Geckos are small to medium-sized lizards belonging to the family Gekkonidae, distinguished by their cylindrical or depressed bodies covered in granular or velvety scales, large protruding eyes with vertical slit pupils protected by a fixed transparent spectacle (brille) rather than movable eyelids, and specialized adhesive pads on their toes consisting of microscopic setae that enable them to climb smooth vertical surfaces and even ceilings with ease.[1]The family Gekkonidae represents the largest and most diverse group within the suborder Gekkota, encompassing approximately 1,700 species across about 64 genera, with a global distribution spanning tropical, subtropical, and some temperate regions on all continents except Antarctica.[2] Highest species richness occurs in countries such as Australia, India, Madagascar, and Malaysia, reflecting ancient evolutionary radiations dating back over 200 million years to the Mesozoic era, with ongoing discoveries continuing to increase known diversity.[3] These lizards inhabit a wide array of environments, from arid deserts and rocky outcrops to humid rainforests, woodlands, and urban areas, often favoring crevices, bark, or foliage for shelter.[1]Ecologically, geckos are predominantly nocturnal or crepuscular predators with a generalist diet focused on arthropods such as insects and spiders, though larger species may consume small vertebrates, nectar, or fruit.[4] They exhibit notable behavioral adaptations, including autotomy of the tail for escape (with regeneration possible), prehensile tails in some species for grasping, and unique vocalizations—such as chirps, barks, or screams—that serve for territorial defense, mating, or alarm, setting them apart from most other lizards.[4]Reproduction is typically oviparous, with females laying one to two hard-shelled or parchment-like eggs per clutch in concealed sites, incubated for several weeks before hatching into miniature versions of adults.[4] While many species thrive in natural habitats, some, like house geckos, have become widespread through human-mediated dispersal, highlighting their adaptability and role in controlling pest insects in human settlements.[3]
Introduction
Etymology
The word "gecko" derives from the Malay "gēkoq" or "ge'koq," an onomatopoeic term imitating the animal's distinctive vocalization, such as the chirping or barking calls made by many species during mating or territorial displays.[5] This Malay word itself is borrowed from the Javanese "tokek," which similarly mimics the sound produced by geckos like the tokayspecies.[6]The term entered scientific nomenclature in 1768 when Ignaz von Laurenti established the genus Gekko in his work Synopsis Naturae, designating Gekko gecko (the tokay gecko) as the type species; this built on Carl Linnaeus's earlier 1758 description of the same animal as Lacerta gecko in Systema Naturae. In English, "gecko" first appeared in print in 1774, reflecting European colonial encounters with Southeast Asian fauna, though earlier variant spellings like "chacco" (1711) and "jackoa" (1727) had been used.[5]In binomial nomenclature, the repetition in Gekko gecko exemplifies tautonymy, directly echoing the onomatopoeic root to honor the animal's vocal traits, a convention permitted under the International Code of Zoological Nomenclature for such cases.[6]Across Southeast Asia, geckos bear diverse cultural names rooted in their sounds or habitats, including "tokek" in Indonesian and Javanese, "tuko" or "butiki" in Filipino languages for larger species, "chee chak" or "chi chak" in Malay, and "jing-joke" in Thai; smaller species are often simply called "house lizards" (e.g., "rumah tokek" in Malay or equivalents in regional dialects) due to their prevalence around human homes.[7]
General characteristics
Geckos are small to medium-sized lizards belonging to the infraorder Gekkota of the order Squamata, encompassing more than 2,300 described species across six families.[8] This diverse clade is characterized by adaptations suited to nocturnal, scansorial (climbing), and primarily insectivorous lifestyles, distinguishing them from many other lizard groups through unique cranial modifications such as fused frontals and an edentulous palate in crown-group members.[9]A suite of unifying traits defines geckos, including the lack of movable eyelids in most species (replaced by a transparent spectacle that they clean with their tongue), nocturnal activity in the majority, the capacity for vocalization through chirping or clicking sounds, and specialized adhesive toe pads composed of lamellae that enable vertical climbing and adhesion to smooth surfaces.[10] These features, present in over 60% of species for the toe pads alone, underscore their evolutionary success in diverse habitats ranging from forests to deserts.[10]Geckos display considerable variation in size, with the smallest species, Sphaerodactylus ariasae (Jaragua dwarf gecko), reaching a total length of just 1.6 cm as adults, representing one of the minimal sizes for terrestrial amniotes.[11] At the opposite extreme, the New Caledonian giant gecko (Rhacodactylus leachianus) can exceed 40 cm in total length and weigh up to 500 grams, making it the largest extant gecko species.[12]Their general body plan features a cylindrical form with soft, delicate skin, short stout limbs adapted for agility, and disproportionately large heads housing vertical pupils and robust dentition.[10]
Physical characteristics
Body structure and size
Geckos exhibit considerable morphological diversity in body shape, adapted to various ecological niches. Robust forms, such as the tokay gecko (Gekko gecko), feature a dorso-ventrally flattened body with powerful, well-defined limbs suited for climbing on vertical surfaces.[13] In contrast, slender body shapes are common in terrestrial or sand-dwelling species, exemplified by Stenodactylus petrii, which facilitates burrowing and rapid movement across open substrates.[14] Leaf-like morphologies, seen in genera like Uroplatus (e.g., U. phantasticus), involve flattened bodies with irregular skin flaps and textured surfaces that enhance camouflage against foliage and bark in forested habitats.[14][15]Limb adaptations in geckos vary with lifestyle, though most species retain the primitive condition of five toes per foot. Fossorial species, such as Chondrodactylus angulifer, often display reduced digit lengths, particularly in the third toe, which shortens the out-lever arm to improve force generation during digging.[13] Prehensile tails, capable of grasping branches, have evolved independently in arboreal genera including Correlophus (crested geckos) and Rhacodactylus, aiding stability during climbing.[16]Skeletal features contribute to the geckos' functional versatility. The skull is notably flexible due to a kinetic construction, including a streptostylic quadrate bone that pivots freely, enabling a wide jaw gape for prey capture.[17] Unlike many lizards with dermal armor, geckos generally lack osteoderms across the body, resulting in smooth cranial bones without extensive sculpturing, which supports lightweight construction and cranial mobility.[18][19]Gecko sizes span a broad range, from the diminutive Sphaerodactylus ariasae at 18 mm snout-vent length (SVL) to the giant Rhacodactylus leachianus reaching 360 mm SVL, reflecting adaptations to microhabitats from leaf litter to tree trunks.[20] Growth follows a determinate pattern, with juveniles achieving adult proportions early; neonates closely resemble miniature adults in form due to direct development without larval stages.[20] Sexual dimorphism in size occurs in several lineages, such as the day geckos (Phelsuma), where males are typically larger than females, often exceeding them in SVL and mass.[21][22]
Skin and molting
Gecko skin consists of a thin, semi-transparent epidermis overlaid with small, granular keratin scales that provide protection while maintaining flexibility essential for climbing and navigating varied terrains. The outer epidermal layer is primarily composed of β-keratin, a protein that forms a durable yet pliable barrier against abrasion and desiccation, distinguishing it from the more rigid α-keratin dominant in other vertebrate skins. This structure allows geckos to thrive in diverse habitats, from arid deserts to humid forests, by balancing toughness with elasticity.[23][24][25]The renewal of this skin occurs through ecdysis, a complete shedding process where the entire outer layer is sloughed off in one continuous piece from the snout to the tail tip, in contrast to the fragmented molting observed in many other lizard species. This holistic shedding typically happens every 2–6 weeks in growing juveniles, influenced by factors such as temperature and metabolic rate, while adults may cycle less frequently, around every 4–8 weeks. The process is hormonally regulated, primarily by thyroid hormones like thyroxine, which accelerate epidermal cell proliferation and separation of the old stratum corneum; disruptions in thyroid function can lead to incomplete or irregular ecdysis.[26][27]Gecko skin displays intricate cryptic patterns of bands, spots, or mottling that facilitate camouflage against predators and prey by mimicking natural backgrounds such as bark or leaf litter. Embedded in the dermis are chromatophores—pigment cells including melanophores for dark tones and xanthophores for yellows—that enable subtle color shifts in response to light, temperature, or stress, enhancing adaptive concealment without the dramatic transformations seen in chameleons. These visual adaptations underscore the skin's multifunctional role in survival.[28][29]Beyond protection and camouflage, gecko skin houses embedded sensory organs, such as cutaneous mechanoreceptors, that detect substrate-borne vibrations transmitted through the body during movement or from distant sources. These low-threshold sensors provide critical tactile feedback for environmental monitoring, complementing visual and auditory cues in nocturnal foraging and predator avoidance. The skin's sensory integration also supports adhesive interactions during locomotion.[30][31]
Adhesive system and locomotion
Geckos possess a remarkable adhesive system located on the ventral surfaces of their toes, consisting of expanded toe pads covered by millions of keratinous setae, which are hair-like structures typically 10-100 μm in length.[32] These setae branch distally into hundreds of nanoscale terminal elements known as spatulae, each approximately 200 nm wide, that enable intimate molecular contact with surfaces.[33] The hierarchical structure of these setae, formed from β-keratin produced by the epidermis, maximizes surface area for adhesion while allowing flexibility.[34]The adhesion mechanism relies on reversible intermolecular forces, primarily van der Waals attractions between the spatulae and substrate, which generate sufficient force to support the gecko's body weight on vertical and inverted surfaces without the need for suction, glue, or wet adhesion.[35] This dry adhesion is highly effective on diverse substrates, including smooth glass, due to the conformal contact achieved by the spatulae, with detachment occurring through a controlled peeling motion at low angles (around 30 degrees).[36] Experimental measurements confirm that a single seta can adhere with forces up to 20 μN, scaling to over 100 N for a full gecko foot, far exceeding the animal's weight.[35]This adhesive system facilitates exceptional locomotion adaptations, enabling geckos to climb smooth vertical walls and ceilings upside-down at speeds up to 1 m/s, as demonstrated in species like Hemidactylus garnotii.[37] Certain species, such as the flat-tailed house gecko (Hemidactylus platyurus), can even run across water surfaces at speeds approaching 1 m/s by combining rapid foot-slapping to generate propulsion with hydrophobic skin that exploits surface tension, supplemented by tail undulations for stability.[38] Additionally, geckos employ tail autotomy as a locomotion-related escapemechanism, voluntarily detaching their adhesive-capable tails to distract predators while continuing to maneuver using their limbs.[36]The gecko's setal adhesion represents an evolutionary convergence with other lizards, such as anoles (Anolis spp.), where similar keratinous setae and spatulae have independently evolved for climbing, though geckos exhibit the most elaborate and versatile system with denser, finer arrays optimized for diverse habitats.[39] This repeated evolution underscores the adaptive value of fibrillar adhesives in arboreal and scansorial lifestyles across squamate reptiles.[40]
Sensory organs
Geckos exhibit highly specialized visual systems adapted to their predominantly nocturnal lifestyles, featuring large eyes without movable eyelids. Instead, their eyes are protected by a transparent spectacle, or brille, formed embryologically from the fusion of the upper and lower eyelids, which provides a fixed corneal covering. This structure is common among many gecko species and helps maintain eye moisture and protection in diverse environments. Most geckos clean and lubricate the spectacle using their tongue, a behavior that prevents debris accumulation and supports clear vision, although some genera like Correlophus and Rhacodactylus may require additional humidity to avoid issues.[41][42]Nocturnal geckos typically possess vertical slit pupils that constrict to narrow apertures during the day to reduce glare and expand at night to maximize light capture, enhancing their sensitivity in dim conditions. Their retinas consist exclusively of cone photoreceptors that have evolved rod-like properties, including larger outer segments and specialized pigments, enabling superior scotopic vision with sensitivity far exceeding that of diurnal lizards. For instance, these adaptations allow geckos to detect movement and prey in near-total darkness through a high density of visual cells and low convergence ratios in the retina. In contrast, some diurnal geckos retain multiple cone types sensitive to different wavelengths, supporting tetrachromatic color vision that aids in foraging and mate selection under brighter photic environments.[43][44][45]Auditory capabilities in geckos are tuned for survival in low-light habitats, with no external ear openings but a functional tympanic membrane that transmits vibrations to the inner ear. This system is particularly sensitive to low-frequency sounds below 1,500 Hz, facilitating the detection of approaching predators or conspecific calls from a distance. The middle ear structure, including an extrastapes connected to the angular process, amplifies these frequencies effectively, though geckos show reduced sensitivity to higher pitches compared to other lizards.[46][47]Chemoreception plays a key role in gecko sensory ecology, primarily through the Jacobson's organ, a paired vomeronasal structure in the roof of the mouth that detects pheromones and environmental odors. Unlike snakes, geckos engage in less frequent tongue flicking to sample chemical cues, often relying on direct contact or air-borne particles transferred via the tongue tip to the organ's ducts. This modality supports behaviors such as territory marking and prey location, with evolutionary modifications in the vomeronasal epithelium enhancing sensitivity to non-volatile compounds in nocturnal settings. Geckos may also briefly reference skin-based vibration sensitivity to integrate tactile cues with chemosensory input, though this is elaborated in skin adaptations.[48][49]
Dentition
Geckos exhibit pleurodont dentition, in which teeth are ankylosed to the medial surfaces of the jaw bones rather than the crest, facilitating attachment along the maxilla, premaxilla, and dentary.[50] This arrangement is typical of most squamate reptiles and contrasts with acrodont dentition seen in some basal or stem-group geckos, such as the Triassic Clevosaurus hadroprodon, where teeth fuse directly to the jaw margin without resorption during replacement.[51] The teeth are homodont, meaning they are uniform in shape across the jaw, generally conical or recurved with one to three cusps at the apex, often featuring a longitudinal groove dividing the tip; this morphology is adapted for piercing and grasping soft-bodied insect prey.[10][52]Geckos are polyphyodont, capable of lifelong toothreplacement, with successional teeth developing lingual to each functional tooth in a single row per quadrant, typically numbering around 40 teeth.[53]Replacement occurs in a wave-like pattern across the jaw, with teeth erupting approximately every 30 days in species like the leopard gecko (Eublepharis macularius), though the cycle can extend to 1-2 months in adults; resorption of the functional tooth precedes eruption of the replacement, ensuring minimal gaps in dentition.[54] This rapid turnover, driven by epithelial stem cells in the dental lamina, contrasts with the monophyodont or diphyodont patterns in mammals and supports the geckos' predatory lifestyle.[55]The jaw mechanics of geckos are enhanced by a streptostylic quadrate, a mobile articulation between the quadrate bone and the skull that allows forward and backward rotation during mouth opening, producing a wide gape of up to 180 degrees in some species.[56] This kinetic system, combined with flexible intramandibular and palatal joints, enables rapid prey capture and handling of soft-bodied invertebrates without requiring crushing force, aligning with their dentition's grasping function.[57]Variations in dentition occur among gecko genera, particularly in the diurnal Phelsuma species, which incorporate nectar, pollen, and fruit into their diet alongside insects; these geckos possess teeth with increased complexity, such as additional cusps or broader crowns, facilitating the processing of tougher plant material compared to strictly insectivorous forms.[58]
Behavior and ecology
Habitat and distribution
Geckos exhibit a cosmopolitan distribution primarily within tropical and subtropical regions across all continents except Antarctica, with native ranges extending from Australia and Southeast Asia through the Indian Ocean islands to Africa, the Americas, and the Pacific. Their highest species diversity occurs in Australasia, which encompasses the greatest number of endemic genera and species, reflecting ancient biogeographic patterns of diversification. Recent discoveries as of 2025 have described over 200 additional species, increasing global gecko diversity to more than 1,500, though Australasia maintains the highest regional richness.[14][59][60]Geckos are notably absent from extreme cold regions and high latitudes, as their thermophilic physiology limits occupancy in areas with prolonged low temperatures below their thermaltolerance thresholds. They thrive in a wide array of habitats, including humid rainforests, arid deserts, rocky outcrops, urban environments, and oceanic islands, demonstrating remarkable adaptability to varied microclimates. Many species are arboreal or scansorial, utilizing trees, walls, and cliffs, while others are terrestrial or semi-fossorial in leaf litter and soil. Nocturnal habits in many lineages facilitate exploitation of desert and urban habitats by reducing daytime heat stress and predation risk.[14][61]Human activities have facilitated the introduction of several gecko species beyond their native ranges, often via shipping and trade, leading to established populations in new regions. For instance, the tropical house geckoHemidactylus mabouia, originally from sub-Saharan Africa, has been introduced to Florida through inadvertent transport, where it now occupies urban and suburban settings.[61][62]Altitudinally, geckos range from sea level to elevations exceeding 4,000 m in mountainous areas such as the Atlas Mountains in Morocco, where species adapt to cooler, drier conditions at higher altitudes through behavioral and physiological adjustments.[14]
Diet and foraging
Geckos are predominantly insectivorous, with their diet consisting mainly of arthropods such as moths, crickets, beetles, and other small invertebrates.[63] This carnivorous focus supports their high metabolic demands, particularly in nocturnal species that hunt under low-light conditions. However, certain taxa exhibit opportunistic omnivory, incorporating plant-based foods like fruit, nectar, pollen, and sap into their diet, especially in resource-scarce environments or during periods of insect scarcity.[64] For instance, species in the genus Rhoptropus and some Australian geckos frequently consume plant exudates alongside animal prey, reflecting adaptive flexibility in trophic niches.[64]Foraging strategies among geckos vary with activity patterns and habitat. Nocturnal species, which comprise the majority, typically employ a sit-and-wait ambush tactic, remaining motionless to detect and lunge at passing prey, minimizing energy expenditure in dim environments.[65] In contrast, diurnal geckos often adopt more active pursuit behaviors, actively scanning and chasing prey across surfaces, which aligns with their higher visual acuity during daylight.[66] Tongue projection plays a limited role in prey capture compared to other lizards; geckos rely more on rapid jaw snaps and adhesive toe pads for close-range strikes rather than extended lingual projection.[67]Prey selection is influenced by gecko body size, with individuals typically targeting items up to approximately 50% of their own body length to optimize handling efficiency and energy gain. Seasonal shifts in diet are common, particularly in temperate or arid regions, where geckos may consume larger or more diverse prey during warmer months when arthropod abundance peaks, transitioning to smaller items or alternative foods in cooler seasons.[68] These adaptations ensure nutritional balance amid fluctuating food availability.Digestive processes in geckos are tuned to their predatory lifestyle, featuring relatively rapid gut transit times that facilitate quick nutrientabsorption, especially in species with elevated metabolic rates.[69] Higher environmental temperatures accelerate digestion, shortening passage times and enhancing overall foraging efficiency, which is crucial for ectothermic predators that must process meals intermittently.[69] This physiological trait supports the high-energy demands of frequent molting and locomotion in active foragers.
Reproduction and development
Geckos exhibit diverse mating behaviors, primarily involving visual courtship displays by males to attract females and deter rivals. These displays often include push-ups, tail waving, and shuddering movements, which signal male fitness and stimulate female receptivity.[70][71] In some species, such as the leopard gecko (Eublepharis macularius), males perform these actions during close encounters, escalating to mounting if the female is receptive. Vocalizations, including chirps and clicks, may accompany these displays to coordinate mating.[72]Most gecko species are oviparous, laying 1-2 hard-shelled eggs per clutch that are typically buried in moist substrate for protection and incubation.[73] However, reproductive modes vary; some, like certain New Zealand diplodactylids (Hoplodactylus and Naultinus spp.), are viviparous, retaining eggs internally until live young emerge after 5-8 months of gestation.[74] Parthenogenesis occurs in select lineages, such as the mourning gecko (Lepidodactylus lugubris), where females produce diploid eggs without fertilization, resulting in all-female clones.[75]Clutch size is characteristically fixed at two eggs in many geckos, though some species produce single-egg clutches early or late in the breeding season, potentially as an adaptive response to resource availability.[76] Females often lay multiple clutches annually, with frequencies ranging from 4-6 in temperate species like the leopard gecko over a 4-5 month breeding period, and up to 8-10 in tropical forms, spaced 15-30 days apart.[77]Incubation periods last 30-90 days, influenced by temperature; optimal ranges of 26-32°C yield higher hatchling viability, while extremes affect sex ratios in temperature-dependent species.[78][79]Hatchlings emerge as fully formed mini-adults with functional limbs, adhesive toe pads, and hunting capabilities, requiring no parental assistance beyond egg deposition.[73]Sexual maturity is reached relatively quickly, typically at 6-12 months of age, coinciding with body lengths of 50-70 mm snout-vent length (SVL) and 35-40 g in the leopard gecko, and around 40-50 mm SVL and 2-3 g in the house gecko (Hemidactylus turcicus).[80][81][82]
Social behavior and vocalization
Geckos exhibit predominantly solitary lifestyles, though certain species display more social tendencies, such as forming loose aggregations or engaging in family living. For instance, the tokay gecko (Gekko gecko) demonstrates social behaviors including prolonged pair associations and biparental care, which facilitate inter-individual interactions in shared habitats.[83] In many species, males are highly territorial, defending resources through agonistic encounters that can be influenced by prior social experience; in leopard geckos (Eublepharis macularius), males exposed to conspecifics show elevated territorial marking and activity levels compared to isolated individuals.[84] Such territoriality often manifests in resident males initiating confrontations, as observed in the Mediterranean house gecko (Hemidactylus turcicus), where adults exclude juveniles from prime areas via aggressive displays.[85]Unlike most lizards, which are largely silent, geckos produce a variety of vocalizations generated by a specialized larynx containing true vocal cords rich in elastic fibers, enabling modulated sounds for communication.[86] Common call types include barks for intimidation, distress calls during threats, short low-intensity chirps for close-range interactions, and extended complex sequences that function in territorial advertisement.[87] These vocalizations often accompany visual displays, with geckos being the most vocally diverse among lizards due to their enhanced auditory sensitivity in the 2–5 kHz range.[86] Alarm calls, such as the distinct pulse trains and sinusoidal waveforms produced by tokay geckos during predator encounters, involve specific laryngeal retraction movements to deter threats.[88]In the tokay gecko, species-specific calls like the iconic "tok-kay" sequence—characterized by multipulse rattles, bi-motifs, and optional grumbles at 0.3–4 kHz—serve primarily to proclaim territory while also signaling to potential mates.[87] These calls exhibit individual and population variations, akin to dialects, that convey identity and status within social contexts.[86] Beyond audition, geckos employ chemical signals, such as pheromones from skinlipids and fecal deposits, for self-recognition and socialdiscrimination; tokay females, for example, direct more tongue flicks toward unfamiliar conspecific odors than their own, aiding in territory marking and kin assessment.[83] Visual signals complement these, including tail wagging, push-ups, and body inflation in species like the eastern spiny-tailed gecko (Diplodactylus vittatus), which escalate during territorial disputes to assert dominance without physical contact.[89]
Predators and defenses
Geckos are preyed upon by a diverse array of predators, including birds such as owls and hawks, snakes, mammals like domestic cats and rats, and larger lizards through intra-guild predation.[90][91] These predators exploit geckos' nocturnal or crepuscular habits and small size, often ambushing them during foraging or when they are exposed on surfaces.To counter these threats, geckos rely on crypsis via camouflage, which enables them to blend seamlessly with bark, leaves, or rocks, thereby evading visual detection by avian and mammalian predators.[92] Banding patterns in many species further disrupt outlines, enhancing this protective mimicry in varied microhabitats.[93]A key anti-predator adaptation is caudal autotomy, in which geckos voluntarily detach their tail to distract attackers, allowing escape; the wriggling tail mimics the body and diverts attention, while the tail regenerates over weeks to months.[94] This pre-capture defense is widespread across gecko lineages and boosts immediate survival rates, though repeated autotomy incurs locomotor and energetic costs.[95]Certain species exhibit chemical defenses, such as tail squirting in Diplodactylus geckos, where specialized caudal glands release a sticky, unpalatable secretion that deters predators upon contact.[96] Geckos also leverage their adhesive setae for swift vertical escapes onto inaccessible surfaces, outmaneuvering ground-based pursuers.In open habitats with sparse cover, geckos face elevated predation pressure from terrestrial mammals like rats, driving strong selection for enhanced crypsis and behavioral vigilance, which can limit population densities compared to forested refugia.[93][97]
Taxonomy and evolution
Classification
Geckos are classified within the order Squamata, the lizards and snakes, specifically in the suborder Gekkota, which encompasses all gecko-like lizards distinguished by features such as vocalization ability and lack of preanal pores in some lineages.[98] The Gekkota comprises seven families, including Gekkonidae as the core and largest family with approximately 1,713 described species across 64 genera, representing the "true geckos" characterized by nocturnal habits and adhesive toe pads in many species.[99] In total, the suborder Gekkota includes about 2,450 species distributed among these families.[100]The family Gekkonidae is divided into two primary subfamilies: Gekkoninae, which includes the majority of species such as house geckos (Hemidactylus) and tokay geckos (Gekko), and Uroplatinae, comprising leaf-tailed geckos like Uroplatus from Madagascar. Species in Gekkoninae typically possess immovable eyelids fused into a transparent spectacle, aiding in eye protection and suited to their often scansorial lifestyles, whereas Uroplatinae exhibit specialized leaf-like tails for camouflage. Related families within Gekkota include Diplodactylidae (e.g., New Zealand geckos with similar adhesive setae but regional distributions) and Sphaerodactylidae (small, New World geckos often lacking extensive toe pads), which together highlight the diverse morphological adaptations across the suborder.Nomenclaturally, the classification traces back to Carl Linnaeus, who described the type species Lacerta gecko in 1758, establishing the genus Gekko via Laurenti in 1768. The family Gekkonidae was formally erected by John Edward Gray in 1825 based on morphological traits like digit structure. Modern taxonomic revisions have been profoundly shaped by molecular phylogenetic analyses, which have redefined family boundaries and revealed paraphyly in earlier groupings, as demonstrated in studies using multi-locus nuclear DNA data.
Phylogenetic history
Geckos, belonging to the clade Gekkota within Squamata, originated in the Late Jurassic approximately 160 million years ago, diverging from other squamate lineages during a period of early lizard diversification. Molecular clock estimates based on multigene analyses place the stem age of Gekkota around 225–180 million years ago, with the crown group radiation aligning with the breakup of Pangaea and subsequent Gondwanan fragmentation.[101] This divergence is supported by phylogenetic reconstructions using nuclear and mitochondrial DNA sequences, which highlight Gekkota as a basal squamate clade characterized by unique traits such as vocalization and adhesive capabilities.[102]The Gondwanan radiation of geckos led to the establishment of major clades across southern continents, with evidence from molecular phylogenies indicating vicariance events predating continental drift. For instance, pygopodid geckos, which exhibit legless, pygopod-like forms, underwent diversification in Australia, forming a monophyletic family sister to other diplodactyloid geckos.[101] Similarly, New Caledonian endemics within Diplodactylidae represent a distinct radiation, with genera like Rhacodactylus and Bavayia showing high speciation rates tied to island isolation.[103] Repeated island colonizations are evident in lineages such as Hemidactylus, which achieved global dispersal through overwater rafting, contributing to polyphyletic distributions in the Indo-Pacific.[101]Studies from the 2010s, employing comprehensive multigene datasets, have revealed widespread polyphyly in traditional gecko genera, necessitating taxonomic revisions; for example, Cyrtodactylus and Gekko are not monophyletic, with multiple independent radiations within these groups.[101] Obligate parthenogenesis has evolved independently at least six times within Gekkonidae, often in hybridogenic complexes, as inferred from nuclear and mitochondrial phylogenies that trace its origins to Cretaceous-Paleogene transitions.[104] Regarding adaptations, hair-like setae—derived from ancestral spinules—are likely plesiomorphic across Gekkota, providing a structural basis for adhesion; however, expanded adhesive toepads bearing these setae have arisen and been lost multiple times (11 gains, 9 losses), with losses occurring in lineages like pygopodids adapted to terrestrial habits.[101] This repeated evolution underscores the lability of adhesive systems in response to ecological pressures.
Fossil record
The fossil record of geckos (Gekkota) is sparse, reflecting their small body size and preference for habitats conducive to poor preservation, such as tropical forests, though some exceptional finds have provided key insights into their evolutionary history. The earliest known stem gekkotans date to the Late Jurassic, approximately 150 million years ago. In Europe, fossils traditionally assigned to Ardeosaurus brevipes from the Solnhofen Limestone of Germany represent basal members of the gekkotan lineage, exhibiting primitive cranial features like a fused postorbitofrontal bone. Similarly, Helioscopos dickersonae, described from a partial skull in the Morrison Formation of Utah, USA, confirms the presence of stem gekkotans in North America during the Tithonian stage, around 149–150 million years ago, with traits such as an edentulous palate and prominent pineal foramen indicating early divergence within the clade.[105][106]Mesozoic diversity is primarily documented from Cretaceous deposits in Asia, highlighting the clade's early radiation and acquisition of key adaptations. The oldest crown gekkotan is Cretaceogekko burmae from ~100-million-year-old Burmese amber (Albian stage), preserving a partial tail and foot with sophisticated scansorial toe pads featuring setae-like structures, evidence of adhesive capabilities akin to those in modern geckos. Other Cretaceous forms include Hoburogekko suchanovi and Gobekko cretacicus from Mongolia, dated to the Aptian–Albian and Campanian stages, respectively, which display unique postcranial and cranial traits such as an open Meckelian canal, suggesting adaptation to mesic and xeric environments. These Asian fossils indicate that geckos were already diverse and specialized for arboreal lifestyles by the mid-Cretaceous, though no unequivocal records exist from other continents during this period.[107]The Cenozoic marks a significant expansion of the gecko fossil record, with modern genera emerging around 50 million years ago in the Eocene and the clade surviving the end-Cretaceous mass extinction. Notable early Paleogene finds include Yantarogekko balticus from Eocene Baltic amber (~44–47 million years ago), a nearly complete specimen from Russia preserving scansorial adaptations and integument, assignable to early diplodactylids. By the Miocene (~23–5 million years ago), fossils referable to extant genera like Sphaerodactylus, Pygopus, and Euleptes appear across Europe, Australia, and the Dominican Republic, often in amber or sedimentary deposits, reflecting broader global dispersal and diversification post-K-Pg. This temporal range underscores geckos' resilience, with over 17 pre-Quaternary species described, though family-level assignments remain tentative due to fragmentary preservation.[107]Significant gaps persist in the gecko fossil record, particularly pre-Cretaceous and Paleogene material, attributable to their diminutive size (often under 20 mm snout-vent length) and habitation in taphonomically challenging tropical or arboreal settings, which limit fossilization opportunities. While amber inclusions have yielded exceptional soft-tissue preservation, sedimentary fossils are mostly disarticulated postcrania like maxillae and dentaries, complicating phylogenetic placements and hindering precise calibration of gekkotan divergence times. Ongoing discoveries, such as those from Laurasian deposits, continue to refine this incomplete history.[107]
Diversity and conservation
Species diversity
Geckos, belonging to the infraorder Gekkota, encompass approximately 2,400 described species, accounting for over 31% of all known lizard species globally. This remarkable diversity is unevenly distributed across seven families, with the family Gekkonidae comprising the majority, around 65% of all gecko species. High levels of endemism characterize certain regions, notably Madagascar with about 150 species and Australia with over 240 species, reflecting hotspots of gecko richness driven by unique biogeographic conditions.[3][108]Patterns of speciation in geckos are strongly influenced by island biogeography and adaptive radiation, particularly in archipelagic environments where isolation promotes divergence. For instance, the genus Cyrtodactylus exemplifies this process, with over 300 species arising through rapid adaptive radiations on islands across Southeast Asia and the Indo-Pacific, often tied to habitat specialization on karst formations and varied microhabitats. These dynamics have led to elevated speciation rates in insular settings, contributing to the overall species richness within Gekkota.Morphological diversity among geckos spans a broad spectrum, from diurnal, arboreal forms like green tree geckos adapted to foliage with vibrant coloration and adhesive toe pads for climbing, to nocturnal ground-dwellers featuring cryptic patterns for camouflage in leaf litter or soil. This variation underscores adaptive responses to diverse ecological niches, including differences in activity patterns, body size, and limb morphology across taxa.[3]Recent surveys and molecular studies indicate cryptic diversity in geckos, especially in tropical biodiversity hotspots. For example, investigations in Southeast Asia have uncovered numerous candidate species within genera like Cyrtodactylus, including at least 25 undescribed species in one analysis of 243 studied, highlighting ongoing discoveries that expand known diversity.[109]
Notable species and genera
The Tokay gecko (Gekko gecko) stands out as one of the largest gecko species, with adult males typically measuring 13–16 inches (35–40 cm) in total length and females reaching 8–12 inches (20–30 cm). Native to tropical rainforests, cliffs, and human-modified habitats across southeast and east Asia—from northeast India to the Indo-Australian archipelago—this arboreal, nocturnal species is highly territorial and solitary. It is renowned for its distinctive, loud "to-kay" vocalizations, which serve for mate attraction, territorial advertisement, and defense, often accompanied by hissing or croaking when threatened. The Tokay gecko's adhesive toe pads enable exceptional climbing ability, supporting weights up to 450 times its body mass, and its popularity in the international pet trade has resulted in established introduced populations in regions like Florida, Hawaii, and Belize, where it sometimes aids in natural pest control by consuming insects.[110][111]The crested gecko (Correlophus ciliatus), endemic to the southern forests of New Caledonia, exemplifies resilience after being presumed extinct for over a century until its rediscovery in 1994 during an expedition to the Isle of Pines. This medium-sized, arboreal species, growing to 8–10 inches (20–25 cm), features prominent eyelash-like crests above its eyes and a prehensile tail, adaptations suited to its nocturnal lifestyle in humid rainforests and shrublands. Unlike many insectivorous geckos, it is omnivorous, consuming a diet that includes arthropods, small vertebrates, fruits, nectar, and pollen, which supports its role in forest ecosystems as both predator and seed disperser. Its ease of captive breeding post-rediscovery has made it a staple in the pet trade, highlighting its adaptability while underscoring the need for habitat protection in its restricted range.[112][113]Day geckos of the genus Phelsuma represent a diverse, diurnal group with over 50 species, most of which are endemic to Madagascar and nearby Indian Ocean islands such as the Comoros, Seychelles, and Réunion. These small to medium-sized (2–12 inches or 5–30 cm) arboreal lizards are celebrated for their striking, iridescent colorations—often vibrant greens accented with red, blue, or yellow stripes and spots—that aid in camouflage among foliage and signaling during social interactions. Active during daylight hours, they forage for insects, nectar, pollen, and soft fruits in a variety of habitats, from pristine rainforests to urban gardens and plantations, demonstrating remarkable tolerance to human-altered environments. Their diurnal habits set them apart from most nocturnal geckos, contributing to their visibility and appeal in both ecological studies and the pet trade.[114]Leaf-tailed geckos of the genusUroplatus, confined to Madagascar's eastern rainforests, are exemplary nocturnal specialists renowned for their extraordinary cryptic camouflage that mimics decaying leaves and bark. Comprising approximately 22 species, these arboreal geckos feature flattened bodies, broad heads with horn-like projections in some cases, and leaf-shaped tails with irregular edges and mottled brown, gray, or reddish hues, allowing them to rest motionless during the day to evade predators. Active at night, they hunt insects and small invertebrates low in the forest canopy, typically within 10 feet (3 m) of the ground, relying on keen eyesight and stealth rather than vocalizations. Iconic species like the satanic leaf-tailed gecko (U. phantasticus), reaching just 3.5 inches (9 cm), showcase extreme adaptations such as skin flaps and a pose that enhances their leaf-like disguise, making them vital indicators of rainforest health.[115][116]
Conservation status
Geckos face varying levels of extinction risk, with approximately 16% of assessed species (374 out of 2,389) classified as threatened (Vulnerable, Endangered, or Critically Endangered) on the IUCN Red List as of October 2025, including over 100 species vulnerable primarily due to habitat destruction in regions like Madagascar, where deforestation has fragmented critical forest habitats for endemic taxa such as day geckos in the genus Phelsuma.[117][118][119]The principal anthropogenic threats include habitat loss from deforestation and agriculture, competition and predation by invasive species, and overexploitation via the international pet trade. For instance, the turquoise dwarf gecko (Lygodactylus williamsi), endemic to a small area in Tanzania, has experienced severe population declines from illegal collection, with thousands of individuals seized in smuggling operations between 2004 and 2012.[120][121]Conservation efforts encompass the designation of protected areas in key biodiversity hotspots, CITES Appendix II listings for over 180 gecko species to regulate international trade, and targeted captive breeding programs. In February 2025, the IUCN Species Survival Commission established the Gekkota Lizard Specialist Group to coordinate global conservation actions for the nearly 2,400 species. Notable programs have bolstered populations of critically endangered species, such as the Christmas Island giant gecko (Cyrtodactylus sadleiri) through ex-situ breeding and reintroduction trials, and L. williamsi via seizures that yielded around 40,000 individuals for propagation.[8][120]Climate change poses an escalating risk, driving range shifts toward cooler elevations and projected local extinctions for many island-endemic geckos by 2050, as thermal limits constrain dispersal on fragmented oceanic habitats.[122]