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Fulmar

Fulmars are a (Fulmarus) of tube-nosed seabirds in the family , consisting of two extant species: the (Fulmarus glacialis) and the Southern Fulmar (Fulmarus glacialoides). These robust, medium-sized birds, resembling but with stiffer wings and tubular nostrils on their stout bills, are adapted to life over cold waters, where they glide effortlessly and scavenge or forage for , squid, crustaceans, and using a keen . The breeds in large colonies on steep coastal cliffs across the North Atlantic, North Pacific, and regions, from and to , the , and , with populations that have expanded dramatically since the due to access to human food waste like discards and carcasses. Monogamous and long-lived—often exceeding 30 years, with some reaching 50 or more—these birds defer breeding until 8–10 years old, laying a single per year in cliff crevices and fiercely defending nests by regurgitating foul-smelling stomach oil. They exhibit plumage polymorphism, ranging from pale gray-and-white to dark gray morphs, and forage over deep waters up to 600 miles from colonies during the breeding season. In contrast, the Southern Fulmar inhabits the around and subantarctic islands, breeding on rocky slopes and cliffs in massive colonies, such as those on the South Sandwich and , with a global population estimated at around 4 million individuals. Paler overall with silvery-gray upperparts, white underparts, and a distinctive white wing patch, it is slightly smaller-billed than its northern counterpart and migrates northward during the non-breeding season, reaching waters off , , , and . Like the , it forms a superspecies pair and shares ecological traits, including surface and attendance at fishing vessels, though its population remains stable despite emerging threats from , which may delay breeding and reduce . Both species are classified as Least Concern by the IUCN, serving as important indicators of due to their tendency to ingest plastics.

Taxonomy and Phylogeny

Etymology and Classification

The name "fulmar" originates from the term fúlmár, translating to "foul " or "foul-mew," a reference to the bird's defensive behavior of regurgitating a foul-smelling oil from its proventriculus to deter predators. The genus Fulmarus was established in 1826 by the English naturalist James Stephens in his work General Zoology, or Systematic Natural History, encompassing tube-nosed seabirds classified within the family . This placement is defined by key morphological adaptations, including the presence of naricorns—elongated, tubular nostrils that house supraorbital glands enabling efficient excretion of excess from diets and enhance olfaction for locating prey over vast ocean expanses. Members of , including fulmars, exhibit a characteristic split in the upper (culmen) that supports their filter-feeding and prey-capture mechanisms, alongside the of oil derived from partially digested marine prey. This oil serves dual purposes: providing a lightweight, energy-dense resource to sustain adults during extended flights across oceans and as a regurgitated nutrient for provisioning chicks in remote colonies. Additionally, these birds possess a highly developed , facilitated by enlarged olfactory bulbs and the naricorn structure, allowing detection of volatile compounds from prey such as and schools. Phylogenetically, Fulmarus is closely allied with other petrels and shearwaters within Procellariidae, forming a monophyletic clade distinct from the albatross family Diomedeidae; genetic analyses, including mitogenomic sequencing, confirm an ancient divergence between these groups estimated at approximately 60 million years ago, reflecting adaptations to different foraging ecologies.

Extant Species

The genus Fulmarus comprises two extant species of seabirds in the family Procellariidae, both adapted to marine environments in polar regions, and considered to form a superspecies. The Northern Fulmar (Fulmarus glacialis) is a medium-sized petrel measuring 45-50 cm in length, with a wingspan of 102-112 cm and a mass of 450-1,000 g. It is polymorphic, occurring in light morphs with predominantly white plumage on the head and underparts contrasted against gray upperparts, and dark morphs that are overall gray; intermediate forms also exist across its range. This species is well-suited to cold northern waters, where its robust build supports scavenging and surface feeding behaviors. In contrast, the Southern Fulmar (Fulmarus glacialoides) is monomorphic, featuring pale gray upperparts, white underparts, and darker wing tips, with a distinctive pale pink bill tipped in black. It measures 45-50 cm in length, has a wingspan of 110-120 cm, and weighs 720-1,180 g on average. Specialized for navigation over pack ice, it possesses morphological traits that enhance efficiency in windy, icy conditions. Key distinctions between the species include bill structure and wing proportions: the has a broader, more robust bill adapted for handling carrion and tougher prey, while the Southern Fulmar exhibits a longer, thinner bill and proportionally longer wings that facilitate sustained soaring over expansive ice fields. Both species are monogamous, forming long-term pair bonds; the can achieve lifespans exceeding 50 years, with some individuals reaching up to 60 years in the wild, while the Southern Fulmar reaches up to 45 years. The polymorphism observed in the is discussed in greater detail in the Plumage Variations section.

Fossil Record and Evolution

The fossil record of the genus Fulmarus is limited but provides key insights into its Miocene origins within the family. Two extinct have been described from middle to deposits in , dating to approximately 15–11 million years ago. Fulmarus miocaenus (, 1984), based on a complete from the Sharktooth Hill Bonebed in Kern County, was smaller than extant Fulmarus , with a humerus length of 78.3 mm, and exhibited a more developed bicipital area and distally placed ectepicondylar prominence compared to modern forms. Similarly, Fulmarus hammeri (, 1968), known from the proximal end of a carpometacarpus and a distal fragment from Laguna Hills in , was larger than living fulmars but differed in subtle morphological features of the wing elements. These specimens represent the earliest definitive records of the genus and indicate that fulmars had achieved a body size and skeletal proportions similar to those of today by the middle . The genus Fulmarus likely evolved from early procellariids during the late to early , around 25–20 million years ago, as part of the broader radiation of the family. This period coincided with the development of key for , including the tubular nostrils characteristic of procellariiforms, which enhance olfaction for detecting prey over vast open-ocean distances. The fulmarine , including Fulmarus, diverged from lineages leading to shearwaters (e.g., ) around 20–15 million years ago, based on molecular phylogenies calibrated with data. Another critical , the production of stomach oil from digested prey, originated in Eocene ancestors of the (approximately 40–50 million years ago) and enabled extended foraging trips by providing a high-energy food source for chicks during parental absences at sea. Mitochondrial DNA (mtDNA) studies support a radiation of procellariid lineages, including fulmars, during Miocene cooling events that expanded polar and subpolar habitats suitable for cold-water foraging. For instance, divergence estimates within Fulmarus place the split between northern and southern forms at 2–3 million years ago, aligning with intensified Pleistocene glaciations, though the core genus morphology remained stable post-Miocene. No Quaternary fossils of Fulmarus have been reported, suggesting a conserved evolutionary trajectory without major speciation events in the last 2.6 million years.

Description

Physical Characteristics

Fulmars are medium-sized seabirds with a stout build that supports their pelagic lifestyle. The (Fulmarus glacialis) measures 39–53 cm in length, with a of 100–112 cm, and weighs 450–1000 g. The Southern Fulmar (Fulmarus glacialoides) is similar in length at 45–50 cm but has a larger of 114–120 cm and weighs 720–1180 g. Their compact, robust frame includes a short that aids in maintaining during windy conditions at sea. This morphology, combined with a relatively short neck and large head, enables efficient energy use while over vast expanses. A defining feature of fulmars, as members of the order, is their distinctive tube-nosed bill equipped with paired naricorns—elongated, double-barreled nasal passages running along the top of the upper . These structures connect to supraorbital salt glands that allow fulmars to drink , excreting excess salt as a concentrated solution through the nostrils, which can amount to significant daily volumes relative to body weight to maintain osmotic balance. The naricorns also enhance olfactory capabilities, funneling scent molecules to olfactory receptors and enabling prey detection via smell from distances exceeding 5 km in some procellariiforms. Fulmars possess long, narrow wings with a high , typically around 10-12, optimized for where they exploit gradients to glide efficiently over long distances with minimal expenditure. Their legs are short and relatively weak, ill-suited for but well-adapted for , with webbed feet that provide when pursuing prey or pattering across the surface for takeoff. The Southern Fulmar has a slightly smaller bill than the . Internally, fulmars exhibit notable adaptations, including enlarged olfactory bulbs that represent one of the largest relative sizes among birds, with a diameter ratio to the of approximately 0.27 (27%) in the . Additionally, a specialized proventriculus in the produces and stores oil derived from dietary , which sustains adults and chicks during extended periods of over 100 days associated with and .

Plumage Variations

Fulmars exhibit notable plumage variations across species, particularly in the (Fulmarus glacialis), which displays polymorphism that influences its appearance and ecological adaptations. The light , more common in Atlantic populations, features a white head and neck contrasting with a grey mantle and wings, providing a distinctive during flight. In contrast, the dark , more common in Pacific populations, presents a uniformly slate-grey overall, with intermediate morphs occurring rarely and blending elements of both extremes. These variations are genetically determined and stable within individuals, though environmental factors may subtly influence expression. The Southern Fulmar (Fulmarus glacialoides), however, lacks such polymorphism, displaying a plumage adapted to its environment. Its body is pale grey with a white face and underparts, accented by black primaries on the wings, creating a subtle against snowy backdrops without the need for morphic diversity. This monomorphic form ensures consistency across populations, emphasizing functionality over variability. Molting cycles in fulmars are synchronized with and , involving a complete annual replacement post-breeding season that typically spans 2-3 months. During this period, birds undergo a gradual molt from head to tail, maintaining flight capability through sequential feather loss. Juveniles closely resemble adults by their first year, though they often appear with worn feathers during initial migrations, which are gradually replaced. Oil , facilitated by the , plays a crucial role in maintaining throughout these cycles, preventing matting and enhancing insulation. These plumage traits carry adaptive significance, with light morphs in the offering enhanced visibility in open oceanic waters for predator avoidance and foraging signaling, while dark morphs provide better concealment on rocky cliffs. Such variations underscore the evolutionary pressures shaping fulmar diversification, though size correlations with morphs are addressed in broader physical descriptions.

Distribution and Habitat

Northern Fulmar

The (Fulmarus glacialis) has a in the , encompassing the with key colonies in , , , as well as the including , the , and the , and extending to islands such as and the Canadian archipelago. The forms dense aggregations at remote coastal sites, with a global population estimated at approximately 7 million pairs (2016). Outside the breeding season, Northern Fulmars adopt a pelagic lifestyle, ranging widely over northern Atlantic and Pacific waters, often following currents and occasionally straying into subtropical regions. Vagrant individuals have been documented rarely in the , with confirmed records south of and off . Northern Fulmars select nesting on exposed cliff ledges and steep slopes, typically at elevations up to 500 meters, favoring sites with minimal vegetation and protection from ground predators. For foraging, they target marine environments over the continental shelf and slope, where water depths range from 200 to 2000 meters, concentrating in areas of nutrient-rich upwellings that enhance prey availability. These birds show a strong preference for windy coastal microhabitats that promote dynamic flight and access to productive feeding zones, such as shelf breaks with cold-water upwellings supporting , , and . Their distribution has expanded inland and to new coastal areas in recent centuries, facilitated by the exploitation of human-generated like discards, which supplements sources. Recent studies indicate potential shifts in breeding distribution due to warming oceans (as of ).

Southern Fulmar

The Southern Fulmar (Fulmarus glacialoides) breeds along the , , and various sub-Antarctic islands including the and , forming large colonies on ice-free cliffs and steep rocky slopes. The global breeding population is estimated at approximately 2 million pairs (corresponding to around 4 million individuals as of 2016), with significant concentrations in the region and South Atlantic Ocean islands, where up to 72% of pairs nest. During the non-breeding season, the species exhibits a circumpolar distribution across the , closely following pack ice edges and marginal ice zones rich in prey aggregations. Rare vagrants occasionally appear further north, reaching as far as , , , , and southern Brazil. This fulmar prefers nesting habitats on steep rocky slopes and ice cliffs that provide shelter from harsh weather, often in close association with penguin colonies where it may scavenge food scraps. It forages primarily in marginal ice zones of the , exploiting areas with high biological productivity near swarms. The species tolerates extreme cold down to -40°C and thrives in environments influenced by katabatic winds, which aid its dynamic over icy terrains. Recent observations suggest potential alterations from reduced due to (as of 2023). Its uniform provides effective against ice, enhancing survival in these polar habitats.

Behavior and Ecology

Breeding Biology

Fulmars are long-lived seabirds that form monogamous , with high site fidelity as breeding pairs typically return to the same cliff ledge annually. This is maintained through mutual and vocal displays upon reunion at the , contributing to reproductive stability over decades, as pairs may together for 40 or more years. initiation varies by and hemisphere: northern fulmars begin arrival in late winter (), with egg-laying peaking in May–June at lower latitudes and later in the ; southern fulmars start in October–November, aligning with the summer. Nesting occurs in colonies on steep cliff ledges or rocky slopes, where pairs scrape shallow depressions in bare rock or scant vegetation, often 7–20 cm across and minimally lined with pebbles or grass. sizes range from small groups of a few pairs to massive aggregations exceeding hundreds of thousands of pairs at major sites, such as those in for the and the for the Southern Fulmar; this coloniality provides safety in numbers but also intensifies competition for prime ledges. Fulmars aggressively defend their nests against intruders, including conspecifics and predators, by regurgitating foul-smelling stomach oil—composed of from their —that can be projected up to 2 meters with surprising accuracy. Each pair lays a single large white , weighing about 10–15% of the female's body mass, with shared by both parents in shifts lasting 2–16 days, totaling 47–55 days until hatching. The , covered in gray down, remains in the nest under guard for the first 1–2 weeks while parents alternate trips, often exceeding 600 km round-trip to provision it with regurgitated , including energy-rich stomach that aids and . Fledging occurs at 50–60 days post-hatching, with the departing independently to ; ends abruptly thereafter, though some post-fledging feeding may occur at sea. Breeding success varies annually with food availability and weather, reaching 70–90% in favorable years where most eggs hatch and chicks fledge successfully, though averages can drop to 40–50% in poor conditions due to starvation or predation. Fulmars exhibit delayed maturity, with most individuals not breeding until 8–10 years old, a trait that buffers populations against short-term environmental fluctuations but leads to lagged responses in overall numbers.

Feeding Habits

Fulmars are opportunistic feeders with diets dominated by and , varying by species and region. In the (Fulmarus glacialis), typically comprise 47–93% of the diet by wet mass, squid around 5–8%, and s such as decapods, amphipods, and copepods 3–13%, with smaller contributions from like pteropods (up to 16%) and carrion. Northern fulmars exhibit greater opportunism, incorporating carrion and where available, though natural prey forms the core. In contrast, the southern fulmar (Fulmarus glacialoides) focuses more on (Euphausia superba), which can account for 36% by mass alongside 63% like Pleuragramma antarcticum in Prydz Bay, with squid and other s minimal at 1–5%. Dietary composition shifts seasonally; for instance, intake increases toward late chick-rearing in some northern populations. Fulmars employ surface-based foraging techniques, including dipping (head submersion while hovering or walking on water), surface-seizing (snatching prey from the surface in flight), and pursuit plunging or to depths of up to 5 m to chase schools of or . They travel substantial distances from colonies, with daily foraging ranges of 100–500 km during and chick-rearing, though trips can extend to 1,000 km or more in some cases. A highly developed aids in detecting prey patches from afar. , where fulmars steal food from other seabirds, occurs rarely but has been observed in mixed foraging flocks. Nutritional adaptations include the production of stomach oil, a lipid-rich substance derived from partially digested prey that provides high-energy fuel for long bouts and chick provisioning, enabling efficient processing of energy-dense meals. During , pairs alternate duties, with one partner on land for up to two weeks while the other forages pelagically; post-fledging, fulmars shift to extended . This strategy supports their endurance flights and variable prey availability.

Flight, Vocalizations, and Sociality

Fulmars exhibit a distinctive flight style characterized by stiff-winged interspersed with shallow flaps, enabling efficient travel over vast oceanic distances. This locomotion relies on , where birds exploit —gradients in wind speed with altitude—to gain energy with minimal flapping, allowing sustained travel at airspeeds averaging around 13 m/s (approximately 47 km/h). Such techniques permit fulmars to cover hundreds of kilometers daily with low energy expenditure, as evidenced by satellite-tracked individuals traveling up to 369 km in a single day during non-breeding periods. Their broad wings, adapted for this , facilitate effortless wheeling in strong winds, often close to the water surface. Vocalizations in fulmars are primarily associated with social interactions rather than long-distance communication at , where they remain mostly silent except for occasional calls from young. During displays, they produce nasal cackling calls, such as the braying "aaark-aark" notes delivered at a rate of about three per second, often interspersed with rasping inspirations. In aggressive contexts, low growls and grunts accompany threats, signaling intent without physical contact. These , and throaty, underscore territorial boundaries and pair . Socially, fulmars are typically solitary or form small flocks while at sea away from food concentrations, contrasting with their dense breeding aggregations. In colonies, aggression manifests through aerial dives toward intruders and regurgitation of foul-smelling stomach oil, which can be projected several meters to deter rivals by matting feathers and impairing flight. Mate displays involve mutual head tossing and bill fencing, where partners wag necks back and forth while touching bills, reinforcing long-term pair bonds annually upon returning to nest sites. These birds exhibit high , with strong fidelity to natal colonies—particularly in the North Pacific—contributing to low dispersal rates. Fulmars boast exceptional , with adults averaging over 30 years and some individuals breeding for more than 40 years, supporting stable pair bonds that enhance .

Conservation Status

The Northern Fulmar underwent a dramatic population expansion during the , with numbers increasing approximately 10-fold across the Northeast Atlantic due to enhanced food availability from discards. In the , this growth was especially pronounced, as breeding pairs rose from roughly 10,000 in the early 1900s to about 1.5 million by 2000, allowing the species to colonize new coastal sites previously unsuitable for large-scale breeding. Recent trends indicate regional variation, with post-2020 declines of around 20% observed in select Scottish and North Atlantic colonies amid broader reductions exceeding 40% since the mid-1980s. In , however, populations remain stable or increasing at major sites such as St. Paul and St. George Islands, comprising about 80% of North American breeders. Southern Fulmar populations have held steady at approximately 4 million individuals worldwide, though breeding success fluctuates with extent and prey availability. Long-term monitoring, exemplified by a 50-year in , demonstrates that fulmar populations respond to ocean climate variations with delays of up to 5 years, as indices like the influence cohort recruitment and overall dynamics through effects on summer temperatures and breeding performance. The global fulmar population is estimated at 15-20 million individuals, reflecting the combined totals of both Northern and Southern . Fulmars' strong breeding site fidelity, often exceeding 90% in adults, can prolong recovery from such perturbations. Avian influenza outbreaks since 2022 have caused mortality in some individuals, with cases reported in areas like the in .

Threats and Protection Measures

Northern fulmars face significant threats from plastic ingestion, with recent Canadian data indicating that 52% of sampled individuals from between 2010 and 2023 contained 0.1 grams or more of in their stomachs. However, recent assessments indicate a significant decline in levels in northern fulmars over the past 20 years (as of ). This arises largely from surface-floating debris mistaken for food during . Bycatch in longline fisheries represents another major risk, with estimates suggesting annual mortality of around 3,920 northern fulmars in North Pacific groundfish fisheries alone between 2007 and 2014, potentially contributing 2-5% to adult mortality in vulnerable populations. exacerbates these pressures by altering breeding and prey availability; Antarctic seabirds, including southern fulmars, now lay eggs an average of 2.1 days later than in the early 1950s, linked to shifting and temperatures. For southern fulmars, declines in abundance due to warming waters have reduced breeding success, with performance notably lower in years of low concentration. Additional risks include oil spills, which can cause mass mortality events among fulmars through direct and habitat contamination. poses a threat, with fulmars susceptible to outbreaks that amplify mortality in dense colonies. Sustainable fishing regulations have diminished the availability of —a key food source for northern fulmars—leading to dietary shifts and increased vulnerability to other stressors. Pollutants accumulated during further compound cumulative physiological stress on both . Conservation efforts classify both northern and southern fulmars as Least Concern on the , with stable global populations as of 2025 assessments. The establishes marine protected areas that safeguard southern fulmar breeding habitats from exploitation and disturbance. coordinates ongoing population monitoring and threat assessments for both species across their ranges. To mitigate , the has implemented gear modifications and best practices in longline fisheries, while the Northwest Atlantic Fisheries Organization () enforces measures like weighted lines and night-setting restrictions. Research on climate adaptation highlights a 5-year in reproductive responses to ocean climate variability in fulmars, informing predictive models for population resilience. reserves provide safeguards for northern fulmar breeding sites, restricting human access and introduction to protect cliff-nesting colonies.

Human Interactions

Historical Exploitation

In the 18th and 19th centuries, the inhabitants of St. Kilda in the relied heavily on the for sustenance and resources, harvesting approximately 100-130 birds per person annually during the seasonal fowling period in late summer. Young fulmars were captured by hand or with fowling hooks on cliff ledges before they could fly, yielding around 12,000 birds in total each year for the island's small population. The meat served as a dietary staple, often dried in special huts for winter storage, while fresh eggs provided seasonal nutrition; feathers were stuffed into pillows and bedding, and the birds' stomach oil—extracted and stored in gut bags—was burned in lamps for lighting and fuel. By the mid-19th century, annual harvests reached about 9,600 birds, producing up to 566 gallons of oil in 1875 alone, which was traded with mainland for goods like cloth and tools. Beyond St. Kilda, indigenous Arctic peoples, including communities, traditionally exploited northern fulmars for food and eggs, consuming the birds' meat, using their oil for fuel, and gathering eggs as a vital protein source during breeding seasons. In , harvesting of fulmars in and the faced temporary bans in the late 1930s due to psittacosis outbreaks, a that could transmit to humans, though regulated resumed thereafter. The evacuation of St. Kilda's remaining 36 residents in 1930 ended intensive local harvesting there, allowing fulmar populations to rebound initially in the absence of human pressure. The held cultural significance in traditions, where its name derives from terms meaning "foul " or "foul-mew," referring to the bird's defensive ejection of foul-smelling stomach oil. In 19th-century , fulmar oil from St. Kilda became a minor trade commodity, bartered or sold for its preservative and lighting properties, though demand waned with the rise of cheaper alternatives like . While such exploitation caused localized population declines around key breeding sites like St. Kilda, the species proved resilient overall, with broader expansions in the North Atlantic facilitated by human fishing discards providing abundant waste as a food source.

Modern Significance and Conflicts

In contemporary environmental monitoring, the Northern Fulmar serves as a crucial bioindicator for marine litter in the North Atlantic and North Sea, with programs like the OSPAR Convention using stomach plastic content from beached birds to monitor pollution levels; as of assessments in the early 2020s, over 50% of North Sea fulmars exceed ecological quality thresholds of 0.1 g plastic per bird. The Southern Fulmar (Fulmarus glacialoides) plays a key role as a bioindicator for marine pollution in Antarctic and sub-Antarctic waters, with studies examining plastic ingestion in its stomach contents to assess debris levels in remote polar ecosystems. As a top predator in the Southern Ocean food web, it also bioaccumulates persistent organic pollutants such as polychlorinated biphenyls (PCBs), with concentrations fluctuating during the breeding season and reflecting dietary exposure from prey like krill and fish. These attributes make the species valuable for tracking long-term contaminant trends in regions where direct sampling is challenging due to ice cover and isolation. Human activities pose several conflicts for fulmar populations. For the Southern Fulmar, scavenging behavior draws birds to vessels, where they ingest fishery discards laced with s, exacerbating loads. Tourism in breeding colonies, such as those at , can cause disturbance by trampling nests and triggering adult flush responses, potentially reducing chick survival rates amid growing visitor numbers exceeding 100,000 annually. Additionally, research efforts involving satellite tagging to study patterns—revealing wide-ranging movements from breeding sites to subtropical grounds—may impose short-term stress, though benefits include informing strategies. Northern Fulmars similarly face risks from plastic ingestion and interactions with fisheries, contributing to their use in monitoring. On the positive side, fulmars contribute to ecosystem services by scavenging fishery waste, helping mitigate organic debris accumulation in ocean environments and indirectly supporting sustainable fishing practices. Antarctic ecotourism, featuring Southern Fulmar colonies as highlights for birdwatchers, generates substantial revenue—estimated at over US$800 million annually across the industry as of 2024—while promoting public awareness of polar conservation. Both species are integral to seabird population models, such as Leslie matrix projections that incorporate climate variability to predict breeding success and demographic trends over decades. Culturally, fulmars appear in wildlife documentaries, showcasing their graceful flight and colonial behaviors to educate global audiences on ocean biodiversity. Recent 2025 monitoring amid high-pathogenicity (HPAI) outbreaks in seabirds, including Southern Fulmars, underscores their resilience, with recorded lifespans exceeding 40 years in the wild enabling population recovery despite disease pressures.

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