^[18]

Size and variation

Spitting spiders in the family Scytodidae exhibit a typical body length ranging from 3 to 10 mm, reflecting their generally small stature among araneomorph spiders.^[19] For instance, the cosmopolitan species Scytodes thoracica measures 3 to 6 mm in body length, with females typically larger at 4 to 5.5 mm compared to males at 3.5 to 4 mm, showcasing sexual dimorphism in size.^[1]^[20] Size variation occurs across genera within the family, with Scytodes species generally smaller, often under 5 mm, while those in Dictis tend to be larger, reaching up to 8 to 9 mm in females and 6 to 7 mm in males.^[21] This intraspecific and intergeneric range allows adaptation to diverse microhabitats, though overall body proportions remain compact with elongated legs contributing to a greater leg span. Coloration in Scytodidae displays polymorphism, ranging from translucent pale yellow or creamy-pink to mottled brown, often accented by black speckles, stripes, or bands on the cephalothorax and legs.^[1]^[20]^[22] These patterns, such as the lyre-like markings on S. thoracica or the bright orange hues in some tropical forms, provide camouflage against bark or foliage, with darker variants more prevalent in humid tropical environments.^[1]^[2] Intraspecific variation is evident in species like the invasive Scytodes fusca, where juveniles appear paler yellow with narrow dark crossbands, maturing to a uniform chestnut brown.^[23] This variation likely aids in blending with varied substrates during dispersal. Compared to related families, Scytodidae are smaller than many orb-weavers in Araneidae, which can exceed 25 mm and build expansive webs, but their size aligns closely with other ground-hunting spiders like those in Lycosidae, despite differences in leg proportions and body mass distribution.^[24]^[25]

Habitat and distribution

Preferred habitats

Spitting spiders (family Scytodidae) primarily inhabit warm and humid environments, including tropical forests, caves, and areas with leaf litter.^[26]^[27] Many species, particularly those studied in tropical regions like Singapore, are maintained under conditions reflecting these preferences, with temperatures around 24°C and relative humidity between 70% and 90%.^[28] They favor low-light microclimates, such as the aphotic zones of caves for troglobitic species like Scyloxes magna and Scyloxes sp.^[29], or twilight areas near entrances for others like Scytodes fusca.^[27] Several species exhibit synanthropic tendencies, readily adapting to human-modified habitats. For instance, Scytodes thoracica is commonly found in dark corners, basements, closets, and cellars of buildings, where it thrives in sheltered, humid indoor spaces.^[1] These spiders prefer vertical substrates and crevices for web construction, such as small funnel-like retreats leading into cracks, allowing them to position above potential prey while avoiding open areas. In both natural and urban settings, spitting spiders associate with microhabitats rich in insect prey, such as attics, kitchens, or forest leaf litter layers, where humidity remains elevated (typically 70–90%) and temperatures range from 20–30°C to support their activity.^[1]^[28] This positioning near prey hotspots enhances their role in controlling arthropod populations in these environments.^[1]

Geographic range and dispersal

The family Scytodidae, comprising spitting spiders, is predominantly distributed across tropical regions, with native ranges centered in the Paleotropics—including parts of Africa, Asia, and Australia—and the Neotropics.^[30] This tropical affinity reflects the family's evolutionary origins and ecological preferences, though some species have achieved broader distributions through historical dispersals driven by climatic factors such as the Indian monsoon, which facilitated overwater movements from Southeast Asia to East Africa between approximately 46.5 and 33.0 million years ago.^[4] Several species within Scytodidae exhibit cosmopolitan distributions, most notably Scytodes thoracica, which originates from the Mediterranean region but has been introduced to all continents except Antarctica via human-mediated transport.^[31] This species' synanthropic lifestyle, favoring human-modified environments, has enabled its establishment in temperate zones, including parts of Europe, North America, and Asia, often through trade routes involving ships and luggage.^[31] Similarly, the Scytodes fusca species complex, native to the Malay Archipelago, has undergone multiple long-distance invasions, spreading from Australasia to the Americas, Madagascar, Southeast Asia, South Asia, and Australia; a single widespread haplotype has been documented in these regions, likely propelled by human activities such as the international trade in plants.^[32] Dispersal in Scytodidae is primarily human-facilitated in modern times, with natural mechanisms playing a lesser role; ballooning is rare, while overwater rafting and wind-assisted movements have contributed to ancient expansions, such as those across the Indian Ocean.^[4] These spiders are notably absent from extreme cold environments, including high Arctic latitudes and polar regions, where low temperatures limit their survival and establishment.^[31] Urban adaptations, such as tolerance for indoor habitats, further enhance their invasive potential in non-native areas.^[32]

Behavior

Hunting technique

Spitting spiders (family Scytodidae) employ a distinctive hunting strategy that relies on rapid projectile ejection rather than web-building. They detect approaching prey primarily through vibrations transmitted via the substrate, prompting the spider to orient toward the disturbance and prepare for attack.^[33] Once prey is within range, the spider ejects a zig-zag pattern of silk glue from specialized ducts near the bases of its chelicerae, immobilizing the target at distances up to 20 mm. This sticky, venomous fluid congeals on contact, forming strands that tether small arthropods such as flies, moths, and silverfish to the substrate in as little as 0.2 to 0.7 seconds, effectively halting their movement without direct contact.^[33] Following immobilization, the spider cautiously approaches the entangled prey and administers a bite to inject additional venom, facilitating digestion and consumption; notably, Scytodidae do not construct webs for hunting or prey capture. This ambush-style predation underscores their adaptation as active, nocturnal foragers in varied environments.^[33] Spitting spiders in the family Scytodidae are predominantly solitary, with adults exhibiting strong territorial behaviors that limit inter-individual tolerance outside of brief mating periods. Adult females aggressively defend their webs against conspecific intruders, often through charging, chasing, or spitting, which underscores their territorial nature. Cannibalism is common among adults, particularly toward juveniles, as a means of resource competition in shared or overlapping spaces.^[34]^[28] Presocial traits emerge in several species, notably through maternal care that fosters temporary family groups. In species such as Scytodes intricata and Singaporean Scytodes sp., females guard egg sacs by carrying them with their chelicerae until hatching and provide brief provisioning by dragging prey to spiderlings, allowing juveniles to feed alongside or independently. These interactions create short-lived colonies consisting of a mother and her offspring, where spiderlings remain in the maternal web until the third or fourth instar, exhibiting cooperative prey capture on larger items through collective spitting and biting. Juvenile aggregations can form in high-density areas, such as communal web peripheries, though aggression and cannibalism increase as spiderlings mature, leading to dispersal. Male-female interactions show limited tolerance, primarily during courtship with web-tapping signals, and males may remain with females briefly post-mating before separation.^[35]^[28]^[36] More advanced sociality is observed in the Scytodes fusca species complex, where tropical populations display communal living in interconnected web complexes, including cooperative prey capture and web-sharing. These groups, often found in human-modified or natural shelters, tolerate multiple adults and juveniles, though cannibalism persists as a regulatory mechanism. Such behaviors represent a rare progression toward sociality among spiders, potentially contributing to the complex's invasive success through enhanced colonization via long-distance dispersal and group-level adaptations.^[30]^[37]^[38]

Reproduction and life cycle

Spitting spiders in the family Scytodidae exhibit solitary lifestyles, with interactions primarily limited to mating periods. Males detect female pheromones through chemosensory hairs on their pedipalps and legs, approaching potential mates cautiously due to the aggressive nature of both sexes.^[1] Copulation is brief, typically lasting minutes, during which males insert both palpal organs into the female's genitalia to transfer sperm; sexual cannibalism during or after mating is rare, allowing both sexes to survive for potential multiple matings.^[39] Following mating, females produce one to three egg sacs annually, each containing 20–35 large eggs enclosed in a simple silk structure.^[1] The female carries the egg sac in her chelicerae or attaches it to her body, providing protection through direct embryogenesis without intermediate larval stages, a characteristic of spider development.^[28] Incubation lasts approximately 2–3 weeks under typical conditions, after which spiderlings hatch and undergo an initial molt while still in or near the sac.^[1] Maternal guarding continues for 2–4 weeks post-hatching, with females sharing prey and tolerating offspring in their web until the spiderlings reach the second or later instars.^[40] Spiderlings disperse shortly after hatching or following their first molt, transitioning to independent foraging and hunting behaviors.^[1] Development proceeds through 5–7 instars, with juveniles molting periodically as they grow; maturity is reached in 2–3 years for females under natural conditions, though this can vary by species and environment.^[1] Most spitting spiders display iteroparity, producing multiple clutches over their lifetime, though some populations exhibit semelparity with reproduction limited to a single event.^[39] The overall lifespan of spitting spiders ranges from 1–2 years for males to 2–4 years for females, with death often resulting from predation, starvation, or exhaustion rather than post-reproductive decline.^[1] Development rates are influenced by environmental factors, including temperature, which accelerates growth in tropical habitats where many species occur year-round.^[40]

Venom and adaptations

Composition of venom and spit

The spit ejected by Scytodidae spiders is a distinctive venom-silk mixture produced in cheliceral glands, combining adhesive silk proteins with bioactive peptides for prey immobilization. Transcriptomic analyses reveal that approximately 50% of expressed sequences encode glycine-rich peptides, which form the primary silken matrix of the spit, providing elasticity and rapid congealing upon contact. These silk-like proteins exhibit high glycine-alanine content (19–40%), contributing to fiber shrinkage of 40–60% during ejection and overall structural integrity.^[15] Venom components constitute about 35% of the proteome, encompassing 13 distinct groups of cysteine-rich peptides (3–9 kDa) with inhibitor cystine knot (ICK) motifs, including neurotoxins classified as scytotoxins that target ion channels in prey. Enzymes such as astacin metalloproteases and longistatin are also present, facilitating tissue liquefaction and digestion post-immobilization, while the mixture's low mammalian toxicity stems from the absence of sphingomyelinase D and limited paralytic effects observed in bioassays. Specific examples include U₃-Sth1a and U₅-Sth1a peptides from Scytodes thoracica, which display structural stability but minimal activity against insect or mammalian receptors at doses up to 350 nmol/g.^[15]^[41] Adhesive properties arise from specialized motifs in the glycine-rich peptides, such as diglutamine (QQ) and dityrosine (YY) sequences, which promote β-sheet formation via hydrogen bonding for fast-drying stickiness. The glands feature dual compartmentalization—anterior reservoirs for venom peptides and posterior for glue-like silk—enabling precise mixing distinct from the abdominal silk glands and cephalothoracic venom glands typical in other spiders. Recent venomics research underscores the diverse peptide repertoire (over 19 toxin families) and biochemical divergence of these components from ancestral silk-producing origins, with conserved transcription networks adapted for toxicity.^[15]^[42]

Evolutionary origins and functions

Spitting spiders (family Scytodidae) belong to the superfamily Scytodoidea within the Haplogynae clade, an early-diverging lineage of araneomorph spiders that represents basal araneomorphs in the spider phylogenetic tree.^[43] This position places Scytodidae among the more primitive araneomorph groups, with the divergence of Haplogynae from other araneomorph lineages estimated around 200 million years ago during the Late Triassic to Early Jurassic.^[44] The family's origins trace back to this haplogyne lineage, where genomic and transcriptomic analyses indicate that spider venom glands, including those modified for spitting in Scytodidae, likely evolved from ancestral silk-producing glands present in early chelicerates.^[42] In Scytodidae, these glands produce a silk-like adhesive combined with venom, enabling the unique ejection mechanism observed today.^[42] The evolution of spitting behavior facilitated adaptive radiation in Scytodidae by serving dual roles in prey capture and predator defense, thereby reducing dependence on traditional web-building strategies common in other spider families. This innovation allowed spitting spiders to exploit diverse microhabitats, such as leaf litter and bark crevices, where active hunting without webs provides a selective advantage over sedentary orb-weavers.^[33] By projecting a rapid, sticky silken projectile from their chelicerae, Scytodidae can immobilize insects at distances up to several body lengths, enhancing foraging efficiency in cluttered environments.^[33] Beyond predation, spitting functions as a primary defense mechanism against threats, with spiders ejecting the adhesive substance to deter predators like larger arthropods or vertebrates, often entangling approaching attackers.^[33] This defensive capability has contributed to the invasive success of certain Scytodes species, such as the Scytodes fusca complex, which has achieved global expansion through multiple long-distance dispersals, facilitated by human-mediated transport and the versatility of their hunting and defense strategy.^[30] Recent comparative venomics studies across spider lineages, including basal groups like Haplogynae, reveal evolutionary shifts in venom composition from primarily predatory to more defensive functions in some taxa, underscoring the adaptive plasticity of these glandular systems.^[45]

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