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Caribbean monk seal

The Caribbean monk seal (Neomonachus tropicalis) was a medium-sized, extinct species of earless seal in the family Phocidae, endemic to the warm waters of the Caribbean Sea and Gulf of Mexico.^[1] Adults typically measured 2 to 2.4 meters in length and weighed 160 to 200 kilograms, with a robust, streamlined body covered in short, dense fur that was grayish-brown on the back and sides, paler yellowish on the underside, and slightly darker in females; newborns were born with a long, black woolly coat.^[2] Known for their docile and curious temperament, these seals hauled out on sandy or rocky beaches and cays to rest, breed, and nurse pups for 30 to 50 days, showing little flight response to human presence.^[3] The species was declared extinct by the International Union for Conservation of Nature (IUCN) in 1996 and delisted from the U.S. Endangered Species Act in 2008, with the last confirmed sighting occurring in 1952 at Seranilla Bank despite extensive subsequent surveys.^[4] First documented by Christopher Columbus in 1494 off the coast of modern-day Haiti during his second voyage to the Americas, the Caribbean monk seal was initially abundant across its range, with pre-exploitation population estimates ranging from 233,000 to 338,000 individuals distributed among at least 13 colonies.^[1]^[5] Taxonomically, it was reclassified in 2014 into the genus Neomonachus alongside the Hawaiian monk seal (N. schauinslandi), based on molecular, genetic, and cranial morphology analyses that distinguished it from the Mediterranean monk seal (Monachus monachus), highlighting its primitive phocid traits reminiscent of fossils from 14 to 16 million years ago.^[1] This reclassification underscored the Caribbean species' role as a key evolutionary link among extant monk seals, all of which are or were adapted to tropical and subtropical environments unlike most other pinnipeds.^[6] Ecologically, the Caribbean monk seal played a significant role as a benthic predator in shallow coastal reefs, consuming reef-associated fish, eels, octopuses, squid, and crustaceans, which required substantial biomass support estimated at 732 to 1,018 grams per square meter—levels that historical reefs provided at 3 to 6 times the capacity of modern Caribbean reefs.^[3]^[5] Colonies were centered in areas like the Bahamas, Greater Antilles, Yucatán Peninsula, and scattered cays such as Serranilla and Roncador Banks, where they formed harems and gave birth year-round, though peaking in winter months.^[1] Their vocalizations included grunts, barks, and snarls, and they exhibited social behaviors similar to other monk seals, but their range contracted rapidly due to human pressures, eliminating breeding sub-populations by the early 19th century.^[3] The primary driver of extinction was relentless human exploitation beginning in the 16th century, when European explorers and later colonists hunted the seals en masse for meat, oil (used for lighting and machinery), and hides, with documented slaughters including over 100 seals per night in the late 1600s and 200 individuals killed in a single 1911 event off Mexico.^[4]^[5] Overfishing of prey species and incidental persecution by fishermen further compounded the decline, restricting the species' range and preventing recovery despite early 20th-century protections under U.S. law; by the 1880s, sightings were rare, and no viable populations remained after the 1952 observation.^[1] As the only marine mammal driven to extinction by humans in tropical waters during historical times, the Caribbean monk seal serves as a stark example of anthropogenic impacts on marine biodiversity.^[5]

Taxonomy and evolution

Classification

The Caribbean monk seal is classified under the binomial name Neomonachus tropicalis (Gray, 1850), a change from its former designation as Monachus tropicalis to reflect its distinct phylogenetic position within the monk seals.^[7] This species belongs to the family Phocidae, comprising the true seals (earless seals or Phocida), and is placed in the subfamily Monachinae, which encompasses the three recognized monk seal species.^[8] Historical synonyms for N. tropicalis include Monachus tropicalis (Gray, 1850) and Phoca tropicalis (Gray, 1850), the latter reflecting early placements within the genus Phoca before taxonomic revisions.^[1] The type specimen, described by John Edward Gray, originated from Pedro Cays off the coast of Jamaica, marking the species' formal scientific recognition in 1850 based on material in the British Museum collection.^[8] The International Union for Conservation of Nature (IUCN) lists N. tropicalis as Extinct (EX), a status officially assessed in 2008 following the absence of confirmed sightings since 1952 and comprehensive surveys confirming no remaining populations.^[4]

Phylogenetic relationships

The Caribbean monk seal (Neomonachus tropicalis) belongs to the genus Neomonachus, which it shares with the Hawaiian monk seal (N. schauinslandi), forming a distinct New World clade within the subfamily Monachinae.^[9] This genus diverged from the Mediterranean monk seal (Monachus monachus) approximately 6.3 million years ago (95% highest posterior density interval: 4.98–7.64 million years ago), marking a deep evolutionary split in the late Miocene.^[9] The separation is supported by morphological and genetic evidence, including skull morphology and mitochondrial DNA (mtDNA) sequences, which highlight unique synapomorphies in the Neomonachus lineage, such as specific dental and cranial features not shared with Monachus.^[9]^[6] Within Neomonachus, the Caribbean and Hawaiian monk seals diverged around 3.7 million years ago (95% highest posterior density interval: 1.90–5.45 million years ago), an event contemporaneous with the final closure of the Isthmus of Panama, which likely influenced their biogeographic isolation across the Atlantic and Pacific.^[9] Genetic studies, particularly analyses of mtDNA cytochrome b sequences extracted from historical museum specimens, confirm that N. tropicalis is the sister taxon to N. schauinslandi, with limited genetic variation observed among Caribbean samples, suggesting a historically small effective population size.^[9] These mtDNA analyses, using maximum likelihood, Bayesian inference, and maximum parsimony methods, robustly place N. tropicalis closer to the Hawaiian species than to the Mediterranean monk seal, resolving prior uncertainties in monk seal phylogeny.^[9] No confirmed evidence of hybridization potential between N. tropicalis and other phocids has been identified in genetic data.^[9] Fossil evidence for the broader monachine clade, to which Neomonachus belongs, traces back to the late Miocene (approximately 10–5 million years ago) in the North Atlantic and Mediterranean regions, with early representatives indicating an origin in paratropical to temperate environments.^[10] Ancestral monachines from this period, such as undescribed forms from European and Atlantic deposits, show morphological traits transitional to modern monk seals, including reduced canines and specialized postcanine teeth adapted for a soft-bodied diet. Direct fossils of N. tropicalis are absent, but the clade's Atlantic diversification aligns with the species' historical range. Subfossil and archaeological records confirm the prehistoric presence of N. tropicalis across the Caribbean, including remains from the Yucatán Peninsula that attest to its exploitation by indigenous peoples.^[11] For instance, a femur fragment from the Postclassic Maya site of Isla Cerritos (ca. 1200–1500 CE) provides direct evidence of the species in coastal Yucatán contexts, while older archaeological bones from Antillean sites, dated to approximately 2,000–3,000 years ago, further document its regional abundance prior to European contact.^[11]^[5] These records, combined with paleontological data, underscore the species' long-term residency in tropical western Atlantic waters.^[5]

Physical description

Morphology

The Caribbean monk seal exhibited a streamlined, fusiform body typical of phocid seals, optimized for efficient swimming, with a robust torso supported by a thick layer of blubber that provided thermal insulation in tropical waters. The head was relatively large and rounded, featuring a short snout equipped with prominent vibrissae that served as specialized sensory organs for detecting hydrodynamic cues and prey movements in low-visibility conditions.^[12]^[13] The foreflippers were short and paddle-like, with five digits bearing well-developed claws on the anterior side and rudimentary nails on the posterior, while the hindflippers were broader and more flexible for primary propulsion during swimming; as in all phocids, there were no external ear flaps, only small pinnae openings covered by skin folds.^[12]^[14] The dental formula consisted of 2/2 incisors, 1/1 canines, 4/4 premolars, and 1/1 molars (totaling 32 teeth), with the posterior premolars and anterior molars forming carnassial pairs adapted for shearing and gripping slippery prey such as fish and squid.^[15] Females lacked prominent external genitalia, with the vulva reduced to a small, inconspicuous slit integrated into the smooth ventral body contour, further enhancing hydrodynamic streamlining alongside the short tail and naked palms and soles of the flippers.^[12]

Size and coloration

The Caribbean monk seal (Neomonachus tropicalis) exhibited a robust build typical of phocid seals, with adults reaching lengths of 2.0–2.4 meters from nose to tail.^[15] Males measured 213–244 cm, while females were slightly smaller at 199–224 cm, indicating modest sexual dimorphism in size but no pronounced differences in coloration.^[15] Estimated adult weights ranged from 170–270 kg, derived from comparisons with Hawaiian monk seals (N. schauinslandi) of equivalent length, though direct measurements were limited to a single captive female at 163 kg.^[16] Newborns measured approximately 75–100 cm in length and weighed 15–25 kg, with near-term fetuses recorded at 85–89 cm and about 17 kg.^[15] In terms of coloration, the dorsal surface was dark brownish-gray, often grizzled from light-colored tips on the short, stiff pelage hairs (6–10 mm long), transitioning to lighter yellowish-brown or pale yellow on the ventral side, sometimes with mottling.^[15] A greenish tint frequently appeared on the back and hind flippers due to algal growth on the fur, while the muzzle could show dusky graying in older individuals.^[15] Pups were born with a uniform dark brown to black woolly lanugo coat, which lightened to yellowish-gray in juveniles.^[15] Compared to other monk seals, the Caribbean species was smaller than the Mediterranean monk seal (Monachus monachus), which could reach up to 2.8 meters in length, but similar in dimensions to the Hawaiian monk seal, reflecting their close phylogenetic ties.^[15]

Habitat and distribution

Historical range

The historical range of the Caribbean monk seal (Neomonachus tropicalis) encompassed the Caribbean Sea, the Gulf of Mexico, and portions of the western Atlantic Ocean, extending from the Bahamas in the north to the Yucatán Peninsula in the west.^[15] Documented populations and sightings confirmed presence at key sites including the Bahamas, Jamaica, Cuba, the Yucatán Peninsula (such as the Triangle Keys and Arrecife Alacrán), and Serranilla Bank, a remote atoll midway between Jamaica and Nicaragua where the last verified sighting of a small colony occurred in 1952.^[15]^[17] The species' distribution also reached the Greater and Lesser Antilles, mainland coasts of Central America, and as far south as Guyana, with occasional records from the southern U.S. coast near Georgia.^[15] Subfossil and archaeological evidence reveals a broader prehistoric expansion across the West Indies and Central America, with remains dating to the Holocene period up to approximately 6,000 years ago, including sites in Florida (such as Long Bayou) and Puerto Rico (Taino Caracoles Midden).^[15] These findings, alongside Late Pleistocene fossils from Florida, indicate that the seals occupied a wide array of coastal and insular habitats long before European contact, likely utilizing both mainland beaches and offshore cays for haul-outs.^[15] Prior to European arrival around 1492, the population was estimated at 233,000 to 338,000 individuals, distributed across at least 13 colonies throughout the Caribbean region.^[18] Intensive hunting led to a drastic decline, with the population reduced to fewer than 500 individuals by 1900, fragmented into isolated remnants primarily in the central Caribbean near the Yucatán, Cuba, and Jamaica.^[4]^[15] The Caribbean monk seal exhibited largely sedentary behavior, with no documented evidence of long-distance migration or extensive seasonal movements; individuals typically remained associated with specific haul-out sites, forming dense colonies of up to several hundred on sandy beaches, rocky islets, or cays, though limited shifts between nearby mainland and offshore locations may have occurred.^[15] This non-migratory pattern aligned with the species' tropical habitat preferences, such as secluded cays and reefs within its range.^[15]

Preferred environments

The Caribbean monk seal (Neomonachus tropicalis) primarily inhabited warm tropical and subtropical waters of the western Atlantic Ocean, including the Gulf of Mexico and Caribbean Sea, where sea surface temperatures typically ranged from 24 to 30°C.^[19] These seals favored shallow, near-shore marine environments protected by coral reefs, which provided shelter and access to productive foraging grounds.^[15]^[5] For terrestrial use, the species preferred low-lying sandy beaches on isolated cays, atolls, and offshore islands for haul-out sites, often in areas with sparse or no vegetation and lacking fresh water sources.^[15] Historical accounts indicate that large groups, sometimes numbering in the hundreds, utilized these sites, including mainland beaches along North and Central American coasts prior to extensive human exploitation. They also occasionally hauled out on near-shore rocks and small rocky islets, though sandy substrates were predominant.^[15]^[12] In the aquatic realm, foraging occurred mainly in coastal waters to depths of up to 200 meters, with most activity concentrated in shallower zones less than 50 meters deep near reef systems.^[1] The seals' reliance on these tropical habitats made their populations centered in the central-western Caribbean, where offshore atolls offered relative isolation from disturbance.^[5]

Behavior and ecology

The Caribbean monk seal exhibited social tendencies, forming loose haul-out groups on sandy beaches and cays, often numbering 20 to 40 individuals, though historical accounts prior to intensive human exploitation describe gatherings of up to 500 seals.^[16]^[15] These groups lacked a rigidly defined hierarchical structure but facilitated non-reproductive interactions such as resting and social contact among adults and pups.^[16] On land, the seals communicated through a range of vocalizations, including growls, dog-like barks, pig-like snorts, moans, roars, and snarls, which likely served to maintain group cohesion and signal alarm or affiliation.^[20] Underwater vocalizations have not been documented and remain unknown for this species.^[4] The species displayed notable docility, characterized by curiosity and an absence of fear toward humans, with no observed aggressive defense behaviors even when approached closely.^[19] This unaggressive temperament contributed to frequent non-defensive interactions during haul-outs.^[21] Activity patterns were crepuscular, with peak activity at dawn and dusk, while much of the daytime was spent resting on land in groups; foraging excursions typically occurred during low-light periods.^[12]

Diet and foraging

The Caribbean monk seal (Neomonachus tropicalis) was a carnivorous piscivore with a diet consisting primarily of reef-associated fish, cephalopods, and crustaceans, inferred from osteological evidence and comparisons to its congeners, the Hawaiian and Mediterranean monk seals. Specific prey likely included reef fishes such as jacks (Carangidae) and grunts (Haemulidae), squid, octopuses, crabs, and lobsters, reflecting an opportunistic, generalist feeding strategy adapted to shallow tropical reef environments.^[22]^[2] No evidence supports herbivory or significant consumption of seabirds in its diet.^[22] Foraging occurred mainly in shallow coastal waters, where the seals engaged in bottom-feeding to capture demersal prey among coral reefs and seagrass beds. As a pierce feeder, N. tropicalis employed a combination of biting and suction to seize and consume prey, facilitated by specialized cranial morphology including an elongated rostrum and robust zygomatic arches for enhanced bite force.^[23] Highly sensitive vibrissae (whiskers) aided in prey detection by sensing hydrodynamic disturbances from nearby fish and invertebrates in low-visibility conditions.^[24] Dives were typically short and shallow, averaging 5-10 minutes at depths of 30-50 meters, similar to observed patterns in the closely related Hawaiian monk seal.^[25] Historical overfishing of reef fish and lobster populations in the Caribbean likely contributed to prey scarcity for N. tropicalis, exacerbating its vulnerability to extinction by reducing food availability in its preferred habitats.^[22] This depletion, combined with direct exploitation, disrupted the seal's trophic role as a mid-level predator influencing reef community dynamics.^[18]

Reproduction and life cycle

The Caribbean monk seal exhibited a polygynous mating system, with dominant males mating with multiple females, similar to its congeners, the Hawaiian and Mediterranean monk seals. Mating likely occurred in shallow waters near haul-out sites, with the peak breeding season spanning December to February, reflecting the species' tropical habitat and asynchronous reproductive patterns.^[15] Gestation lasted approximately 11 months, including a period of delayed implantation typical of phocid seals, allowing females to time births to favorable conditions.^[26] Birthing occurred primarily on sandy beaches or rocky shores, with a long pupping season peaking in early December, as observed in historical records from Mexico.^[19] Females typically gave birth to a single pup, though twins were rarely reported; newborns measured about 1 meter in length and weighed 16–18 kg, covered in a sleek black lanugo coat.^[15] Lactation lasted 4–7 weeks (30–50 days), during which mothers nursed pups on open beaches before weaning, with pups reaching approximately 50 kg by that stage based on patterns in closely related monk seals.^[3] Pups became independent shortly after weaning and were active like adults by around 9 months.^[15] Sexual maturity was attained at 3–5 years of age, with the smallest mature female recorded at 199 cm in length.^[15] In the wild, individuals had an average lifespan of about 20 years.^[19] The species' low reproductive rate, with females producing a pup approximately every other year rather than annually, limited population growth. This was compounded by high juvenile mortality, primarily from predation by sharks and killer whales, the main natural threats to young seals.^[27]

Human interactions

Historical exploitation

The Caribbean monk seal (Monachus tropicalis) faced initial human exploitation shortly after European contact in the Americas. During Christopher Columbus's second voyage in 1494, his crew encountered the seals—described as "sea wolves"—off the coast of present-day Hispaniola and killed at least eight individuals for meat to provision the expedition.^[4] Indigenous peoples in the region had likely hunted the seals to a lesser extent for meat and skins prior to European arrival, though records of such activities are sparse.^[28] Post-1492 colonization rapidly intensified exploitation, as the seals' docile nature and tendency to haul out in large groups on isolated beaches and cays made them easy targets for early mariners and settlers seeking fresh food during voyages.^[5] By the late 17th century, commercial harvesting escalated, driven by the seals' value for multiple resources. Blubber was rendered into oil for lighting lamps, lubricating machinery—particularly sugar cane mills on Caribbean plantations—and other industrial uses, while meat served as a reliable food source for sailors, fishermen, and colonists.^[28] Skins were processed into leather for clothing, bags, and straps, though this was secondary to oil production.^[4] Exploitation peaked in the 1800s, especially around the Bahamas and Cuba, where large rookeries in the western Caribbean and Gulf of Mexico were systematically targeted by organized hunting parties that clubbed seals on haul-out sites without regard for sustainable quotas.^[28] Hunting methods capitalized on the species' fearlessness, with groups approaching resting seals en masse and slaughtering them directly on beaches, often processing blubber on-site into barrels for transport.^[5] Oil and other products were traded internationally, with exports shipped to Europe and the United States to meet demand in lighting and manufacturing sectors.^[28] This unchecked harvesting decimated populations; historical models estimate the pre-exploitation population at 233,000 to 338,000 individuals across 13 major colonies, with hunting accounting for the near-total loss by the early 20th century.^[5]

Discovery and documentation

The earliest evidence of the Caribbean monk seal (Neomonachus tropicalis) comes from archaeological remains in pre-Columbian Maya sites on the northern coast of the Yucatán Peninsula, including a femur fragment and other bones from Isla Cerritos dating to around 600–900 CE, indicating indigenous awareness and possible utilization of the species long before European arrival.^[11]^[18] The first European record occurred during Christopher Columbus's second voyage in August 1494, when his crew encountered seals resting on the beaches of Alta Velo Island, near Cuba, and killed eight individuals—referred to as "sea wolves"—for food, marking the initial documentation in Western accounts.^[6]^[29] Formal scientific description of the species was provided by British zoologist John Edward Gray in 1850, based on a skin specimen collected from Jamaica and housed in the British Museum, which he named Monachus tropicalis in his Catalogue of the Specimens and Drawings of Mammals and Whales in the Collection of the British Museum.^[15]^[2] In the late 19th century, U.S. government scientific expeditions documented remaining populations, notably the December 1886 scientific expedition led by H.A. Ward and F. Ferrari Perez for the Mexican government to the Triangle Islands (also known as Serpientes Keys) off Mexico's Yucatán Peninsula, where 49 seals were killed and collected as specimens, providing key morphological data.^[20]^[28] Further surveys by the U.S. Bureau of Fisheries in 1910 targeted the same area but found no seals, signaling rapid decline.^[20] Captive records began in 1897 when two live Caribbean monk seals, captured from the Triangle Islands, were transported to the New York Aquarium; one female survived five and a half years until 1903, offering the first prolonged observations of the species in captivity, though it exhibited health issues and died young.^[20]^[30] Additional specimens arrived in 1909, but none thrived long-term.^[20] Throughout the 20th century, sporadic sightings by local fishermen provided the primary documentation, including reports from the Yucatán coast in the 1920s and 1930s, and a confirmed group of about 12 individuals observed in 1952 at Serranilla Bank, the last verified wild encounter.^[28]^[20] Post-2000 genetic analyses of museum specimens have advanced understanding, with studies extracting DNA from 19th-century skins and bones to confirm the species' distinct evolutionary lineage separate from other monk seals, as detailed in a 2014 phylogenetic analysis using mitochondrial and nuclear markers from six Caribbean specimens.^[9]^[6]

Extinction

Causes

The primary cause of the Caribbean monk seal's decline and extinction was direct hunting by humans, who targeted the species for blubber to produce oil, as well as for meat and skins, beginning in the late 15th century with early European explorers and intensifying during the 16th and 17th centuries.^[5]^[3] This exploitation drastically reduced the population, with historical estimates indicating 233,000–338,000 individuals across 13 colonies prior to intensive hunting, but only about 500 remaining in the northern Caribbean by 1836—a decline of over 99% within two centuries.^[5] Indirect effects of human activities, such as disturbance to breeding and haul-out sites, further exacerbated the impact by disrupting social structures and reproduction.^[4] Prey depletion through overfishing also contributed significantly to the seals' vulnerability, as their diet relied heavily on reef-associated species including groupers, snappers, and invertebrates, requiring an estimated 732–1,018 g/m² of biomass to sustain populations.^[5] Commercial fishing intensified from the 19th century onward, reducing Caribbean reef fish and invertebrate biomass to less than 25% of historical levels, which likely starved remnant seal populations and altered ecosystem dynamics by removing a key predator.^[5] Secondary factors included habitat loss from coastal development and human encroachment, which led to the abandonment of traditional rookeries and increased exposure to threats.^[4] While the species showed susceptibility to parasites like nasal mites in captivity, no major disease outbreaks were documented in wild populations, and there is no evidence that hurricanes or competition with introduced species played a substantial role in the pre-extinction decline.^[4] Climate change did not contribute to the extinction, as anthropogenic global warming effects were negligible during the period of decline.^[5]

Timeline and confirmation

The Caribbean monk seal was historically abundant in the early 1800s, with large populations reported across its range in the Caribbean Sea and Gulf of Mexico, but underwent a sharp decline due to intensive exploitation for oil, meat, and scientific collections following European colonization.^[4] By the mid-1880s, the species had become rare, with sightings limited to isolated areas such as the Triangle Keys in the Gulf of Campeche, where 49 individuals were killed during a scientific expedition in 1886.^[4] Further collections exacerbated the decline, including the killing of approximately 200 seals off the coast of Mexico in 1911 for scientific study.^[4] The last confirmed sighting in U.S. waters occurred in 1922 near Key West, Florida, where an individual was killed by a fisherman.^[4] The final authoritative sighting of the species took place in 1952 at Serranilla Bank in the western Caribbean, between Jamaica and the Yucatán Peninsula, marking the point after which no verified observations have been documented.^[4] Unconfirmed reports of possible monk seals persisted into the 1960s through the 2000s, primarily from local fishermen and divers in areas such as northern Haiti, northeastern Jamaica, and occasionally the Bahamas, though none were substantiated through photographic evidence or specimens.^[4] Efforts to locate surviving populations included aerial surveys by the U.S. Fish and Wildlife Service in 1973 across the Yucatán Peninsula to Jamaica, which yielded no evidence, as well as ground and aerial surveys in 1984 from the Gulf of Mexico to Campeche, Mexico, and in 1985 in northern Haiti, all without confirmed sightings.^[4] The National Oceanic and Atmospheric Administration (NOAA) conducted a comprehensive 5-year review starting in 2006, incorporating extensive historical analysis and recent field efforts through 2008, which found no evidence of persistence.^[31] The International Union for Conservation of Nature (IUCN) declared the species extinct in 1996, a status reaffirmed by NOAA in 2008 when it removed the Caribbean monk seal from the U.S. Endangered Species Act list due to extinction.^[4] In 2025, Mexico and the United States proposed deleting the species from Appendix I of the Convention on International Trade in Endangered Species (CITES) at the 20th Conference of the Parties (CoP20), recognizing its extinction.^[32] Since the 2008 declaration, no verified remains, live individuals, or environmental DNA have been detected in the wild, with genetic studies relying solely on historical museum specimens to analyze the species' phylogeny and population structure.^[9] Speculative discussions on de-extinction using genetic techniques, such as cloning from preserved samples, have occasionally surfaced in scientific literature, but these remain unfeasible due to technological limitations and ethical concerns over reintroducing a species into altered ecosystems.^[33]

References

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Nothing is retrieved...<|control11|><|separator|>
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