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Uria is a of seabirds in the family Alcidae (/ˈɔːkɪdiː/), commonly known as murres or guillemots, comprising two extant species: the (Uria aalge) and the (Uria lomvia). These are medium- to large-sized auks, typically 38–50 cm in length with a of about 61–73 cm, featuring sleek black-and-white plumage during the breeding season—dark above and white below—and a more mottled appearance in winter. Adapted for a pelagic lifestyle, members of the genus are powerful swimmers and divers, using their wings as flippers to pursue , squid, and crustaceans at depths exceeding 100 m, sometimes reaching over 200 m. The genus Uria has a distribution in the , breeding in massive colonies—often numbering hundreds of thousands to over a million pairs—on steep cliffs and rocky ledges along and coasts of the Atlantic and Pacific Oceans, from and to and . Outside the breeding season, they migrate southward to continental shelf waters, with the extending to temperate latitudes as far south as and . These birds do not build nests, instead laying a single, pyriform directly on bare rock to prevent rolling; the egg's pointed shape is a key for this precarious habitat. Chicks, known as "penguin stage" due to their precocial but flightless departure from the at 2–4 weeks old, jump from cliffs into the and complete development while being cared for by the male parent during a post-fledging dispersal period. Uria species are among the most abundant seabirds globally, with the alone supporting over 8 million breeding pairs, though populations face threats from oil spills, , , and historic hunting. Fossil records indicate the genus originated in the , with a rich history of adaptation to marine environments, making it a key component of northern ecosystems where it serves as both predator and prey. The two are distinguished primarily by bill shape—the having a slender, pointed bill and the a stouter, more decurved one—reflecting subtle differences in diet and depth.

Taxonomy and systematics

Etymology and classification history

The genus Uria was introduced in 1760 by the French zoologist in the first volume of his Ornithologie, ou, Méthode contenant la division des oiseaux en ordres, sections, genres, especes & leurs variétés, with Uria aalge (the ) established as the by subsequent tautonymy. Brisson's classification system, which followed Carl Linnaeus's introduced in 1758, grouped Uria among based on morphological similarities such as bill shape and foot structure, though his work used a quasi-binomial format. The name Uria derives from the term ouria (οὐρία), denoting an unspecified waterfowl or , as referenced by the ancient writer in his . initially incorporated species now assigned to Uria into the broader Alca Linnaeus, 1758 (encompassing the and ), reflecting a less refined separation of auk species in the 10th edition of Systema Naturae (1758) and later editions. Brisson's establishment of Uria thus represented an early effort to distinguish murre-like auks from other members of Alca, setting the stage for more precise generic boundaries. In the early 19th century, the Uria was formally placed within the family Alcidae, erected by British zoologist in 1820 to unite wing-propelled diving birds of the , including related genera such as Alca () and Alle (). This familial grouping, detailed in Leach's Systematic Catalogue of the Specimens of the Indigenous Mammalia and Birds of and subsequent works, emphasized shared adaptations like compact bodies and short wings for underwater propulsion. 19th-century ornithologists, including in his (1827–1838) and Elliott Coues in Key to North American Birds (1872), further solidified Uria's classification by documenting its distinction from Alca through and geographic distribution, while recognizing two extant within the genus.

Extant species

The genus Uria includes two extant species of seabirds in the auk family Alcidae: the (Uria aalge) and the (Uria lomvia). These species are closely related, sharing similar overall body plans but differing in key morphological traits and geographic preferences. The (Uria aalge) is divided into five recognized subspecies, reflecting adaptations to its extensive breeding range across low-Arctic and boreal waters from through the Pacific to and the North Atlantic. These subspecies are U. a. aalge, which occurs along the North Atlantic coasts from to , , and the ; U. a. albionis, restricted to the ; U. a. ibericus, found on the and northwest ; U. a. hyperborea, breeding from to , , and northern ; and U. a. inornata, distributed across the North Pacific from the and to the , Kamchatka, the , and . A primary distinguishing feature of the species is its slender, pointed bill that tapers gradually to the tip. The (Uria lomvia) comprises four , primarily occupying high waters in a circumpolar distribution across , , and . The include U. l. lomvia, breeding from northeast to in the Atlantic sector of the ; U. l. eleonorae, in the eastern to the ; U. l. heckeri, on ; and U. l. arra, in the and of the Pacific sector. This species is characterized by a thicker, shorter bill with a hooked tip and more abrupt taper compared to the . It favors higher latitudes than the , with breeding concentrated in colder environments. Comparatively, the two species differ notably in bill morphology, with the common murre's slender bill suited to its broader latitudinal range in subarctic to temperate zones, while the thick-billed murre's robust, hooked bill aligns with its specialization in high-Arctic conditions. These distinctions aid in field identification and reflect ecological partitioning within the genus.

Phylogenetic relationships and fossil record

The genus Uria occupies a basal position within the Alcidae subfamily Alcinae, forming a monophyletic with the closely related genera Alca (), Alle (), and the extinct Pinguinus (), based on comprehensive molecular analyses of mitochondrial and nuclear DNA sequences from all extant alcid . This fish-eating lineage is sister to a planktivorous comprising genera such as Fratercula (puffins) and Aethia (auklets), with the overall Alcidae radiating in the Early Eocene approximately 53 million years ago. More recent analyses estimate the divergence of Alcidae from its (Stercorariidae) around 35 million years ago in the late Eocene. Molecular clock estimates indicate that Uria and its relatives (Alca, Alle, and Pinguinus) form a that diverged from other alcids in the , with further diversification in the , calibrated using constraints. The fossil record of Uria documents early diversification within subtropical regions of the North Pacific, predating the genus's modern and boreal distributions. The earliest known species, Uria brodkorbi, was described from a partial preserved as impressions in diatomite from the (approximately 11–7 million years ago) Monterey Formation in , featuring a comparable in size to those of extant Uria species and exhibiting a depressed, ovoid pectoral crest for enhanced muscle attachment. This fossil suggests that Uria originated in warmer coastal environments before migrating northward, consistent with broader Alcidae patterns of radiation along Pacific margins. Additional Uria fossils, such as Uria affinis from the , further indicate persistence and adaptation in North American waters through the . Evolutionary adaptations in Uria for wing-propelled diving are evident in humeri, which show dorsoventral compression of the shaft and reinforced proximal ends to withstand hydrodynamic stresses during , traits shared with other Alcidae and calibrated to early divergences around 34 million years ago. Hypotheses for the genus's origin propose an initial Pacific diversification during the , potentially facilitated by tectonic changes and ocean current shifts that enabled northward dispersal, though direct evidence for connections via the proto-Panama remains limited to broader charadriiform patterns. These two extant represent the modern endpoints of this lineage.

Physical characteristics

Morphology and size

Species of the genus Uria exhibit a streamlined body form optimized for aquatic locomotion, with a length ranging from 38 to 48 , a of 61 to 81 , and a body mass of 736 to 1481 g. in size is minimal, with males averaging slightly larger than females in most linear measurements but showing no significant differences in overall proportions. The body structure features short, robust wings adapted for propulsion during underwater dives, functioning like flippers to enable efficient swimming at depths exceeding 100 m. Legs are positioned far posteriorly, providing leverage for powerful kicks in water while contributing to an upright, penguin-like stance on that facilitates balance during brief terrestrial movements. Among anatomical adaptations, Uria possess enlarged supraorbital salt glands that excrete excess from ingested and marine prey, preventing osmotic imbalance in their saline environment. Additionally, their strong, curved toenails aid in gripping steep cliff faces during nesting and colony navigation. Comparative morphology between the two extant reveals subtle differences: U. aalge () is generally slightly smaller, with a mean length of 38–43 cm and mass of 800–1125 g, whereas U. lomvia () has a stockier build, averaging 40–48 cm in length and up to 1481 g in mass, accompanied by a proportionally heavier bill. These variations support distinct ecological roles without altering the core diving-adapted physique shared across the .

Plumage, coloration, and variation

Species of the genus Uria, including the (Uria aalge) and (Uria lomvia), exhibit distinct seasonal s adapted to their marine environments, with adults undergoing two complete molts annually to transition between breeding and winter appearances. In breeding , both display blackish-brown upperparts, including the head, back, and wings, contrasting sharply with white underparts. U. aalge typically features a uniformly dark head in non-bridled individuals, while U. lomvia has a black head with a thin white streak extending from the base of the bill rearward and upward toward the eye. During winter, the of Uria becomes greyer on the upperparts, with the white underparts retained but a partial white throat and cheeks appearing, along with a dark streak or extending from the eye rearward. This non-breeding aids in during pelagic life. Adults of both undergo a complete prebasic molt post-breeding in late summer, rendering them flightless for 4–6 weeks as are replaced, followed by a prealternate molt in late winter to regain breeding coloration. Plumage variation occurs across and geographically within Uria. In U. aalge, the bridled form—characterized by prominent arcs and a white line extending rearward from the eye, resembling spectacles—appears in up to 70% of adults in northern Atlantic populations, with frequency increasing latitudinally from near 0% in southern regions; for example, the U. a. albionensis in the shows bridled traits in a notable proportion of adults. Overall, Pacific populations of U. aalge exhibit blacker upperparts compared to the browner tones in Atlantic forms, reflecting a west-to-east in coloration intensity. Similarly, U. lomvia shows subtle clinal variation, with North Pacific individuals displaying more intensely blackish than their Atlantic counterparts. Eggs of Uria species are uniquely pyriform (conical or pear-shaped), laid singly on bare cliff ledges without constructed nests, and feature highly variable coloration for camouflage and individual recognition amid dense colonies. Ground colors range from white and pale brown to or dark green, overlaid with diverse mottled patterns such as speckles, blotches, or streaks in browns, blacks, or reds, ensuring each egg's distinct appearance to prevent misplacement by parents.

Distribution and habitat

Geographic range

The genus Uria is distributed circumpolarly across the , with its two extant occupying distinct but partially overlapping ranges in marine environments. The (Uria aalge) has a broad distribution spanning to temperate zones, breeding along northern coasts from southward extensions in the eastern Pacific to northern in the western Pacific, and northward into the including , the , and . In the Atlantic, its range extends from western and southward to the , with small, isolated breeding populations reaching the . The (Uria lomvia), by contrast, is a high specialist with a more northerly and restricted range, breeding primarily in and waters from 46° to 82°N in the Atlantic and Oceans, and 50° to 72°N in the Pacific. Its distribution includes coastal and island sites from and eastward through , , and to and the , largely avoiding southern temperate latitudes below 50°N. Overlap between the two species occurs in shared breeding colonies, notably in the Bering Sea—where both nest on cliffs in the eastern Pacific—and in the North Atlantic, such as at sites along the Labrador coast and Newfoundland, facilitating potential interspecific interactions. These zones represent areas of sympatry within the broader circumpolar framework. Following the Pleistocene glaciations, both Uria species expanded their ranges northward, colonizing deglaciated northern coasts as ice sheets retreated around 10,000–12,000 years ago. Population genetic analyses of U. aalge reveal patterns consistent with post-glacial recolonization from southern refugia, leading to current distributions in the Atlantic and Pacific. For U. lomvia, evidence from colony sites in Hudson Strait and northern Hudson Bay indicates rapid post-glacial establishment in high Arctic breeding grounds previously covered by ice.

Habitat preferences and requirements

Uria , comprising the (Uria aalge) and (U. lomvia), exhibit specialized preferences shaped by their need for secure breeding sites and productive grounds in cold marine environments. For breeding, both favor steep sea cliffs, rocky islands, and offshore stacks in and regions, where they form dense colonies on narrow ledges or flat surfaces, often nesting shoulder-to-shoulder without constructing nests. These sites provide essential protection from mammalian predators such as foxes and mustelids by elevating nests above ground level, while allowing easy access to surrounding waters. Colony densities can reach extraordinary levels, with some sites supporting over 1 million individuals, facilitating social benefits like enhanced vigilance against aerial threats. Foraging habitats are centered in pelagic zones over continental shelves, particularly in areas of that concentrate prey such as small fish and invertebrates. Both species avoid warm tropical waters, restricting their range to cooler marine systems where sea surface temperatures typically remain below 15°C, as higher temperatures correlate with reduced prey availability and physiological stress. Access to reliable within 60–70 km of breeding colonies is critical, enabling efficient provisioning during the energetically demanding breeding season. Habitat preferences differ between the two species, reflecting their distributional niches. The (U. lomvia) shows a stronger affinity for ice-edge habitats in high- waters, where it often rests on floes and forages in adjacent pack ice zones during non-breeding periods. In contrast, the (U. aalge) prefers more temperate coastal environments, tending to avoid extensive pack ice and concentrating in shelf waters closer to shore. These distinctions allow partial in Pacific arctic colonies while minimizing competition in the Atlantic.

Ecology and behavior

Diet and foraging strategies

Uria species are piscivorous seabirds that primarily consume small schooling fish such as capelin (Mallotus villosus) and sand eels (Ammodytes spp.), along with crustaceans including amphipods and euphausiids (Thysanoessa spp.), and occasionally mollusks and squid. Their diet exhibits seasonal shifts, with a greater reliance on zooplankton like large copepods during winter when fish availability decreases. These prey items are typically micronektonic, ranging from 2–25 cm in length, and are selected for their abundance in pelagic waters near breeding colonies. Foraging strategies involve pursuit diving, where individuals propel themselves underwater using wings in a flight-like motion to chase prey. Dives commonly exceed 100 m, with U. lomvia capable of reaching up to 200 m and lasting over 3 minutes. Birds often forage in large flocks, including multispecies groups, to exploit concentrated prey schools, enhancing efficiency in open marine environments. Daily food intake for adults ranges from 200–500 g, equivalent to 20–50% of body weight, to meet high energetic demands; prey is stored in the bill or before consumption or regurgitation for transport. Differences between species reflect : U. lomvia favors Arctic prey like Arctic cod (Boreogadus saida), comprising up to 99% of diet mass in some regions, while U. aalge targets a broader array of temperate schooling .

Reproduction and breeding biology

Uria , including the (Uria aalge) and (Uria lomvia), exhibit highly synchronized breeding seasons that vary by latitude, typically spanning May to July in regions and commencing earlier in southern populations where retreats sooner. These seabirds form socially monogamous pairs that often reunite annually and defend small egg-laying sites within dense colonies. Breeding occurs in large colonies on steep cliff ledges or slopes, where pairs lay eggs directly on bare rock without constructing nests, relying on the colony's collective density for protection. Each pair produces a single large, pyriform (pear-shaped) egg per breeding season, with the conical form minimizing rolling risk on narrow ledges and facilitating a tight circular path if displaced. Eggs exhibit polymorphic coloration and patterning, ranging from white to blue-green with spots or streaks, which provides disruptive camouflage against the rocky substrates and enhances survival by reducing detectability to avian predators. Both parents share incubation duties, which last 30–35 days, with shifts typically ranging from 12 to 24 hours depending on environmental conditions and colony location. Hatching success is notably high, often 70–90% in favorable years, and recent studies link elevated rates to increased neighbor density, which deters gull predation, and proximity to foraging resources, allowing sustained parental attendance. Anti-predator spacing within colonies further bolsters success by diluting individual risk through synchronized vigilance and mobbing behaviors. Upon hatching, chicks are semi-precocial, covered in down and capable of limited , but remain flightless and confined to the ledge in a "grow-to-fly" stage. Both parents initially provision the chick with carried crosswise in the bill, but after about 15–20 days, the female departs the , leaving the male to deliver the final feedings and guard the chick exclusively. Fledging occurs at 20–25 days, when the chick, at roughly 25–30% of adult mass, leaps from the cliff , guided by the male who then provides at-sea care for 1–2 months until independence. Overall reproductive success varies from 60–96% across sites but averages 70–90% in resource-rich environments, influenced by egg , spacing, and prey availability that supports rapid chick growth.

Social behavior and vocalizations

Uria species exhibit highly colonial social structures, breeding in dense aggregations on steep cliff ledges where individuals nest in close proximity, often shoulder to shoulder, without building nests or using nesting material. These colonies can include up to 20 pairs per square meter, with synchronized breeding that facilitates collective anti-predator vigilance and . Within colonies, dominance hierarchies emerge based on arrival order, as early-arriving birds secure preferred sites near protective rock faces or "club stones," leading to aggressive interactions that condense individual territories into larger units over days. Vocalizations are essential for communication in these noisy colonies, varying between species in pitch and . The (U. aalge) produces a including soft purring notes for chick recognition, growls and croaks during agonistic encounters, and moans on breeding grounds. In contrast, the (U. lomvia) emits deeper, gruffer calls, such as crow-like "ha ha ha" utterances and frequency-modulated growls, with six distinct adult call types showing individual variation that supports parent-offspring recognition. These vocal differences, with U. lomvia calls being lower-pitched and more nasal, aid in territory defense and social coordination. Outside the breeding season, Uria form large flocks at , engaging in allopreening among group members to maintain social bonds and reduce ectoparasites, particularly targeting the head and neck regions. Agonistic displays include bill-jabbing, where birds thrust their bills at rivals, often accompanied by open-bill at higher intensities, escalating to bill-locking or wing-beating in prolonged disputes. For anti-predator defense, colonial nesting enables behaviors against avian threats like and skuas, where groups collectively harass intruders to deter and chick predation.

Migration and population dynamics

Migration patterns

Uria species exhibit post-breeding dispersal as their primary migration type, with adults and chicks moving southward from and breeding colonies to exploit seasonal prey resources and avoid harsh winter conditions. For the (Uria aalge), this involves longer southward dispersals to mid-latitudes, such as populations from and reaching waters off . In contrast, the (Uria lomvia) undertakes shorter, more localized movements to ice-free waters, often remaining in northern regions like the or shifting to the marginal ice zone in the North Pacific. These patterns reflect adaptations to differing environmental constraints, with U. aalge tolerating milder temperate waters and U. lomvia prioritizing access to open seas. Migration typically commences in August through October, coinciding with the completion of breeding and the onset of molt, when birds become flightless and rely heavily on swimming. Departures follow major ocean currents to facilitate efficient travel, such as the for Atlantic populations or the for Pacific ones, aiding southward progression while minimizing energy expenditure. Irruptive movements, where birds deviate from typical routes, occur in response to localized food shortages, as seen during marine heatwaves that disrupt prey availability and prompt unexpected southward irruptions. Return migrations northward begin in spring, often by March–April for U. aalge in the Atlantic, aligning with ice melt and prey blooms. Navigation during these dispersals relies on a combination of for directional orientation, visual landmarks near coasts, and wind patterns for aerial adjustments during partial flights. Seabirds like murres detect via proteins in the , enabling positional awareness over vast oceanic expanses. to wintering grounds is evident, with individuals showing consistent return to the same areas across years. Dispersal distances vary by species and population but can extend up to 5000 km for some U. lomvia groups, such as those from the traveling to the (approximately 3972 km on average). For U. aalge, distances are generally shorter, averaging around 1852 km from northern breeding sites to eastern North Pacific winter areas. These movements often involve initial swimming phases by parent-chick pairs, covering at least 1000 km in cases like northern populations. Prior to the 2014–2016 northeast Pacific , the global breeding of the (Uria aalge) was estimated at over 8 million pairs, with the majority in the Pacific (approximately 5 million pairs) and the remainder in the Atlantic (about 3 million pairs) (as of 2021). However, the heatwave caused the death of approximately 4 million common murres in (about 50% of the regional ), with aerial surveys confirming persistent colony reductions through 2024. For the (U. lomvia), the global breeding is around 11 million pairs, predominantly in regions (as of 2021). Breeding colonies of both range in from several thousand pairs to over 1 million pairs, with some of the largest concentrations historically occurring in the , where individual colonies supported hundreds of thousands of breeding pairs prior to recent declines. At a global level, populations of U. aalge and U. lomvia were relatively stable prior to recent events, but regional variations are evident, including declines of 20-30% at certain Atlantic sites for U. lomvia between 2000 and 2020. These shifts often involve birds relocating to nearby colonies in response to disturbances, such as increased predation or human activity, which can prompt temporary or permanent changes in colony occupancy. Local movements among colonies are typically short-distance and driven by factors like fluctuating food resources and predation risks, allowing birds to exploit optimal conditions within their breeding range. Banding and GPS tracking studies indicate strong , with about 90% of individuals exhibiting site fidelity by returning to the same colony across breeding seasons. Ongoing monitoring of population densities and trends employs aerial photographic surveys to large colonies efficiently, complemented by programs that contribute ground-based observations and data up to 2025.

Conservation

Threats and challenges

Uria species, including the (Uria aalge) and (Uria lomvia), face significant threats from oil spills, which can cause direct mortality and long-term reproductive impairments. The 1989 in , , resulted in the deaths of approximately 240,000 seabirds, with 74% being murres, leading to substantial population declines and reduced breeding success in affected colonies for years afterward. Bycatch in fishing gear, particularly gillnets, poses another major anthropogenic threat, entangling and drowning thousands of Uria individuals annually. Globally, gillnet fisheries are estimated to cause the of tens of thousands of alcids, including murres, with regional hotspots in the North Pacific and Atlantic contributing to cumulative mortality rates that exacerbate population declines. Climate change alters prey distribution and abundance, forcing Uria to forage farther or switch to less nutritious food sources, which reduces chick provisioning and overall survival rates. In the North Pacific, warming oceans have shifted like and , contributing to mass die-offs and breeding failures observed during the 2014–2016 . Other challenges include outbreaks of avian influenza, which have caused widespread mortality in murre populations. Highly pathogenic avian influenza (HPAI) H5N1, circulating since 2021, has led to mass die-offs during breeding seasons, with up to 46% infection rates in sampled murres in the northwestern Atlantic (Canada) by 2022; impacts continue in Alaska as of 2025. Invasive predators, such as introduced mammals like foxes and rats on breeding islands, prey on eggs and chicks, increasing nest failure rates in vulnerable colonies. Overfishing of key prey species, particularly capelin (Mallotus villosus), has depleted stocks in regions like the Barents Sea, resulting in nutritional stress and population decreases for Uria. As of 2025, emerging issues include , which impairs the and survival of prey such as euphausiids that supplement Uria diets, potentially reducing foraging efficiency in acidified waters. Increased storm frequency and intensity, driven by , disrupt breeding by causing colony abandonment and egg loss, with single events reducing success by approximately 9% in North Sea murre populations. These threats interact synergistically, compounding impacts on Uria ; for instance, climate-driven prey shifts combined with reduce diving success by altering energy budgets, while storms and contaminants together lower viability through increased predation and developmental abnormalities. Such cumulative effects have contributed to observed population declines across Uria ranges.

Conservation status and efforts

Both species of the genus Uria, the (U. aalge) and (U. lomvia), are classified as Least Concern on the global as of 2025, reflecting their large overall population sizes and wide distributions across the North Atlantic and North Pacific. However, regional subpopulations face heightened risks; for instance, the population of U. aalge has experienced significant declines due to historical factors, prompting targeted conservation under regional frameworks like the HELCOM Red List, though it remains part of the global Least Concern assessment. Key breeding and foraging habitats for Uria species are safeguarded through networks of protected areas, including sanctuaries in such as the Alaska Maritime National Wildlife Refuge, which encompasses critical colonies like those on the Semidi Islands. In the , 's protected zones, such as the North-East Svalbard Nature Reserve and bird sanctuaries like Storøya, provide essential safeguards for U. lomvia colonies. Similarly, in Newfoundland, sites like the Witless Bay Ecological Reserve and Terra Nova protect major U. aalge breeding grounds. These areas are supported by international agreements, including the OSPAR Convention for the North-East Atlantic, which designates marine protected areas for , and the African-Eurasian Migratory Waterbird Agreement (AEWA), which promotes coordinated conservation for migratory Uria populations across their flyways. Conservation efforts emphasize mitigation of human-induced pressures, with bycatch reduction initiatives focusing on gear modifications such as acoustic pingers and visual alerts on gillnets, which have demonstrated approximately 50% decreases in U. aalge entanglement in trials in . Oil spill response protocols, coordinated by organizations like the U.S. and Wildlife Service and International Bird Rescue, include rapid rehabilitation centers and standardized techniques that have improved post-release survival rates for oiled murres to over 70% in recent incidents. Habitat restoration projects, particularly on islands, involve predator removal—such as eradicating introduced mammals like foxes and rats—to facilitate colony re-establishment; for example, efforts at Devil's Slide Rock off have successfully attracted U. aalge breeders since 2005 by eliminating terrestrial predators. Ongoing research initiatives in 2025 leverage advanced technologies for monitoring, including AI-driven image analysis from drone and CCTV footage to automate counts for Uria , enabling precise tracking of numbers at remote sites with over 95% accuracy compared to manual surveys. Complementary studies on breeding success examine how nest and influence rates, revealing that U. aalge chicks at high- in Newfoundland experience up to 30% higher success due to enhanced neighbor vigilance and prey proximity.

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