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Gekko

Gekko is a genus of geckos belonging to the family Gekkonidae, consisting of over 90 species that are primarily distributed across Southeast and East Asia, from northeast India through Indonesia, the Philippines, and into parts of China and Japan. These "true geckos" or "calling geckos" are notable for their distinctive vocalizations, which distinguish them from many other geckos, and their specialized adhesive toe pads composed of setae that enable them to climb vertical surfaces and even walk on ceilings. The genus includes some of the largest gecko species, such as the iconic Tokay gecko (G. gecko), which can reach lengths of up to 40 cm and is known for its loud, barking calls resembling "to-kay." Taxonomically, Gekko has undergone significant revision in recent decades, with phylogenetic studies identifying six major morphological species groups: the G. gecko, G. japonicus, G. monarchus, G. petricolus, G. porosus, and G. vittatus groups, supported by molecular and morphological data. The type species, Gekko gecko (Linnaeus, 1758), exemplifies the genus's diversity, while ongoing discoveries have expanded the recognized species count, with approximately 25 species native to as of 2025. Distribution patterns reveal a concentration in tropical and subtropical forests, urban areas, and rocky habitats, though some species have been introduced to other regions like the via the pet trade. Members of Gekko are predominantly nocturnal and arboreal, feeding on , small vertebrates, and occasionally fruits, with powerful limbs and a robust body adapted for their lifestyle. Their skin is soft and granular, often featuring cryptic patterns for , and many exhibit tail as a defense mechanism against predators. The genus's ecological role includes in settlements, but loss and overcollection for pose threats to several populations.

Taxonomy

Etymology and History

The genus name Gekko originates from the term "gekko," derived from the Indonesian-Malay word gēkoq, which itself is borrowed from Javanese tokek and serves as an onomatopoeic imitation of the vocalization produced by the (Gekko gecko), a prominent species in the . This etymological root reflects the audible "gecko" or "tokay" calls characteristic of several Southeast Asian , emphasizing the auditory distinctiveness that first drew scientific attention to the group. The taxonomic history of Gekko commenced with Carl Linnaeus's 1758 description of the type species Lacerta gecko (now Gekko gecko) in Systema Naturae, based on specimens from Southeast Asia, though the genus itself was formally established by Ignaz Laurenti in 1768, who designated Lacerta gecko as the type by tautonymy. Throughout the 19th and early 20th centuries, the genus expanded significantly with European explorations and collections from Southeast Asia; by the late 1800s, George Albert Boulenger's 1885 Catalogue of Lizards in the British Museum recognized only eight species, providing a foundational systematic overview that cataloged their morphological variations and distributions. Further growth occurred in the mid-20th century through fieldwork by Edward H. Taylor (1920s–1960s), who added five species (four of which remain valid), and Walter C. Brown and Angel C. Alcala (1960s–1970s), who described three additional species from the Philippines, reflecting increased focus on regional biodiversity. In the late 20th and early 21st centuries, the underwent substantial revision, with Arnold G. Kluge listing 28 in 2001, followed by 14 more additions since 2004 driven by intensified surveys in as of 2011. Since then, numerous additional have been described, bringing the total to approximately recognized as of 2025. Molecular phylogenetic analyses, incorporating mitochondrial and nuclear data, have since refined boundaries, identifying six distinct phenotypic groups and revalidating taxa like G. reevesii (previously synonymized), while reassigning some former members from related genera such as Luperosaurus. Key contributors include Herbert Rösler, who described like G. ernstkelleri and co-authored phylogenetic syntheses, and Aaron M. Bauer, noted for delineating G. nutaphandi and advancing integrative through combined morphological and genetic approaches.

Classification and Phylogeny

The genus Gekko is classified in the kingdom Animalia, phylum Chordata, class Reptilia, order , suborder Gekkota, family , subfamily Gekkoninae. Within , Gekko occupies a basal phylogenetic position, forming a to other Asian geckos in the subfamily Gekkoninae. This placement is supported by molecular analyses, including sequences of 16S rRNA and , which indicate early divergences within the family during the . For instance, studies using mtDNA and nuclear markers have resolved Gekko as part of the Southeast Asian mainland and clade, with strong support for its deep branching relative to more derived groups like Philippine endemics. A 2011 study subdivided the genus into six morphological and genetic species groups: the G. gecko group, G. japonicus group, G. monarchus group, G. petricolus group, G. porosus group, and G. vittatus group. A more recent 2020 phylogenetic analysis using thousands of ultraconserved elements proposed a revised subgeneric classification comprising seven subgenera: Gekko (sensu stricto), Japonigekko, Gekkotula, Hemidactylus-like group wait no, actually the subgenera are Gekko, Ptychozoon (as subgenus), Luperosaurus (some), but precisely: the paper defines seven subgenera including Gekko, Japonigekko, Gekkoella, Monarchus, Ptychozoon, and others to accommodate the diversity. Recent taxonomic revisions based on genetic data have elevated former subspecies to full species status, such as G. japonicus (previously considered a subspecies of G. gecko) and the revalidation of G. reevesii. These splits reflect cryptic diversity uncovered by , emphasizing the role of genetic analyses in refining Gekko's .

Description

Morphology

Members of the genus Gekko are characterized by a robust build, including a large head distinct from the neck, a cylindrical body that is somewhat dorsoventrally compressed, and a that can exceed the snout-vent length (SVL). These exhibit a size range with SVL typically spanning 40–180 mm across species, with larger forms like G. gecko reaching up to 177 mm SVL. Males possess distinctive preanal and , numbering 20–40 in total, arranged in a continuous or interrupted series along the ventral midline and thighs, which serve as glandular structures. The limbs of Gekko species are well-developed and robust, supporting an arboreal , with each foot bearing five digits equipped with expanded lamellae on the ventral surface. lamellae vary across species groups, with some exhibiting reduced scansors adapted to specific habitats. These lamellae consist of millions of microscopic setae—hair-like projections approximately 100 μm long, with terminal spatulae tapered to 0.2–0.5 μm at the tip—that enable via van der Waals intermolecular forces, allowing the to climb smooth vertical surfaces without residue. This setae microstructure maximizes contact area and force distribution, with adhesion strengths of 20–65 N/cm² reported for gecko feet. Sensory adaptations in Gekko include large, prominent eyes with vertical pupils and no movable eyelids; instead, a transparent covers the , which the clean and moisten by licking with their . Unique among , many Gekko species possess true in the , enabling production of loud, species-specific calls for communication, a trait shared with other "true geckos" in the . Skeletal traits of Gekko feature a robust cranium and adapted for agility, with some developing osteoderms—dermal bony plates—that form a cephalic shield over the skull and scattered postcranial elements, appearing postnatally at SVL around 100 mm. Fossil records of Gekkota, the clade including Gekko, extend back to the (approximately 125–100 million years ago), with specimens known through the (23–5 million years ago), underscoring the ancient origins of these morphological features. Coloration patterns in Gekko often integrate with this body form to enhance on or foliage.

Coloration and Variation

Species in the genus Gekko typically display cryptic coloration adapted for blending into arboreal environments, featuring a mottled gray-brown or bluish-gray base with spots, bands, or flecks that mimic bark and foliage. For instance, Gekko gecko exhibits a gray body overlaid with brownish-red to bright red spots and smaller flecks, while other species like Gekko reevesii show variations such as reddish-olive dorsum marbled with grayish tones. These patterns facilitate background matching and disruptive to evade predators. Sexual dimorphism in coloration is evident in several Gekko species, with males generally possessing brighter hues and more pronounced patterns than females. In Gekko gecko, males display vivid red or orange spots on a background, often accompanied by colorful throat patches, whereas females have duller, less saturated colors for the same base pattern. This dimorphism extends to other congeners, where male-specific brighter pigmentation may enhance visibility during interactions, though females maintain subtler tones. Ontogenetic changes in coloration occur within the , with juveniles often exhibiting more vivid and contrasting spots or bands that fade or become less distinct in adults. For example, young Gekko gecko show intensified spotting for enhanced during early vulnerability, which mellows with maturity to a more uniform mottled appearance. Geographic variation further diversifies patterns, such as island populations of Gekko gecko displaying distinct spot hues or band intensities compared to mainland forms, reflecting local environmental adaptations. The adaptive significance of these colorations lies in predator avoidance through visual , where spotted and banded patterns disrupt outlines against arboreal substrates, and in through physiological color changes observed in species like Gekko gecko, allowing skin to lighten or darken to match surroundings.

Distribution and Habitat

Geographic Range

The Gekko is native primarily to , encompassing regions from northeastern and southern through Indochina, the , , and the , with extensions northward to southern Japan (including the ) and westward across the Pacific to islands such as , the , and . This distribution spans diverse biogeographic zones, including the , continental , and the Sunda and shelves. Introduced populations of Gekko species, most notably G. gecko, have become established outside their native range due to the international pet trade, accidental transport via shipping, and human-mediated dispersal. These include feral populations in Florida and Hawaii (United States), Guam, New Zealand, and parts of Africa such as Madagascar and South Africa. Biogeographic patterns within the reveal high levels of , with many of its approximately 120 recognized restricted to specific islands or regions, driven by historical vicariance events like changes and overwater dispersal via , supplemented by recent spread. richness hotspots occur in , the , and , where over 20 are documented in and high diversity is found in the Philippine and Indonesian archipelagos, reflecting their role as centers of diversification. The continues to see new descriptions as of 2025.

Habitat Preferences

Species of the genus Gekko primarily inhabit tropical and subtropical ecosystems across and parts of , favoring tropical rainforests, secondary forests, and disturbed areas such as plantations and urban environments. These geckos are predominantly arboreal, utilizing trees, cliffs, and rock faces, though some species exhibit saxicolous preferences in landscapes. For instance, Gekko gecko thrives in both primary and secondary forests as well as human-modified habitats like buildings and agricultural zones, demonstrating high adaptability to disturbances. Microhabitat selection emphasizes sheltered, elevated sites for refuge and foraging, often in close proximity to water sources such as wetlands or . Gekko species commonly occupy bark crevices, tree holes, and rock fissures in natural settings, while in areas, they exploit structures like walls, ceilings, and for . This versatility allows them to persist in fragmented landscapes, including plantations and suburban orchards, where they perch at heights of approximately 2 meters on vertical surfaces. The occupies a broad altitudinal gradient, from to elevations exceeding 2000 meters, with some montane species recorded up to 2542 meters above . Lower-elevation species like Gekko gecko are typically found below 1200 meters in lowland forests and human habitations, while higher-altitude taxa, such as those in the Japonigekko, inhabit montane forests. Climate adaptations center on high-humidity environments (typically 70-90%) characteristic of tropical regions, where nocturnal activity helps mitigate daytime heat and risks. Gekko species require warm, humid conditions, with G. gecko observed in areas averaging 32°C and 58-90% relative humidity, often near water bodies to maintain hydration. This preference for moist microclimates supports their and adhesive locomotion on wet surfaces.

Behavior and Ecology

Activity and Locomotion

Gekko species, such as the (Gekko gecko), exhibit strictly nocturnal circadian patterns, remaining sheltered in crevices or foliage during daylight hours to avoid predation and , and emerging shortly after sunset for peak activity. This nocturnal rhythm aligns with their in tropical and subtropical environments, where cooler nighttime temperatures facilitate movement and . Territorial defense during these active periods often involves vocalizations, including the iconic repetitive "gecko-gecko" or "tokay" calls produced by males, which can reach intensities of approximately 70 at 1 meter and serve to deter intruders over distances exceeding 100 meters. Locomotion in Gekko geckos is characterized by exceptional adhesive capabilities, enabling rapid on vertical and inverted surfaces at speeds ranging from 0.6 to 1.2 m/s, primarily achieved through increasing stride rather than length. These movements rely on the morphological adaptations of their subdigital setae, which generate van der Waals forces for without reliance on sticky secretions. As an escape strategy, individuals can voluntarily detach their tails via caudal when grasped by predators, allowing the wriggling detached tail to distract the threat while the gecko flees; the tail regenerates over several weeks, though the regenerated structure is structurally weaker. Sensory behaviors in Gekko support and environmental in low-light conditions, with chemosensory tongue flicking playing a key role in detecting chemical cues from substrates or conspecifics for orientation and territory assessment. Cutaneous sensilla distributed across the body, particularly on the and digits, provide mechanoreceptive feedback essential for precise and substrate contact, potentially contributing to broader sensory including gradients through general integumentary sensitivity. Social interactions among Gekko are predominantly territorial, featuring displays such as vocal barking, body inflation, and tail waving to assert dominance, though overt physical aggression remains minimal outside of breeding periods when males may engage in combat over mates or resources. These behaviors help maintain spacing in dense habitats, reducing competition without frequent injurious encounters.

Diet and Foraging

Gekko species are predominantly insectivorous, with their diet consisting mainly of arthropods such as moths (), beetles (Coleoptera), crickets, and other orthopterans. For instance, the Tokay gecko (Gekko gecko) frequently preys on locusts, grasshoppers, , , and mosquitoes, reflecting a preference for mobile, nocturnal . They display opportunistic feeding, occasionally consuming small vertebrates like and frogs in certain species. Foraging in the genus Gekko typically involves a sit-and-wait strategy, where individuals perch motionless on vertical surfaces like trunks or walls, ambushing passing prey with rapid strikes. This behavior is supported by acute visual hunting capabilities, enabled by large eyes with high sensitivity to low light—up to 350 times greater than in humans—allowing precise detection of during their nocturnal activity periods. Ecologically, Gekko geckos serve as important predators in their habitats, helping to regulate insect populations in agricultural and settings through their consumption of economically significant arthropods like and mosquitoes. They occasionally engage in , particularly larger individuals preying on juveniles, which may occur under conditions of resource limitation or high densities.

Reproduction

Gekko species exhibit polygynous systems, where males defend territories and court multiple females during the season, which typically spans 4-5 months. Males engage in aggressive with rivals, employing visual displays such as arching the body, extending limbs, and producing rasping acoustic calls around 4000 Hz to establish dominance. involves males approaching females with multiple chirp calls (frequencies 500-6000 Hz) followed by visual displays, including raising the body, arching the back, distending the throat to reveal coloration, and lateral head movements to attract mates. Copulation occurs when the male grasps the female's , and females can store sperm to fertilize subsequent clutches. All Gekko are oviparous, with females laying clutches of two hard-shelled, eggs that are glued to vertical surfaces for protection. A single female typically produces 1-4 clutches per year, depending on environmental conditions and nutritional status, with eggs laid approximately 30 days after . lasts 90-120 days at temperatures of 26-30°C (80-86°F), though lower temperatures can extend this to 200 days; optimal conditions yield independent hatchlings measuring about 8-10 cm in total length. Sexual maturity is reached at 1-2 years of age, with males often maturing slightly earlier than females based on size and weight thresholds around 90 grams. In the wild, Gekko geckos have a lifespan of 7-10 years, though captivity can extend this to 15-20 years due to reduced predation and consistent resources. is absent, as hatchlings are fully independent upon emergence and receive no post-hatching attention from adults. Parthenogenesis is rare in the Gekko , with primarily sexual and genotypic sex determination prevailing across species. Most species possess a diploid number of 2n=38, consisting of a mix of metacentric, submetacentric, subtelocentric, and telocentric elements, though slight variations (e.g., 2n=34-42) occur in some taxa due to evolutionary rearrangements. , such as larger size and brighter throat coloration in males, aids in mate attraction during .

Conservation

Threats

Habitat destruction poses a primary threat to Gekko populations across their Southeast Asian range, driven by for , , and . In regions like and , rapid forest clearance has led to significant population declines, with studies documenting up to 50% reductions in local abundances of species such as Gekko gecko over recent decades. plantations and urban expansion particularly displace arboreal species reliant on tree bark and rock crevices, fragmenting habitats and reducing available refugia. The international pet and trade exacerbates these pressures, with overcollection targeting popular like the (G. gecko). Indonesia alone exported approximately three million individuals annually from 2015 to 2021, primarily as dried specimens for medicinal use, though much of this trade is illegal or unsustainable. Live exports for the pet market add further strain, with quotas often exceeded and laundering of wild-caught animals common, threatening endemic in and elsewhere. Additional anthropogenic factors include , which alters levels critical for Gekko function and , potentially shifting suitable ranges for various endemic species. use in agricultural areas reduces prey availability, indirectly impacting insectivorous Gekko populations. In introduced or disturbed habitats, competition from invasive geckos can displace native Gekko through resource monopolization. Natural predators, including birds of prey, snakes, and larger lizards, exert secondary pressure on Gekko populations but are overshadowed by human-induced threats.

Status and Protection

The conservation status of species in the genus Gekko varies widely, with assessments primarily conducted by the International Union for Conservation of Nature (IUCN) Red List. Of the approximately 120 recognized species, a significant portion—particularly recently described endemics—are listed as Data Deficient due to insufficient data on population sizes, distributions, and trends, with around 20% of assessed species classified as threatened (Vulnerable or higher). Ongoing discoveries, such as the description of Gekko septentrionalis in 2025, highlight the need for rapid assessments, as many new species are initially classified as Data Deficient. For instance, Gekko gigante, endemic to the Gigantes Islands in the Philippines, is assessed as Vulnerable owing to habitat loss and limited range, and Gekko palawanensis, restricted to Palawan, is categorized as Near Threatened based on ongoing declines from deforestation and collection pressures. These evaluations highlight the genus's vulnerability, especially for island endemics, though widespread species like Gekko gecko remain Least Concern despite local declines. Legal protections for Gekko species focus on regulating and habitat safeguarding. The (G. gecko), heavily exploited for and pets, has been listed in Appendix II of the since 2019, requiring export permits to monitor and prevent overexploitation. In range countries, endemic species benefit from national legislation; for example, Philippine endemics such as G. gigante are protected within marine and terrestrial reserves like the Protected Landscape, while Indonesian populations are safeguarded in areas like . These measures aim to curb illegal , which briefly contextualizes the need for such interventions amid rising demand. Conservation actions for Gekko emphasize ex situ and efforts to bolster populations. Captive breeding programs, particularly for G. gecko, have been developed in regions like , , to supply sustainable sources for trade and reduce wild harvesting, with successful reproduction protocols established to support reintroduction potential. Habitat restoration initiatives in the target forests critical for narrow-disked geckos, involving and control to enhance suitable microhabitats. Additionally, research on is advancing, using molecular tools to assess connectivity among fragmented populations and guide targeted interventions for threatened endemics. Monitoring of Gekko species employs non-invasive techniques to track elusive populations, especially for taxa. Camera traps have proven effective for detecting nocturnal activity in forested habitats, providing density estimates for species like G. gecko in Southeast Asian reserves. (eDNA) sampling from soil and water is increasingly applied to rare Philippine endemics, enabling detection without disturbance and aiding in mapping distributions for planning. These methods facilitate ongoing assessments and to address knowledge gaps across the genus.

Species

Diversity

The genus Gekko comprises 120 recognized as of November 2025, reflecting ongoing taxonomic revisions and discoveries in . Since 2020, at least eight new have been described, including Gekko fengshanensis and Gekko septentrionalis in 2025, highlighting the genus's dynamic alpha taxonomy amid intensified surveys in habitats. Species diversity is concentrated in the Indo-Malaya region, where over 60 occur across , including mainland countries such as and , and island nations such as , driven by habitat heterogeneity and historical biogeographic barriers. Island endemism is pronounced, particularly in the , which harbors approximately 15 restricted to its , such as those adapted to fragmented volcanic and ecosystems. Morphologically, the genus exhibits wide variation, ranging from dwarf forms under 10 cm in snout-vent length (e.g., Gekko japonicus) to giant exceeding 30 cm (e.g., Gekko gecko), with differences in scalation, limb structure, and adhesive toe pads reflecting ecological adaptations. Evolutionary diversification within Gekko accelerated post-Miocene, approximately 10–5 million years ago, coinciding with tectonic uplift and sea-level fluctuations that fragmented habitats and promoted across archipelagos. Hybridization remains rare but has been documented in contact zones, such as between Japanese congeners like Gekko japonicus and Gekko tawaensis, where genomic analyses reveal limited despite . Taxonomic history involves significant synonymy issues, with over 100 names historically lumped under broad species concepts like G. gecko; molecular phylogenies using mitochondrial and nuclear DNA have resolved many of these, elevating former subspecies to full species status and stabilizing the genus's boundaries.

Notable Species

The (Gekko gecko) is the largest in the genus, reaching lengths of up to 40 , and is widely distributed across southern , , and , where it inhabits tropical rainforests, cliffs, and structures. This has been introduced to various regions outside its native range, including , , and the , where it establishes feral populations and is considered invasive in some areas due to its adaptability and predatory behavior. In , G. gecko holds significant medicinal value in , where dried specimens are used to treat ailments such as and impotence, contributing to substantial trade volumes estimated at up to 200,000 individuals annually in some markets. Culturally, the features prominently in n folklore as a symbol of and , often interpreted as an when heard calling at night, and valued for controlling household pests like insects. Gekko japonicus, known as Schlegel's Japanese gecko, is primarily associated with , where it forms relictual populations in urban and forested areas, and is designated as a special with total protection status due to threats from habitat loss driven by urban development and potential hybridization with non-native lineages. This species exhibits low in some regions, reflecting historical bottlenecks, and is classified as Least Concern on the , underscoring its vulnerability in Japan's fragmented landscapes. The giant forest gecko (Gekko smithii), also called Smith's green-eyed gecko, is a large attaining up to 35 cm in length and is specialized for arboreal life in the humid lowland and hill forests of and , where it forages nocturnally on trees and occasionally enters dwellings. Its striking green eyes and robust build distinguish it among Gekko , with populations in representing key hotspots for the . In 2025, Gekko shiva was described as a new limestone-dwelling species from caves and hills in , eastern , near the Cambodian border, highlighting ongoing discoveries in karst ecosystems amid . Preliminary traits include a robust body with long limbs, short digits, a long head bearing pinkish tongue and relatively large eyes, and a greenish-golden iris, differentiating it from congeners like G. pradapdao through scalation and meristic counts such as 80–86 mid-dorsal scale rows. Similarly, Gekko alpinus, a high-altitude specialist in the subgenus Japonigekko, was discovered in 2024 from the Basin in the of , at elevations of 2,400–2,542 m, marking the highest recorded range for the subgenus and emphasizing montane refugia for diversification. This species exhibits mosaic-like scalation and adapted for cold, rocky terrains, with preliminary observations noting its nocturnal habits in shrublands.

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