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Stridulation

Stridulation is the production of sound through the friction generated by rubbing two hardened body parts together, a mechanism that generates vibratory and acoustic signals primarily for communication in various animal species.^[1] This process, known as stridulation, evolved independently multiple times across arthropods, enabling functions such as mate attraction, territorial defense, and predator deterrence.^[2] The resulting sounds vary from audible chirps and rasps to ultrasonic frequencies, depending on the species and environmental context.^[3] In insects, particularly within the order Orthoptera (including crickets, grasshoppers, and katydids), stridulation is achieved by scraping a file-like structure on one body part against a ridge or scraper on another, such as wings or legs, to produce species-specific songs that play crucial roles in sexual selection and species recognition.^[4] For example, male crickets rub their forewings together, with the file on one wing interacting with the scraper on the other to create rhythmic chirps that attract females and repel rivals.^[5] Similar mechanisms occur in other insects like ants, where workers rub a plectrum on the abdomen against a file to produce alarm signals during nest disturbances.^[6] Beyond insects, stridulation is widespread in non-insect arthropods, including crustaceans such as fiddler crabs, which rub specialized appendages to generate sounds during reproductive and territorial displays, and spiders or scorpions that use chelicerae or pedipalps for substrate-borne vibrations.^[7]^[1] Although less common outside arthropods, stridulation appears in certain reptiles and mammals, highlighting its convergent evolution.^[3] In snakes, species like the saw-scaled viper (Echis carinatus) rub serrated lateral scales against each other to produce a harsh, rasping warning sound when threatened.^[8] The lowland streaked tenrec (Hemicentetes semispinosus), a mammal endemic to Madagascar, uniquely employs stridulation among mammals by rubbing specialized quills on its back together to emit high-pitched ultrasonic clicks for group communication and defense.^[9] These examples underscore stridulation's versatility as an acoustic strategy, often integrated with other sensory modalities to enhance survival and reproductive success across diverse taxa.^[2]

Definition and Mechanisms

Definition

Stridulation is the act of producing sound or vibration by rubbing together certain body parts, typically involving friction between specialized hard or sclerotized structures such as a file-like ridge (pars stridens) and a scraper (plectrum).^[10] This mechanism generates acoustic signals through repeated frictional contact, often amplified by resonant body parts.^[11] The term "stridulation" originates from the Latin verb strīdō, meaning "to make a harsh, creaking, or shrill sound," with the noun form entering English in the 1830s to describe such noisy friction.^[12] It specifically denotes tribulatory sound production, distinguishing it from other bioacoustic methods like tymbalation, which relies on the buckling and snapping of a taut membrane, or percussion, which involves striking body parts or substrates to create impact sounds.^[13] Stridulation was historically recognized and first systematically described in insects during the 19th century by entomologists, with Jean-Henri Fabre providing early detailed observations of the rubbing of dry membranes in grasshoppers to produce their characteristic rustling sounds.^[14] This behavior is primarily associated with arthropods as the dominant group exhibiting stridulation.^[3]

Mechanisms of Sound Production

Stridulation involves the mechanical interaction of specialized body parts, typically a ridged surface known as the file and a scraping structure called the plectrum, which rub against each other to generate sound through friction.^[3] The file consists of a series of ridges or teeth, while the plectrum is a hardened edge or projection that engages these ridges sequentially.^[15] This rubbing action creates intermittent contact points, producing a series of vibrations as the plectrum passes over each ridge.^[16] The primary mechanism of sound production relies on these friction-induced vibrations, where each ridge strike generates a brief pulse of mechanical energy.^[17] These pulses propagate as either airborne sound waves or substrate-borne vibrations, depending on the organism's anatomy and environment, with the vibrations coupling to the surrounding medium for transmission.^[15] The resulting sound is often characterized by a pulsed structure, forming chirps or trills composed of discrete, repeatable impulses.^[16] Key factors influencing the acoustic properties include the spacing of ridges on the file and the speed of the plectrum's movement, which together determine the fundamental frequency of the sound—typically ranging from a few kilohertz to over 100 kHz in various systems.^[3] The amplitude, or intensity, is primarily governed by the force applied during the rubbing action, with greater pressure yielding louder pulses, often reaching up to 100 dB at close range.^[17] These parameters allow for modulation of the signal's pitch and volume through controlled variations in motion.^[15] Biomechanically, stridulation is powered by specialized muscles that drive the rhythmic, oscillatory motion of the plectrum across the file, enabling precise timing and repetition rates.^[16] Body structures, such as exoskeletal plates or membranes, often function as resonators, amplifying the vibrations and enhancing efficiency by matching the pulse frequency to natural resonant modes.^[15] Variations in the system include linear files with evenly spaced, continuous ridges for sustained rubbing versus peg-based configurations using discrete projections, which can produce sharper, more irregular pulses.^[18]

Stridulation in Arthropods

In Insects

Stridulation is a widespread sound-producing mechanism among insects, particularly prevalent in the orders Orthoptera, Hemiptera, and Coleoptera, where it facilitates communication through diverse anatomical adaptations.^[3] In Orthoptera, encompassing crickets and grasshoppers, stridulation serves as the primary method for generating acoustic signals, often involving friction between specialized body parts to produce species-specific calls.^[19] Hemiptera, such as certain cicadas, predominantly rely on tymbal organs for sound production, though some species incorporate stridulatory mechanisms, especially in females, to generate supplementary signals during interactions.^[20] Similarly, in Coleoptera, beetles employ stridulation for distress or courtship signals, adapting the rubbing action to their hardened exoskeletons.^[21] In crickets (family Gryllidae within Orthoptera), males typically produce chirps via tegminal stridulation, where a file—a series of ridges on the underside of one forewing—is rubbed against a hardened scraper vein on the opposing forewing during rapid wing movements.^[22] This mechanism generates pulsed sounds that convey information for species recognition, allowing females to distinguish conspecific males amid choruses of sympatric species through unique temporal patterns in the chirps.^[23] The process amplifies the low-frequency wing strokes (around 20-30 Hz) into audible carrier frequencies of 3-6 kHz via mechanical resonance of the wings.^[24] Grasshoppers (suborder Caelifera, also Orthoptera) utilize a distinct stridulatory apparatus, primarily involving the hind legs: a row of pegs or teeth on the inner surface of the hind femur is rubbed against a thickened vein on the forewing tegmen to create rasping or buzzing sounds.^[3] This leg-tegmen friction produces broadband noise rather than tonal chirps, with variations in peg density and leg movement speed tailoring the acoustic output for mate attraction or territorial signaling in different species.^[4] Among beetles (Coleoptera), the deathwatch beetle (Xestobium rufovillosum, family Anobiidae) exemplifies stridulation adapted for substrate-borne communication, where the insect rubs a file on the prosternum against a plectrum on the head or thorax, resulting in rhythmic tapping sounds that propagate through wood.^[25] These ticks, produced in series of 4-8 pulses, function in courtship and may deter predators by signaling distress within confined larval galleries.^[26] Sexual dimorphism in stridulatory structures is common across these groups, with males often possessing more pronounced files, scrapers, or pegs to enable louder or more frequent calling, while females typically exhibit reduced or absent organs but respond to male signals using their antennae for close-range detection.^[27] In crickets, for instance, courting females position their antennae near the male's vibrating wings and abdomen to assess song quality before mounting, highlighting the antennae’s role in tactile and vibratory cue integration.^[27] The acoustic properties of insect stridulation vary by taxon but emphasize temporal patterns for effective communication; in crickets, calling songs feature pulse rates of 20-40 per second within chirps, enabling females to recognize and orient toward appropriate males based on interpulse intervals and syllable durations.^[28] These rates, modulated by temperature and species, underscore stridulation's efficiency in producing discriminable signals despite the insects' small size and limited muscle power.^[29]

In Other Arthropods

In arachnids, stridulation is prevalent among certain spiders, where specialized structures facilitate sound production primarily for courtship or defense. For instance, wolf spiders of the genus Schizocosa generate seismic signals through palpal stridulation, involving flexion of the pedipalp's tibio-cymbial joint, with a hardened scraper on the cymbium rubbing against a file on the tarsus base.^[30] These vibrations, peaking at frequencies around 1000 Hz, are produced during courtship displays and may convey information about male quality.^[30] Similarly, trapdoor spiders in the family Idiopidae exhibit stridulation using specialized textured structures on the palpal tibia, creating vibratory signals often associated with territorial behavior.^[31] In some species, chelicerae or pedipalps are rubbed against the exoskeleton to produce courtship vibrations, leveraging the sclerotized exoskeleton common to arthropods. Scorpions in families such as Buthidae produce stridulatory sounds by rubbing the chelicerae or pectines against the tergites or other exoskeletal parts, generating substrate-borne vibrations for defense or courtship communication.^[1] Among crustaceans, stridulation occurs in semi-terrestrial and aquatic species, often producing substrate-borne or water-propagated sounds adapted to their environments. Fiddler crabs (Uca spp.) employ claw stridulation during agonistic interactions and courtship, where males rub specialized tubercles on the major chela against the body to generate warning rasps.^[32] Ghost crabs (Ocypode spp.), such as O. quadrata, use a similar mechanism on the major claw, with a pars stridens of tubercles on the dactylus scraping against a plectrum on the ischium, yielding pulses at 30-40 Hz for territorial defense.^[32] Snapping shrimp (Alpheidae), while primarily known for snaps produced by claw closure inducing cavitation bubbles that collapse to create broadband pulses, also possess stridulatory devices in over 20 crustacean families, including rubbing of chelae to produce rasping sounds supplementary to snaps. These mechanisms highlight adaptations for communication in watery media, where airborne sound transmission is limited. Stridulation in myriapods is rare but documented in certain centipedes (Chilopoda), serving mainly defensive roles through substrate vibrations. Some species, including those in Scutigeromorpha and Pleurostigmophora, produce sounds by rubbing leg segments together, generating low-amplitude rasps that deter predators via vibratory cues.^[33] This contrasts with more aerial forms in insects, as myriapod stridulation emphasizes ground-transmitted signals suited to their terrestrial, often nocturnal habits. Unlike the predominantly airborne stridulation in many insects, that in other arthropods frequently relies on substrate-borne vibrations, particularly in semi-aquatic crustaceans where water conduction enhances signal propagation over distance.

Stridulation in Vertebrates

In Reptiles and Fish

Stridulation in reptiles is primarily observed in certain venomous snakes, where specialized scales are rubbed together to generate defensive buzzing or rasping sounds. Similarly, some vipers, including the saw-scaled viper (Echis carinatus), employ a comparable mechanism by coiling and rubbing keeled scales on their flanks and sides, creating a distinctive sawing or sizzling sound as a threat signal.^[34] These acoustic displays serve ecological roles in deterrence, often accompanying postural changes like body inflation or strikes. In fish, stridulation occurs through friction of bony structures, particularly in catfishes and syngnathids, adapting to aquatic environments for communication or alarm. Marine catfishes of the family Ariidae, such as Arius maculatus, generate grunting sounds by vibrating their pectoral fin spines against grooves in the shoulder girdle during handling or distress.^[35] Seahorses (Hippocampus spp.) produce pop-like clicks via stridulatory organs in the head and tail; in the tiger-tail seahorse (H. comes), head stridulation involves the supraoccipital bone rubbing against the coronet during rapid head movements, while tail mechanisms similarly rely on bony articulations for distress signals.^[36] Pimelodid catfishes, like Iheringichthys labrosus, use pectoral girdle friction where ridges on the spine's dorsal process scrape against a groove, yielding pulsed stridulatory sounds often paired with swimbladder drumming in defensive contexts.^[37] Acoustic characteristics of stridulation differ markedly between reptiles and fish due to medium-specific adaptations. Snake stridulations, such as hisses or rasps, typically feature higher frequencies spanning 3–13 kHz, suited for airborne propagation in terrestrial habitats.^[38] In contrast, fish stridulations exhibit lower dominant frequencies, often 500–1100 Hz in catfishes (e.g., ~656 Hz in pimelodids and ~1130 Hz in ariids), facilitating transmission through water where low frequencies attenuate less rapidly.^[37]^[35] Seahorse pops are brief broadband pulses with energy concentrated below 2000 Hz, emphasizing short-duration distress calls.^[36] This form of sound production in reptiles and fish represents evolutionary convergence with arthropod stridulation, arising independently through analogous friction-based mechanisms but utilizing scales or fins rather than appendages like legs or wings.^[2] In vertebrates, these adaptations likely evolved separately in ectothermic lineages to exploit body structures for threat signaling, contrasting with the exoskeletal files and pegs common in arthropods.

In Mammals

Stridulation is exceptionally rare among mammals and is best documented in the lowland streaked tenrec (Hemicentetes semispinosus), a small insectivorous mammal endemic to the humid forests of eastern Madagascar. This species produces sound by rubbing together specialized quills located in a stridulating organ on the mid-dorsal region of its back, consisting of typically 14 to 18 enlarged, hollow, and ridge-bearing spines that lack barbs and are less detachable than surrounding quills.^[39] The quills are vibrated through contractions of thickened cutaneous muscles symmetrical about the sagittal plane, with fast-twitch myosin fibers enabling rapid movement; the organ measures approximately 16.8 mm long and 8.55 mm wide, and body movements such as partial quill erection facilitate the friction.^[40] These rasping sounds are high-pitched and ultrasonic, spanning a broad frequency band from 2 kHz to 200 kHz, with peak energy concentrations between 20-30 kHz and pulsed components around 12-15 kHz, rendering them inaudible to humans but detectable by conspecifics over distances exceeding 4 meters.^[39] This form of stridulation serves primarily for intraspecific communication within multi-generational family groups, coordinating movements between mothers and offspring during foraging in dense leaf litter habitats, where visibility is low.^[41] The sounds also function in antipredator contexts, acting as alarm signals that promote group arousal and cohesion when predators approach, often accompanying defensive postures like crest erection, elevated stance, and hissing; playback experiments have shown these calls can even attract certain predators initially, potentially as part of a deimatic display.^[39] Ecologically, this adaptation suits the tenrec's nocturnal, fossorial lifestyle in Madagascar's understory, where family units of up to 25 individuals huddle and forage together, using the stridulation to maintain contact and deter threats in cluttered environments.^[41] The lowland streaked tenrec represents a remarkable case of convergent evolution with arthropods, particularly insects, as it employs true stridulation—friction between rigid body structures—despite lacking an exoskeleton, making it the only confirmed mammal to do so.^[39] No other mammalian species exhibits verified stridulation, although some rodents, such as kangaroo rats, generate friction-based sounds through foot-drumming or tooth-grinding that superficially resemble it but do not involve dedicated stridulatory organs.^[39] This uniqueness underscores the tenrec's isolated evolutionary history on Madagascar, where such traits have arisen independently from those in other vertebrates.^[41]

Biological Functions

In Communication and Mating

Stridulation serves as a primary mechanism for acoustic signaling in mating and social interactions among arthropods, enabling precise species recognition and coordination between individuals. In crickets, males generate species-specific chirp patterns via forewing stridulation to attract females from afar, with song characteristics conveying information about male quality and fitness. Females typically exhibit strong preferences for longer chirps and higher chirp rates, traits that correlate with larger body size and better nutritional status, respectively, thereby selecting for healthier mates.^[42] These preferences drive female phonotaxis, where receptive females orient toward and approach the most attractive signals in natural settings.^[43] Courtship in katydids often features interactive stridulation sequences that synchronize male and female behaviors, facilitating mutual assessment and reducing mating errors. Males initiate with calling songs produced by rubbing specialized wing structures, prompting receptive females to respond with short, high-frequency ticks or clicks via their own stridulatory organs.^[44] This duetting alternates rapidly, with female responses occurring within milliseconds of the male signal, allowing pairs to confirm compatibility and escalate to physical contact. In species like the tropical katydid Neoconocephalus spiza, such alternating patterns emerge from male-male competition but ultimately benefit female choice by highlighting leading signals that females preferentially follow.^[45] Grasshoppers employ stridulation for territorial communication, where males produce harsh, rasping sounds to assert dominance and repel intruders, thereby securing mating resources. These aggressive signals, generated by hind-leg friction against wings, function in acoustic contests that alternate between rivals, minimizing energy costs of physical clashes while advertising resource-holding potential.^[46] In band-winged grasshoppers, such rasps integrate with visual displays to maintain territory boundaries during peak reproductive periods.^[47] Across taxa, stridulation supports broader social cohesion and mate recognition. In certain fish, like the pictus catfish, stridulatory sounds from pectoral fin movements contribute to choruses that enhance group aggregation and coordination, potentially aiding collective foraging or predator avoidance in social settings.^[48] For acoustic mate recognition, variations in stridulation parameters—such as pulse rate and carrier frequency—act as species-specific filters; in crickets, females' auditory neurons selectively respond to conspecific pulse intervals (around 20-50 ms) and frequencies (4-5 kHz), preventing attraction to heterospecific signals and hybridization.^[49] These traits arise from subtle differences in stridulatory file tooth spacing, enabling reliable discrimination.^[19]

In Defense and Warning

Stridulation plays a crucial role in defense by eliciting startle responses in predators through sudden, intense sound bursts that mimic threats from larger animals. In insects like tenebrionid beetles, individuals produce rasping sounds by rubbing their abdomen against the elytra when handled or attacked, deterring predators such as lizards by interrupting their attack sequence and providing an opportunity for escape.^[50] This disturbance stridulation is widespread among Coleoptera, where it functions primarily as an anti-predator mechanism rather than for other social purposes.^[10] In toxic arthropods, stridulation serves as an aposematic warning signal to advertise unpalatability and chemical defenses, reinforcing visual cues to condition predators against future attacks. Certain cerambycid beetles employ stridulation as a disturbance or warning signal during encounters with predators, enhancing the effectiveness of their chemical repellents.^[51] Among vertebrates, analogous mechanisms appear in the lowland streaked tenrec (Hemicentetes semispinosus), where specialized quills are rasped together to generate high-frequency sounds that alarm mammalian predators or confuse them during close encounters.^[52] In scorpions, stridulation via rubbing pectines or other structures produces warning vibrations to deter predators.^[1] During high-threat situations, stridulation intensity increases, with louder and more erratic patterns amplifying the signal's urgency to maximize deterrence or group response.^[53]

Evolution of Stridulation

Phylogenetic Origins

Stridulation has evolved independently numerous times across arthropod lineages, with estimates indicating at least 84 independent origins within Heteroptera alone and at least 25 within Orthoptera, contributing to a broader pattern of repeated evolution throughout the phylum.^[2] This multiplicity underscores its polyphyletic nature, arising in diverse clades such as insects, spiders (at least 57 times), and non-insect pancrustaceans (at least 40 times).^[2] Fossil evidence supports an ancient history, with the earliest known stridulatory structures appearing in orthopteran insects during the Triassic period, over 200 million years ago, though wing venation patterns suggestive of acoustic capabilities trace back to Carboniferous winged insects around 300 million years ago.^[54]^[55] Within insects, phylogenetic mapping reveals stridulation's prevalence in derived groups like Ensifera (crickets and katydids), where tegmino-tegminal mechanisms evolved in the common ancestor approximately 300 million years ago and persist widely, and Acrididea (a major grasshopper lineage within Caelifera), where it arose at least 10 times independently.^[56] In contrast, it is absent in basal hexapods, such as entognathous orders like Collembola and Protura, reflecting its emergence after the divergence of these primitive lineages.^[57] Distribution patterns further highlight its bias toward terrestrial environments, where airborne sound propagation facilitates effective communication, compared to aquatic lineages where vibrational signals attenuate more rapidly in water and alternative mechanisms predominate.^[7] In vertebrates, stridulation is exceedingly rare and confined to isolated lineages, with single apparent origins in Squamata (certain snakes rubbing specialized scales),^[2] Siluriformes (catfish stridulating via pectoral spines),^[58] Aves (club-winged manakin rubbing specialized wing feathers),^[59] and Tenrecidae (streaked tenrecs vibrating quills).^[9] These instances represent convergent adaptations outside the arthropod dominance, emphasizing stridulation's sporadic phylogenetic footprint beyond invertebrates.

Evolutionary Drivers

Sexual selection has been a primary driver in the evolution of stridulation, particularly in insects, where female choice favors males producing complex acoustic signals that indicate genetic quality or vigor.^[2] The energetic demands of stridulation, such as elevated metabolic rates during prolonged calling, serve as honest signals of male fitness, as only robust individuals can sustain these costly displays without compromising survival.^[60] In orthopterans like crickets and katydids, this selection pressure has led to elaborated song repertoires, enhancing mating success for males with more intricate patterns.^[2] Predation pressure has similarly promoted stridulation as an anti-predator adaptation, evolving through mechanisms like aposematism and startle responses in chemically defended arthropods.^[2] In heteropterans and spiders, stridulatory sounds function defensively by warning predators or eliciting discomfort, reducing attack rates in species with noxious secretions.^[2] This selective force is evident across arthropod lineages, where stridulation correlates with higher predation risk environments, balancing survival costs against evasion benefits.^[2] Environmental factors, particularly terrestrial habitats, have favored the evolution of airborne stridulatory signals over substrate-borne vibrations, enabling long-range communication in open or vegetated settings.^[2] Sexual dimorphism in stridulatory organs is often linked to polygynous mating systems, where males invest in specialized structures for signaling, while females exhibit reduced or absent traits due to lower reproductive variance.^[61] Convergent evolution of stridulation underscores its biomechanical efficiency, with similar file-scraper mechanisms arising independently in distant arthropod clades like ensiferans and heteropterans.^[62] This polyphyly reflects shared selective advantages in producing vibroacoustic signals via simple rubbing of sclerotized body parts.^[2] Despite these benefits, stridulation incurs significant costs and trade-offs, including heightened predation risk from conspicuous sounds that attract eavesdropping predators, offset by reproductive gains in mate attraction and defense.^[2] High-frequency signals limit transmission distance, constraining use to close-range interactions and favoring evolution in structured habitats like foliage or soil.^[2]

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