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Iberian ribbed newt

The Iberian ribbed newt (Pleurodeles waltl), also known as the Spanish ribbed newt, is a large-bodied salamander species belonging to the family Salamandridae, characterized by its broad, flattened head, granular skin with small warts, and a distinctive defensive mechanism where sharp costal ribs can protrude through lateral pores in the skin when threatened, combining with skin toxins for protection.^[1] Native to the central and southern Iberian Peninsula (Spain and Portugal) and northwestern Morocco, it inhabits a variety of aquatic environments including permanent and temporary ponds, slow-moving streams, marshes, lagoons, and ditches, often preferring deeper water with some vegetation and tolerating moderate organic pollution or salinity.^[1] Adults are largely aquatic, rarely venturing onto land except during rainy migrations between breeding sites, and can grow to lengths of up to 312 mm in males and 286 mm in females, with a gray-brown dorsum sometimes marked by yellowish or orange spots aligned with the ribbed tubercles.^[1] The species is carnivorous, feeding on aquatic invertebrates such as mollusks, worms, and insects, and reproduces by laying clusters of 150–1,300 eggs in water, with larvae hatching after about 13 days and metamorphosing in 3–4.5 months depending on temperature.^[1] Although resilient to some environmental stresses, populations face threats from habitat loss, water pollution, and infrastructure development, leading to its classification as Near Threatened on the IUCN Red List due to inferred declines of less than 30% over ten years in fragmented habitats.^[1]^[2]

Taxonomy and Description

Taxonomy

The Iberian ribbed newt (Pleurodeles waltl) is classified within the kingdom Animalia, phylum Chordata, class Amphibia, order Caudata, family Salamandridae, subfamily Pleurodelinae, genus Pleurodeles, and species P. waltl.^[3] This placement positions it among the true salamanders, characterized by their tailed, lizard-like form and aquatic-terrestrial life cycles. The species was first described by Ignaz Rudolf Michahelles in 1830 based on specimens from near Cádiz, Spain.^[3] The genus name Pleurodeles originates from the Greek words pleuron (meaning "side" or "rib") and delos (meaning "evident" or "visible"), alluding to the prominent, rib-like costal grooves visible along the sides of its body.^[4] The specific epithet waltl honors the German naturalist Joseph Waltl (1805–1888), who collected early specimens of the species during expeditions in Spain.^[1] Historical synonyms include Salamandra funebris Bory de Saint-Vincent, 1828; Triton costatus Wagler, 1830; Salamandra major Gray in Cuvier, 1831; Bradybates ventricosus Tschudi, 1838; and Pleurodeles exasperatus Duméril, Bibron, and Duméril, 1854, with common misspellings such as Pleurodeles waltlii.^[3] Phylogenetically, Pleurodeles belongs to the diverse family Salamandridae, with the genus diverging from its closest living relatives, such as the Asian genus Tylototriton, approximately 8.6–10 million years ago during the Late Miocene.^[5] Within the genus, P. waltl forms a distinct clade separate from North African congeners like P. poireti and P. nebulosus (revalidated in 2020 based on molecular and morphological data), with an estimated divergence time of around 5.3 million years ago from P. poireti.^[5]^[6] The genus likely originated in the Iberian Peninsula, as evidenced by upper Miocene fossils (approximately 11–5 million years ago), before dispersing to northwest Africa via temporary land bridges.^[1] The genus Pleurodeles currently includes three recognized species: P. waltl, P. poireti (endemic to northeastern Algeria), and P. nebulosus (distributed in Algeria and Tunisia).^[6] Intraspecific genetic analyses reveal two main clades within P. waltl—one in southeastern and eastern Spain plus northern Morocco, and another in Portugal and southwestern/central Spain—with divergence estimated at 3.2–2 million years ago in the Pliocene.^[5]

Physical Characteristics

The Iberian ribbed newt (Pleurodeles waltl) is the largest species of newt in Europe, with adults reaching a maximum total length of 312 mm in males and 286 mm in females in the wild, though specimens from North Africa tend to be smaller than those from Europe.^[1] In captivity, individuals typically grow to 150–200 mm, reflecting constrained conditions compared to natural habitats.^[7] The body is robust and dorsoventrally flattened, with a tail length generally shorter than the snout-vent length, and females exhibiting proportionally shorter tails than males.^[1] Dorsally, the newt displays a dark gray to brown coloration, often accented by irregular dark spots and a series of 7–10 yellow-orange warts along the dorso-lateral ridges.^[1] The ventral surface is lighter, typically beige or yellowish with scattered black spots, providing camouflage in aquatic environments.^[8] Larvae, hatching at approximately 11 mm, are pale and translucent with dark flecks, adopting an elongated, fish-like form before developing limbs.^[1] Key anatomical features include a broad, flattened head with moderately large eyes positioned dorsally, and a skin texture covered in small, granular warts rich in poison glands.^[1] Along the flanks, the sharp ends of the costal ribs are conspicuous and can protrude through the skin via specialized muscular contractions.^[1] The limbs are sturdy with webbed feet adapted for swimming, and males develop black nuptial pads on the forelimbs during the breeding season.^[1] Larvae possess external gills for respiration, while adults rely on lungs and cutaneous breathing, supporting a predominantly aquatic lifestyle.^[9]

Distribution and Habitat

Geographic Distribution

The Iberian ribbed newt (Pleurodeles waltl) is native to the central and southern Iberian Peninsula, spanning Portugal and much of Spain, as well as northwestern Morocco.^[1] In Portugal, the species occurs throughout the country, with higher abundances in the southern regions characterized by warmer climates. In Spain, its range covers the southern half of the peninsula, extending northward to near León, crossing the Duero River in multiple locations and the Ebro River along the Catalan coast, with the eastern limit on the Mediterranean shore and the southern boundary at Tarifa near the Strait of Gibraltar. In Morocco, populations are confined to a triangular area in the northwest, encompassing humid, semi-humid, and semi-arid zones between Talmagayt, Safi, and Île d'Essaouira.^[1] Fossil records trace the origins of the genus Pleurodeles to the Iberian block during the upper Miocene epoch, approximately 11.6 to 5.3 million years ago, with evidence of migration to Africa via land bridges during the Messinian salinity crisis around 5.6 million years ago.^[1] Subsequent differentiation of North African lineages occurred in the Pliocene epoch following the formation of the Strait of Gibraltar. Quaternary fossils, including early Pleistocene remains attributed to P. cf. waltl, have been found in Moroccan sites such as Irhoud, supporting continuity in the African range.^[10] Overall, the species is not considered abundant throughout its distribution, with eastern Spanish populations particularly scattered.^[11] Post-glacial dynamics indicate limited range expansion beyond southern refugia established during the Pleistocene, maintaining the current bounded extent without major northward shifts.^[12]

Habitat Preferences

The Iberian ribbed newt (Pleurodeles waltl) is a highly aquatic species that inhabits a variety of still-water environments across its range, including permanent ponds, lagoons, marshes, slow-moving streams and rivers, and man-made structures such as cisterns and wells. These habitats are typically characterized by calm, low-current waters, with a preference for depths of at least 1 meter to support year-round occupancy. The newt favors deep, cool, vegetated waters that provide cover and foraging opportunities, though it can occupy sites with sparse or no aquatic vegetation.^[1]^[13]^[7] Water quality plays a key role in habitat selection, with the species tolerating moderate organic pollution and low to moderate salinity but avoiding fast-flowing or heavily contaminated waters. Adults remain fully aquatic throughout their lives, rarely venturing onto land except during brief migrations to nearby moist refuges when primary water bodies dry during droughts; such movements occur primarily on rainy days to locate new aquatic sites. Larvae develop in shallow margins of ponds.^[1]^[7]^[14] Seasonal variations influence habitat use, with individuals overwintering in deeper, stable water bodies to avoid surface freezing or drying. Breeding occurs predominantly in temporary pools with extended hydroperiods during spring and summer, when rainfall replenishes shallow sites suitable for egg-laying and larval growth. These pools often form in Mediterranean scrub, open woodlands, or cultivated lands, ensuring access to prey-rich environments post-metamorphosis.^[1]^[14]^[15]

Behavior and Ecology

General Behavior

The Iberian ribbed newt (Pleurodeles waltl) exhibits primarily crepuscular and nocturnal activity patterns, with individuals becoming more active during twilight and nighttime hours, particularly outside of their extended aquatic phases. This behavior aligns with their secretive nature, allowing them to forage and move while minimizing exposure to diurnal predators. Although predominantly aquatic throughout much of their life, they occasionally emerge to bask on emergent vegetation or shorelines during favorable conditions, such as mild weather. In the wild, these newts have a lifespan of 7–15 years, reflecting their resilience in varied Mediterranean environments.^[16]^[1] Socially, P. waltl is largely solitary, with individuals maintaining independent territories and showing no observed territorial aggression outside of brief breeding aggregations. Interactions are minimal except during the reproductive season, when males and females congregate in aquatic habitats, but even then, post-mating dispersal leads to resumed isolation. This solitary structure supports their low-density populations in fragmented habitats.^[1] The life cycle of P. waltl includes a distinct larval stage lasting 3–4.5 months, during which hatchlings develop external gills for aquatic respiration and grow to 30–60 mm in length before undergoing metamorphosis. Metamorphosis transforms larvae into juvenile forms, typically occurring in spring or early summer, with some individuals exhibiting paedomorphosis—retaining larval traits such as gill-like structures or neotenic morphology into adulthood under certain environmental conditions. This variability enhances adaptability in unstable water bodies.^[16] Physiological plasticity in P. waltl enables seamless transitions between aquatic and terrestrial phases, driven by environmental cues like seasonal rainfall, temperature, and pond desiccation. During wet periods, they favor fully aquatic lifestyles for feeding and breeding, while in dry seasons, they aestivate terrestrially under rocks or in burrows, reducing metabolic demands until water returns. This biphasic cycle underscores their resilience to Mediterranean climate fluctuations.^[17]

Defense Mechanisms

The Iberian ribbed newt (Pleurodeles waltl) employs a remarkable mechanical defense through rib protrusion, where sharp rib tips penetrate the skin to form defensive spines when threatened. This mechanism involves the rotation of ribs anteriorly up to 65 degrees via a specialized two-headed costo-vertebral joint, allowing the pointed ends to pierce through lateral skin "warts" without causing significant self-damage, as the punctures are shallow and heal rapidly.^[18]^[19] The process is triggered by physical stimuli, such as handling or attack, and is faster in the aquatic larval and juvenile stages compared to the terrestrial adult phase, reflecting adaptations to different predation risks.^[17] Complementing the physical spines, the newt secretes a poisonous milky mucus from granular skin glands, which coats the protruding ribs and enhances their deterrent effect by injecting irritants into attackers. These toxins, including steroidal compounds lethal to small mammals like mice when injected, cause pain and inflammation without intoxicating the newt itself, likely due to physiological resistance.^[19] The orange-colored warts serve as an aposematic signal, drawing predator attention to the toxin-laden areas, while the bright yellow ventral surface with dark spots functions as warning coloration, particularly visible when the newt arches its body in defense.^[17] In addition to active defenses, the newt exhibits escape behaviors such as rapid bursts of swimming in aquatic environments or burrowing into soft mud to evade pursuit on land.^[20]^[1] This integrated defense strategy is unique among salamanders, with the rib protrusion having no direct analogs in other species, and proves effective against avian predators like herons and storks, as well as mammalian ones such as otters, by redirecting attacks to protected wart regions and delivering toxins that reduce palatability. Experiments using plasticine models demonstrate that conspicuous warts attract over 60% of simulated attacks to those sites, minimizing damage to vital areas.^[17]^[1]^[21]

Diet and Predators

The Iberian ribbed newt (Pleurodeles waltl) is strictly carnivorous, preying on a variety of small aquatic organisms including insects such as beetle larvae and chironomid larvae, annelid worms, mollusks like gastropods, microcrustaceans (e.g., cladocerans, ostracods, and copepods), tadpoles, and occasionally small fish.^[1]^[16]^[22] Adults are opportunistic foragers that employ an ambush strategy in water, using a rapid suction-feeding mechanism to draw prey into the mouth via hyobranchial depression.^[23]^[24] Larval diet focuses on planktonic and benthic invertebrates, with microcrustaceans comprising the majority by number, supplemented by aquatic insect larvae such as those of culicids and chironomids; larvae filter or actively select available prey, adapting to rapid changes in pond succession during the flooding period.^[22]^[25] In the episodic Mediterranean ponds that characterize their range, feeding ecology reflects seasonal and habitat variability; during aquatic phases, the newts target abundant invertebrates.^[25]^[26] As mid-level predators in these aquatic food webs, Iberian ribbed newts regulate invertebrate populations, exerting top-down control that influences pond community dynamics, though they face low predation pressure in fishless temporary waters.^[22]^[25] Natural predators of the Iberian ribbed newt include birds such as herons and storks, mammals like the Eurasian otter (Lutra lutra), and reptiles including the viperine snake (Natrix maura), which target adults in aquatic and riparian zones.^[1]^[27]^[28] Fish, particularly invasive species like the largemouth bass (Micropterus salmoides), pose significant threats by preying on eggs and larvae in permanent waters, contributing to local population declines.^[2] These predatory pressures are mitigated by the newt's defensive adaptations, such as rib protrusion and skin toxins.^[1]

Reproduction

Breeding Cycle

The breeding season of the Iberian ribbed newt (Pleurodeles waltl) typically spans from winter to spring, varying by region but often occurring between November and May, particularly in areas like Extremadura where it aligns with October to May.^[1] This period is primarily triggered by increased rainfall and rising temperatures, which prompt adults to migrate to breeding sites such as temporary ponds and slow-moving streams.^[1] In northern regions like León, mating may be more concentrated from February to April, while southern populations can exhibit extended or biannual cycles including a summer peak.^[1]^[7] Mating occurs in shallow, vegetated waters where males develop nuptial pads on their forelimbs and a reddish dorsal hue to attract females. Males also produce sex pheromones encoded by Sodefrin Precursor-like Factor (SPF) genes, which function as whole proteins released during amplexus to facilitate mating.^[29] Courtship involves the male pursuing the female, rubbing her throat, and grasping her forelimbs in a prolonged axillary amplexus that can last from hours to several days.^[1]^[30] During this embrace, the male may perform circular swimming motions before releasing a spermatophore—a packet of sperm—onto the substrate, which the female picks up with her cloaca for internal fertilization.^[7]^[1] Females can store viable sperm in their spermatheca for up to five months, allowing delayed fertilization triggered by hormonal cues such as environmental changes.^[31] Following fertilization, females lay eggs in small clusters of 9 to 20, each surrounded by a gelatinous envelope approximately 5–7 mm in diameter, attaching them individually or in strands to submerged vegetation or other objects.^[1]^[30] A single female typically deposits 150 to 1,300 eggs over 2 to 3 days, with averages around 200–400 in natural conditions.^[1] Eggs hatch into larvae after 2 to 3 weeks, depending on temperature (e.g., 13 days at 18°C), emerging as 11 mm-long, limbless individuals with external gills.^[1] Larval development proceeds rapidly in warm waters, with front limbs appearing around two weeks and hind limbs by three weeks; metamorphosis, involving gill resorption and tail fin reduction, completes in 3 to 6 months (100–110 days at 18°C), resulting in juveniles 53–110 mm in snout-vent length.^[1]^[30] In permanent water bodies, some larvae exhibit paedomorphosis, retaining aquatic larval traits into adulthood rather than undergoing full metamorphosis.^[1]

Sex Determination

The Iberian ribbed newt (Pleurodeles waltl) exhibits a genetic sex determination system based on ZW sex chromosomes, where males possess the homogametic ZZ genotype and females the heterogametic ZW genotype, with the W chromosome carrying a dominant female-determining factor.^[32]^[33] This system aligns with female heterogamety observed in many urodele amphibians, where gonadal differentiation primarily occurs during the larval stage, influenced by genetic cues from the sex chromosomes.^[34] The ZW configuration ensures that ovarian development is triggered in genetic females, while testicular differentiation proceeds in ZZ individuals unless overridden by environmental factors.^[35] Environmental temperature plays a critical role in modulating this genetic determination, particularly during a thermosensitive period from developmental stages 42 to 54 in larvae. Exposure to high temperatures, specifically 32°C, during this window induces complete female-to-male sex reversal in genetic ZW individuals, resulting in functional neomales capable of producing viable sperm.^[32]^[35] This temperature-dependent override demonstrates the plasticity of the sex determination pathway, where heat inhibits the female-determining influence of the W chromosome and promotes testicular development.^[36] Conversely, lower temperatures reinforce the genetic sex, highlighting how P. waltl integrates environmental signals with chromosomal cues during early gonadogenesis.^[33] Hormonal mechanisms further underpin gonadal differentiation, with estrogens and androgens acting as key regulators. Estrogens, synthesized via aromatase activity, are essential for ovarian differentiation and can induce male-to-female sex reversal when administered to ZZ larvae, emphasizing their role in promoting female pathways.^[35] Androgens support testicular development but can also contribute to reversal if aromatized to estrogens, illustrating the interconnected steroid signaling in this species.^[37] This hormonal plasticity extends into adulthood, where rare spontaneous sex reversals have been documented under environmental stress, allowing limited gonadal reconfiguration in response to physiological challenges. Due to its ZW system combined with temperature and hormonal sensitivities, P. waltl serves as an influential model organism for investigating environmental influences on sex determination in amphibians, providing insights into evolutionary adaptations and plasticity in vertebrate reproduction.^[32]^[34] Studies on this species have elucidated mechanisms of thermosensitive reversal and steroid-mediated differentiation, contributing to broader understanding of how external factors can override genetic sex in ectothermic vertebrates.^[35]

Regeneration

Regenerative Capabilities

The Iberian ribbed newt (Pleurodeles waltl) exhibits remarkable regenerative abilities, capable of fully restoring complex structures including limbs, spinal cord, heart tissue, brain cells, the lens of the eye, and jaws following injury or amputation. Limb regeneration involves the complete regrowth of fore- and hindlimbs, typically completing in 1-2 months under standard laboratory conditions, resulting in functional appendages with proper patterning and no scarring. Spinal cord regeneration restores neural connectivity and function after transection, with ependymal cells proliferating to bridge the lesion and support axonal regrowth. Heart tissue regenerates after cryo-injury, with the damaged area fully recovering within approximately 8 weeks through cardiomyocyte proliferation and remodeling without fibrotic scarring. Brain regeneration includes the replacement of lost neurons and glia via activation of quiescent neural stem cells in the ependymal layer. The lens regenerates uniquely from dorsal iris pigment epithelial cells through transdifferentiation, forming a new, functional lens. Jaw regeneration rebuilds both upper and lower structures, including bone and associated tissues, maintaining craniofacial integrity. The regenerative process in P. waltl begins with wound healing and the formation of a blastema, a mass of undifferentiated progenitor cells derived from local dedifferentiation of multiple cell types near the injury site. This blastema proliferates and patterns the new tissue, guided by conserved signaling pathways such as Wnt/β-catenin, which promotes cell proliferation and proximal-distal outgrowth, and fibroblast growth factor (FGF) signaling, which supports blastema maintenance and mesenchymal condensation. Unlike mammalian wound healing, this process avoids scar formation entirely, enabling perfect structural and functional restoration across all regenerable tissues. Regenerative capacity varies by life stage, with larvae demonstrating faster regeneration rates compared to adults; for instance, larval lens regeneration initiates more rapidly post-lentectomy, though both stages achieve full lens replacement from iris-derived cells. This age-related difference highlights developmental plasticity, yet P. waltl retains robust regeneration throughout adulthood, unlike many vertebrates where abilities decline. In neural regeneration, P. waltl effectively restores brain architecture and spinal cord functionality through ependymoglial activation, similar to other urodeles such as the axolotl. The species' large 20 Gb genome, one of the largest among vertebrates, facilitates detailed molecular studies of these processes by enabling identification of regeneration-associated genes and transposable elements. A chromosome-scale assembly of the 20.3 Gb genome, published in 2025, has further advanced these investigations by improving annotation of repetitive elements and regulatory regions involved in regeneration.^[38]

Factors Influencing Regeneration

The regeneration process in the Iberian ribbed newt (Pleurodeles waltl) is modulated by various environmental factors that influence the rate and quality of tissue repair. Temperature plays a critical role, with regeneration typically studied at 18-22°C, as this range supports cellular proliferation and patterning in laboratory settings. Water quality affects overall health and regenerative efficiency, as contaminants like heavy metals (e.g., aluminum, chromium, cadmium) disrupt development in P. waltl. Nutrition is essential, with nutrient deprivation such as starvation allowing initial blastema formation but significantly retarding subsequent limb outgrowth and patterning due to limited energy for cell proliferation.^[39] Physiological variables further shape regenerative outcomes in P. waltl. Age impacts efficiency, with juveniles regenerating limbs faster and with fewer abnormalities than adults, where processes slow due to reduced proliferative capacity and increased fibrosis risk.^[40] Injury size influences regeneration, as larger wounds elicit a more robust blastema formation and gene expression response compared to smaller injuries, though excessive damage may impair efficiency.^[40] Hormonal levels, particularly thyroid hormones, play a role in metamorphosis and can reduce regenerative rate and fidelity post-metamorphosis in urodeles.^[41] Genetic and epigenetic mechanisms underpin variability in P. waltl regeneration. The gene Prod1, a cell-surface protein, is crucial for proximodistal limb patterning in urodeles, as its graded expression guides blastema cell sorting and outgrowth direction during regeneration.^[42] Epigenetic modifications, such as dynamic changes in histone methylation (e.g., H3K27me3), facilitate dedifferentiation by altering chromatin accessibility and enabling progenitor gene activation in regenerating tissues. Experimental interventions reveal additional modulators. X-irradiation inhibits regeneration in P. waltl by damaging DNA in blastema precursor cells, preventing dedifferentiation and blastema formation even at localized doses.^[43] Retinoic acid exposure can alter regenerative potential in newts by proximalizing blastema positional identity, often inducing supernumerary limb structures.

Scientific Research

Space Experiments

The Iberian ribbed newt (Pleurodeles waltl) has served as a key model organism in space biology since the 1980s, with experiments focusing on the impacts of microgravity on regeneration, development, and physiological adaptations. These studies, conducted primarily on Russian biosatellites and the Mir space station, have utilized both larval and adult specimens to investigate how weightlessness influences processes like tissue repair and reproduction. Ground-based controls were consistently employed to compare outcomes, with post-mission analyses involving histological examinations of tissues to assess cellular proliferation, structural changes, and functional integrity.^[44] The inaugural mission occurred aboard Bion 7 in 1985, where ten adult newts were exposed to microgravity for seven days to observe initial effects on limb regeneration following amputation. This experiment marked the first use of P. waltl in space and provided baseline data on unaltered development under orbital conditions. Subsequent flights built on these foundations: Bion 10 in 1992-1993 examined tail regeneration, including spinal cord, muscle, and bone repair in adults, revealing altered regenerative patterns such as modified neural reconnection and tissue differentiation compared to ground controls. Bion 11 in 1996 targeted neural retina regeneration after optic nerve lesioning, demonstrating enhanced cell proliferation rates in early regenerative stages.^[45]^[46]^[44] Longer-duration studies on the Mir space station from 1996 to 1999, including the FERTILE I, FERTILE II, and NEUROGENESIS missions, exposed larvae and adults to microgravity for periods up to five months, investigating embryonic development, fertilization, and neurogenesis. These experiments confirmed that fertilization rates remained comparable to terrestrial conditions, with post-flight adults exhibiting intact fertility and producing viable offspring without genetic abnormalities. The Foton-M2 mission in 2005 further explored hematopoietic tissue and stem cell responses in intact and operated newts over 16 days, highlighting subtle shifts in proliferative activity without severe disruptions.^[47]^[31]^[48] A consistent finding across missions was accelerated tissue regeneration in microgravity, with cell proliferation increasing 1.5- to 2-fold during early stages of lens, retina, limb, and tail repair, attributed to reduced gravitational constraints on cellular migration and dedifferentiation. Spinal cord regeneration showed distinct alterations, including enhanced ependymal cell activity but delayed functional integration post-flight. Vestibular adaptations were evident in larvae, with modified sensorimotor development and balance recovery upon re-entry, indicating gravity's role in neural circuit formation. No teratogenic effects were observed in developing embryos, as cleavage, gastrulation, and organogenesis proceeded normally, providing insights into gravity-independent developmental thresholds. These outcomes underscore microgravity's promotion of regenerative processes while revealing its modulatory effects on neural and reproductive systems.^[44]^[45]^[49]^[47]

Genomic and Biomedical Studies

The genome of the Iberian ribbed newt (Pleurodeles waltl) was sequenced at chromosome-scale resolution in 2025, resulting in a high-quality assembly of approximately 20.3 Gb, which encompasses 18,799 protein-coding genes.^[38] This assembly highlights the species' large genome size—six times that of humans—dominated by repetitive elements that contribute to its regenerative capabilities, including insights into non-coding RNA landscapes active during limb regeneration, facilitating the identification of genes involved in tissue repair and limb regeneration.^[50] The sequencing effort, published in Cell Genomics, provides a foundational resource for comparative genomics and evolutionary studies of urodele amphibians.^[51] Husbandry protocols for captive breeding and laboratory maintenance of P. waltl emphasize optimal environmental conditions to support its biphasic life cycle, with temperatures maintained at 18–22°C to mimic natural habitats and promote health.^[52] A varied diet consisting of gut-loaded insects, earthworms, and commercial amphibian feeds is recommended to meet nutritional needs across larval, juvenile, and adult stages.^[53] Recent 2025 updates detail life cycle phases, including metamorphosis and breeding cues, enabling efficient long-term colony management in research settings.^[53] As a model organism, P. waltl is widely used in biomedical research for regenerative medicine, particularly in studying limb, heart, and neural tissue repair due to its robust regenerative abilities.^[29] It serves as a platform for investigating neurogenesis, with studies demonstrating ongoing neural stem cell activity in the adult brain and spinal cord, offering insights into vertebrate brain plasticity.^[54] Additionally, the species has been employed in toxicity testing, including genotoxicity assessments of chemicals like cadmium on development.^[55] A 2025 study in Current Biology explores its phenotypic plasticity, highlighting adaptations between aquatic and terrestrial lifestyles that inform broader applications in developmental biology.^[56] Emerging research utilizes CRISPR-Cas9 editing in P. waltl to target regeneration pathways, such as Hox genes and Yap signaling, revealing mechanisms of genetic compensation that maintain regenerative proficiency despite mutations.^[57]^[58] These efforts enable direct comparisons to human tissue repair processes, underscoring conserved genes that could advance therapies for injury recovery and degenerative diseases.^[59]

Conservation

Status and Threats

The Iberian ribbed newt (Pleurodeles waltl) is classified as Least Concern by the IUCN Red List, based on a 2020 global assessment.^[60] This status upgrade from Near Threatened (recorded in 2009) reflects a generally stable global population, though localized declines persist in parts of its range.^[60]^[1] However, the 2025 European Red List assessment classifies the species as Near Threatened (NT A2c) for its European populations.^[61] Fragmentation is particularly evident in Moroccan subpopulations where suitable habitats are isolated.^[60] Primary threats include habitat destruction from urbanization and agricultural activities, which eliminate or degrade temporary ponds essential for breeding.^[60]^[1] Water pollution, particularly from pesticides and heavy metals, contaminates breeding waters and reduces larval survival rates.^[60]^[1] Introduced invasive species exacerbate risks, as the largemouth bass (Micropterus salmoides) and red swamp crayfish (Procambarus clarkii) actively prey on eggs and larvae, leading to recruitment failures in affected sites.^[60]^[11]^[27] Climate change compounds these pressures through intensified droughts that diminish ephemeral breeding habitats.^[60]^[62] Temperature increases may further disrupt population dynamics by skewing sex ratios, given the species' reliance on temperature-dependent sex determination and phenotypic plasticity.^[60]^[63]

Conservation Measures

The Iberian ribbed newt (Pleurodeles waltl) benefits from several legal protections across its range. In Spain, the species is safeguarded by national legislation, including royal decrees that prohibit collection and habitat disturbance, ensuring its inclusion in protected areas such as national parks and Natura 2000 sites.^[1] It is listed under Appendix III of the Bern Convention, which promotes cooperation for monitoring and trade regulation among signatory states, particularly relevant for transboundary populations in Iberia and North Africa.^[64] Additionally, while not explicitly in the EU Habitats Directive annexes for strict protection, it receives indirect safeguards through Annex IV provisions in Spain and Portugal via habitat management requirements in designated wetlands.^[64] In Morocco, populations are conserved within protected areas like Parc National de Souss-Massa, where habitat preservation limits exploitation.^[64] Conservation initiatives focus on habitat enhancement and population reinforcement. Habitat restoration projects, such as those under the EU LIFE program (LIFE05 NAT/E/000060), involve creating artificial ponds and restoring degraded wetlands in eastern Spain, leading to rapid colonization by P. waltl and other amphibians.^[65] In regions like Valencia, efforts include controlling invasive species, such as red swamp crayfish (Procambarus clarkii), which prey on eggs and larvae, through targeted removal to protect breeding sites.^[1] Captive breeding programs have been successful, with facilities in Spain producing individuals for reintroduction into restored sites, as demonstrated in Valencian Community projects that aim to bolster fragmented populations.^[15] These initiatives emphasize maintaining aquatic habitats amid agricultural pressures. Monitoring efforts include regular amphibian surveys across Iberia to track population trends and habitat use. In eastern Spain, multi-year pond monitoring has documented P. waltl abundances and interactions with co-occurring species, informing adaptive management.^[66] Genetic studies, leveraging the species' 2017 genome assembly and 2019 transcriptome resources, assess diversity in Iberian populations to guide reintroduction efforts and prevent inbreeding in isolated groups.^[14] Spanish and Portuguese authorities conduct ongoing surveys, highlighting the need for continued vigilance in northern ranges.^[2] Future strategies prioritize climate adaptation and awareness. Projects like ADAPTECCA in Spain focus on creating resilient habitats, such as elevated ponds to counter drought and flooding from climate change, ensuring long-term breeding site availability.^[67] Public education campaigns, integrated into LIFE Renaturwat initiatives, promote pollution reduction through community involvement in wetland cleanups and agrochemical awareness, fostering sustainable land use in rural Iberia.^[68] These approaches aim to mitigate ongoing habitat fragmentation while building resilience against emerging threats.^[69]

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[11]
(PDF) Pleurodeles waltl - ResearchGate
... Pleurodeles waltl. The IUCN Red List of Threatened. Species 2009: e ... status. Away from Djebel Edough, P. poireti consists of three distinct clades ...
[12]
Integrative inference of population history in the Ibero-Maghrebian ...
In this study, we combine SDMs and molecular data to identify glacial refugia and reconstruct the population history of the Iberian ribbed newt Pleurodeles ...
[13]
[PDF] AMPHIBIANS OF NORTHWEST AFRICA - Smithsonian Institution
Position of palatal teeth and choanae in Pleurodeles poireti (a) and Pleurodeles waltl (b). ... wells and cisterns. Active by day and night. They feed on.
[14]
[PDF] Pond area and availability safeguard amphibian genetic diversity ...
Apr 15, 2024 · Owing to their preference for temporary ponds with long hydro- period or permanent aquatic habitats, P. perezi and P. waltl show a high degree ...
[15]
[PDF] Pleurodeles waltl
Habitat and Ecology Information: This is a highly aquatic species of Mediterranean type habitats including scrub, open wood and cultivated land.
[16]
Abstract - Wiley Online Library
Dec 29, 2022 · In this article, I explored the relationships between different antipredator defences in the Iberian ribbed newt, Pleurodeles waltl. This ...
[17]
Salamander Pleurodeles waltl (Salamandridae) - jstor
NOWAK AND EDMUND D. BRODIE, JR. The sharp rib tips of Pleurodeles waltl penetrate the lateral warts which do not contain granular glands.
[18]
Hurt yourself to hurt your enemy: new insights on the function of the bizarre antipredator mechanism in the salamandrid Pleurodeles waltl
**Summary of Rib Defense Mechanism in Pleurodeles waltl**
[19]
Defense Mechanisms of the Spanish Ribbed Newt - Caudata.org
In a study published in 2009, Egon Heiss and colleagues observed and documented the anatomical adaptations that allow rib protrusion in the Spanish ribbed newt.
[20]
Close-up of ventral view of some predated Pleurodeles waltl (© R....
We report an event of massive predation of ribbed newts ( Pleurodeles waltl ) and Iberian spadefoot toads ( Pelobates cultripes ) by a pair of otters ( Lutra ...
[21]
[PDF] Diet of larval Pleurodeles waltl (Urodela
Oct 8, 2019 · We first assessed the extent to which the food items found represent the entire dietary range of the larvae of P. waltl. To do this we ...<|separator|>
[22]
Functional trade-offs in the aquatic feeding performance of ...
The purpose of this research was to quantify the diverse morphology and suction feeding performance of aquatically feeding salamandrids. We hypothesized that a ...Missing: diet | Show results with:diet
[23]
Spanish Ribbed Newt - Reptilia Zoo
Scientific Name: Pleurodeles waltl. Names: N/A. Locations: Whitby, Vaughan ... Average lifespan. In the wild, about 7–10 years; in captivity, 8–12 years ...<|separator|>
[24]
Pleurodeles waltl Newt Larvae Diet Regarding Available Prey
Dec 18, 2012 · We studied the diet of larvae of Pleurodeles waltl when exogenous feeding begins and throughout the flooding period in an episodic Mediterranean ...
[25]
Functional morphology of terrestrial prey capture in salamandrid ...
Nov 1, 2017 · Representative images of the cleared and stained hyobranchial apparatus of Chioglossa lusitanica, Salamandra salamandra, Pleurodeles waltl, ...
[26]
Species page of Pleurodeles waltl - Moroccoherps
Sep 23, 2012 · It is mainly abundant in stagnant places but less abundant in low current both rivers and streams. It can get used to live in high anthropic- ...Missing: seasonal overwinter
[27]
(PDF) Impact of otter (Lutra lutra) predation on amphibians in ...
Aug 10, 2025 · The high number of dead amphibians killed by two otters in only several hours suggests that the presence of these predators may pose a serious ...
[28]
Breed Spanish Ribbed Newts - Reptiles Magazine
### Summary of Breeding Spanish Ribbed Newts (Captive Conditions)
[29]
Pleurodeles waltl, amphibian, Urodele, is a suitable biological ...
Pleurodeles waltl (amphibian, Urodele) is an appropriate biological model for space experiments on a vertebrate. One reason for interest in this animal ...Missing: etymology Michael
[30]
Sexual Development of the Urodele Amphibian Pleurodeles waltl
Jun 20, 2008 · Pleurodeles waltl is a urodele amphibian that displays a ZZ/ZW genetic mode of sex determination involving a putative W-borne dominant ...
[31]
Female-enriched expression of ERα during gonad differentiation of ...
This member of the salamandridae family shows a ZZ/ZW genetic mode of sex determination in which the W chromosome induces a female differentiation. The gonadal ...
[32]
Lifelong testicular differentiation in Pleurodeles waltl (Amphibia ...
Mar 5, 2009 · The Iberian ribbed newt Pleurodeles waltl has been studied extensively as a model for sex determination and differentiation. However, the ...
[33]
Steroids, aromatase and sex differentiation of the newt Pleurodeles ...
Female to male sex reversal occurs when ZW larvae are reared at 32 degrees C during a thermosensitive period (TSP) that takes place from stage 42 to stage 54 of ...
[34]
Lifelong testicular differentiation in Pleurodeles waltl(Amphibia ...
Mar 5, 2009 · The thermosensitive period is the developmental window during which genetically female larvae (ZW) have to be reared at 32°C to obtain sex ...
[35]
Steroids, aromatase and sex differentiation of the newt Pleurodeles ...
In the newt Pleurodeles waltl, genetic sex determination obeys female heterogamety (female ZW, male ZZ). In this species as in most of non-mammalian ...
[36]
Temperature preference during forelimb regeneration in the red ...
Environmental temperatures, however, can increase or decrease the red-spotted newt's metabolic processes, including their rate of tissue regeneration; whether ...
[37]
Bioassays for testing effects of Al, Cr and Cd using development in ...
Nov 7, 2008 · Bioassays for testing effects of Al, Cr and Cd using development in the amphibian Pleurodeles waltl and regeneration in the planarian Dugesia ...
[38]
Altered metabolic state impedes limb regeneration in salamanders
Our results demonstrate that under adverse conditions, salamanders can exploit alternative strategies to secure blastema formation for limb regeneration.
[39]
Hallmarks of regeneration - ScienceDirect.com
Sep 5, 2024 · Stressful conditions affect hair follicles through increased B adrenergic signaling, driving hyperproliferation, depletion of melanocyte ...
[40]
Mechanisms of urodele limb regeneration - PMC - PubMed Central
Effects of localized X‐irradiation on regeneration of the urodele limb. ... The inhibition of limb regeneration in urodele larvae by localized irradiation ...
[41]
A cross-species analysis of systemic mediators of repair and ...
Apr 1, 2021 · Thyroid hormone has been implicated in the regeneration of a variety of physiological structures, including appendages, heart, and nerves. The ...
[42]
https://evodevojournal.biomedcentral.com/articles/10.1186/s13227-015-0006-6
[43]
Changes in global histone modifications during dedifferentiation in ...
Sep 16, 2010 · Histone modifications are dynamically changed during dedifferentiation of PECs. A coordination of gene activation-related modifications, ...Missing: epigenetic | Show results with:epigenetic
[44]
https://pubmed.ncbi.nlm.nih.gov/12528722/
[45]
https://www.sciencedirect.com/science/article/pii/S0273117702003927
[46]
effects upon organ and tissue regeneration - PubMed
The most recent experiment flew aboard the Bion-11 biosatellite. This test was performed for study on microgravity effect on neural retina regeneration ...
[47]
Urodelean amphibians in studies on microgravity - ScienceDirect.com
The most recent experiment flew aboard the Bion-11 biosatellite. This test was performed for study on microgravity effect on neural retina regeneration ...
[48]
Tissue regeneration in space (spinal cord, muscle and bone) in the ...
... Pleurodeles (TJrodeles amphibians) sent aboard the Russian satellite Bion 10. After landing, regeneration blastemas were removed and subjected to ...
[49]
Microgravity and hypergravity effects on fertilization of the ... - PubMed
Effects of microgravity (microG) on fertilization were studied in the urodele amphibian Pleurodeles waltl on board the MIR space station.Missing: teratogenic | Show results with:teratogenic
[50]
Studies on hemopoietic tissue of ribbed newt, Pleurodeles waltl after ...
The effect of 16-day spaceflight aboard the Foton-M2 satellite on the hematopoietic tissue of P waltl newts was studied in flown intact animals F-int and in ...
[51]
Development of neuronal and sensorimotor systems in the absence ...
Neurobiological experiments on 4 animal species (Xenopus laevis, Pleurodeles waltl, Drosophila melanogaster, Acheta domesticus) were performed to study effects ...
[52]
Chromosome-scale genome assembly reveals how repeat elements ...
Jan 27, 2025 · Among DNA repeat elements, hATs have a significant expression in the early phases of limb regeneration in P. waltl (Figure S15). To further ...
[53]
Chromosome-scale genome assembly reveals how repeat elements ...
Jan 27, 2025 · We present here the chromosome-scale as- sembly of the 20.3 Gb genome of the Iberian ribbed newt, Pleurodeles waltl, with a hitherto ...
[54]
Chromosome-scale genome assembly reveals how repeat elements ...
We present here the chromosome-scale assembly of the 20.3 Gb genome of the Iberian ribbed newt, Pleurodeles waltl, with a hitherto unprecedented ...
[55]
Keeping the Spanish Ribbed Newt - Vet Verified
Aug 14, 2025 · This guide provides a detailed overview of the Spanish ribbed newt's care, including housing, feeding, health, and breeding. ... August 16, 2025.Missing: husbandry protocols
[56]
Husbandry of Spanish Ribbed Newts (Pleurodeles waltl)
Aug 6, 2025 · Here, we show that the Iberian ribbed newt (Pleurodeles waltl) has outstanding properties as a laboratory newt. We developed conditions ...
[57]
The inbred newt genome unveils molecular mechanisms of behavior ...
Oct 17, 2025 · waltl after divergence from the common ancestor of Salamandridae 56 million years ago. These findings raise intriguing questions about the ...
[58]
Establishment of neural stem cell culture from the central nervous ...
We report the formation of spherical cell aggregates similar to neurospheres in vitro from the brain and spinal cord of post‐metamorphic Pleurodeles waltl.
[59]
Endocrine-disrupting effects and reproductive toxicity of low dose ...
Flament et al. Effect of cadmium on gonadogenesis and metamorphosis in Pleurodeles waltl (urodele amphibian) ... Cardiotoxicity in African clawed frog ...
[60]
Iberian ribbed newts: Current Biology - Cell Press
Jan 20, 2025 · And like all newts, the Iberian ribbed newt (Pleurodeles waltl) belongs to a subfamily of salamanders known as Pleurodelinae (Figure 1A).
[61]
Multiple gene knockouts in newts reveal novel functions of 5' Hox ...
Therefore, we generated 5' Hox knockout newts (Pleurodeles waltl) using CRISPR-Cas9. Results: Individual knockouts of Hox9, Hox10, and Hox12 ...
[62]
Genetic compensation response contributes to Pleurodeles waltl ...
May 31, 2025 · We found that in the highly regenerative salamanders, Yap knockout does not compromise the limb regeneration due to genetic compensation response (GCR).
[63]
Reading and editing the Pleurodeles waltl genome reveals novel ...
Dec 22, 2017 · Here, we present the ~20 Gb genome and transcriptome of the Iberian ribbed newt Pleurodeles waltl, a tractable species suitable for laboratory research.<|control11|><|separator|>
[64]
The IUCN Red List of Threatened Species
Insufficient relevant content. The provided URL (https://www.iucnredlist.org/species/59463/11926338) does not contain detailed information on geographic range, distribution, population trends, or historical aspects for *Pleurodeles waltl*. The page appears to be a placeholder or lacks the specific data requested. No citations or exact numbers are available from the content provided.
[65]
Spanish ribbed newt facts, distribution & population - BioDB
This amphibian can be found across a wide geographical range, including Portugal, southern and central Spain, and Morocco.
[66]
Effects of global warming on species with temperature‐dependent ...
As global warming continues, the TSD mechanism could lead to heavily skewed offspring sex ratios towards one sex, which in turn threatens populations' ...Missing: drought | Show results with:drought
[67]
[PDF] Near Threatened Amphibian Species
It lives in lowland moist forest and in degraded habitats near more mature forest and can be found in small groups along forest watercourses, often in ...
[68]
Restoration of priority habitats for amphibians - European Union
Outcomes were effective and resulted in almost immediate colonisation of the ponds by amphibians, especially Pleurodeles waltl, Pelodytes punctatus, Alytes ...
[69]
Are Iberian ribbed newts Pleurodeles waltl Michahelles, 1830 ...
Jan 20, 2021 · The newt Pleurodeles waltl is commonly found in Mediterranean ponds in the Iberian Peninsula, where it preys on a wide variety of organisms. We ...
[70]
Adaptation, restoration and creation of habitats for amphibians ...
This project consists in an intervention for the conservation and improvement of the populations of the eleven species of amphibians inhabiting the Montes de ...
[71]
LIFE Renaturwat project celebrates World Water Day ... - Noticia UPV
"The reintroduction of Pleurodeles waltl is an important step in conservation efforts in the area and demonstrates the commitment of the LIFE project ...
[72]
Effective conservation planning of Iberian amphibians based on a ...
Nov 1, 2022 · Amphibians are severely affected by climate change, particularly in regions where droughts prevail and water availability is scarce.