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Gannet
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| Gannet Temporal range: Early Miocene to recent
| |
|---|---|
| Northern gannets (Morus bassanus) on Heligoland | |
| Northern gannet calls recorded on Grassholm Island, Wales | |
| Scientific classification | |
| Kingdom: | Animalia |
| Phylum: | Chordata |
| Class: | Aves |
| Order: | Suliformes |
| Family: | Sulidae |
| Genus: | Morus Vieillot, 1816 |
| Type species | |
| Pelecanus bassanus Linnaeus, 1758
| |
| Species | |
| Synonyms | |
|
Moris | |
Gannets are seabirds comprising the genus Morus in the family Sulidae, closely related to boobies. They are known as 'solan' or 'solan goose' in Scotland. A common misconception is that the Scottish name is 'guga' but this is the Gaelic name referring to the chicks only.
Gannets are large white birds with yellowish heads, black-tipped wings and long bills. Northern gannets are the largest seabirds in the North Atlantic, having a wingspan of up to two metres (6+1⁄2 feet). The other two species occur in the temperate seas around southern Africa, southern Australia, and New Zealand.
Etymology
[edit]"Gannet" is derived from Old English ganot, meaning "gander."[1]
Taxonomy
[edit]Morus is derived from Ancient Greek μωρός moros "stupid"[2] or "foolish" due to lack of fear shown by breeding gannets and boobies, allowing them to be easily killed.[3]
Behaviour
[edit]Hunting
[edit]Gannets hunt fish by diving into the sea from a height of 30 m (100 ft) and pursuing their prey underwater, and have a number of adaptations:
- They have no external nostrils; they are located inside the mouth, instead.
- They have air sacs in the face and chest under the skin, which act like bubble wrap, cushioning the impact with the water.
- The position of their eyes is far enough forward on the face for binocular vision, allowing them to judge distances accurately.[4]
Gannets can achieve speeds of 100 km/h (62.13 mph) as they strike the water, enabling them to catch fish at a much greater depth than most airborne birds.[5]
The gannet's supposed capacity for eating large quantities of fish has led to "gannet" becoming a description of somebody with a voracious appetite.[6]
Mating and nesting
[edit]Gannets are colonial breeders on islands and coasts, normally laying one chalky-blue egg. They lack brood patches and use their webbed feet to warm the eggs.[7] They reach maturity around 5 years of age. First-year birds are completely black, and subsequent subadult plumages show increasing amounts of white.
Northern gannets
[edit]The most important nesting ground for northern gannets is the United Kingdom, with about two-thirds of the world's population. These live mainly in Scotland, including the Shetland Isles. The rest of the world's northern-gannet population nests in Canada, Ireland, the Faroe Islands, and Iceland, with small numbers in France (they are present in the Bay of Biscay), the Channel Islands, Norway, and a single colony in Germany on Heligoland. The biggest northern-gannet colony is on Scotland's Bass Rock in the Firth of Forth; in 2014, this colony contained some 75,000 pairs.[8] Sulasgeir off the coast of the Isle of Lewis, St Kilda, Grassholm in Pembrokeshire, Bempton Cliffs in the East Riding of Yorkshire, Sceilig Bheag, Ireland, Cape St Mary's, Newfoundland, and Bonaventure Island, Quebec, are also important northern-gannet breeding sites.
Systematics and evolution
[edit]The three gannet species are now usually placed in the genus Morus, Abbott's booby in Papasula, and the remaining boobies in Sula. However, some authorities believe that all nine sulid species should be considered congeneric, in Sula. At one time, the various gannet species were considered to be a single species.
| Common name | Scientific name and subspecies | Range | Size and ecology | IUCN status and estimated population |
|---|---|---|---|---|
| Northern gannet (also known as "solan goose") | Morus bassanus (Lichtenstein, MHC, 1823) |
North Atlantic on coasts influenced by the Gulf Stream |
Size: Habitat: Diet: |
LC
|
| Cape gannet | Morus capensis Milne-Edwards, 1882 |
Southern Africa in three islands off Namibia and three islands off South Africa |
Size: Habitat: Diet: |
EN
|
| Australasian gannet | Morus serrator (Gray, GR, 1843) |
Coasts of New Zealand, Victoria, and Tasmania |
Size: Habitat: Diet: |
LC
|



Most fossil gannets are from the Late Miocene or Pliocene, when the diversity of seabirds in general was much higher than today. The cause the decline in species at the end of the Pleistocene is not clear; increased competition due to the spread of marine mammals may have played a role.
The genus Morus is much better documented in the fossil record than Sula, though the latter is more numerous today. The reasons are not clear; boobies possibly were better adapted or simply "lucky" to occur in the right places for dealing with the challenges of the Late Pliocene ecological change, or many more fossil boobies could still await discovery. Notably, gannets are today restricted to temperate oceans, while boobies are also found in tropical waters, whereas several of the prehistoric gannet species had a more equatorial distribution than their congeners of today.
Fossil species of gannets are:
- Morus loxostylus (Early Miocene of EC USA) – includes M. atlanticus
- Morus olsoni (Middle Miocene of Romania)
- Morus lompocanus (Lompoc Late Miocene of Lompoc, USA)[9]
- Morus magnus (Late Miocene of California)
- Morus peruvianus (Pisco Late Miocene of Peru)
- Morus vagabundus (Temblor Late Miocene of California)[9]
- Morus willetti (Late Miocene of California) – formerly in Sula[9]
- Morus sp. (Temblor Late Miocene of Sharktooth Hill, US: Miller 1961) – possibly M. magnus
- Morus sp. 1 (Late Miocene/Early Pliocene of Lee Creek Mine, US)
- Morus sp. 2 (Late Miocene/Early Pliocene of Lee Creek Mine, US)
- Morus peninsularis (Early Pliocene)
- Morus recentior (Middle Pliocene of California, US)
- Morus reyanus – Del Rey gannet (Late Pleistocene of W US)[9]
Cultural references
[edit]In many parts of the United Kingdom, the term "gannet" is used to refer to people who steadily eat vast quantities of food, especially at public functions.[10]


Young gannets were historically used as a food source, a tradition still practised in Ness, Scotland, where they are called "guga". Like examples of continued traditional whale harvesting, the modern-day hunting of gannet chicks results in great controversies as to whether it should continue to be given "exemption from the ordinary protection afforded to sea birds in UK and EU law". The Ness hunt is currently limited to 2,000 chicks per year and dates back at least to the Iron Age. The hunt is considered to be sustainable, since between 1902 and 2003 gannet numbers in Scotland increased dramatically from 30,000 to 180,000.[11][12]
In The Bookshop Sketch, originally from At Last the 1948 Show (1967), a customer (Marty Feldman) asks the bookshop proprietor (John Cleese) for "the expurgated version" of Olsen's Standard Book of British Birds, "the one without the gannet", because he does not like gannets owing to their "long nasty beaks". Desperate to satisfy the customer, the proprietor tears the page about the gannet out of the book, only for the customer then to refuse to buy it because it is damaged.[13][14] The sketch is reprised in Monty Python's Contractual Obligation Album, where the customer (Graham Chapman) says he does not like the gannet because "they wet their nests."[15]
In Series 1, Episode 3, of The F Word, Gordon Ramsay travels to the northwestern coast of Scotland and is shown how to prepare, cook and eat gannet.[16]
References
[edit]- ^ "gannet". Oxford English Dictionary (Online ed.). Oxford University Press. (Subscription or participating institution membership required.)
- ^ Liddell, Henry George; Scott, Robert (1940). "μωρός". A Greek-English Lexicon. Perseus Digital Library.
- ^ Jobling, James A (2010). The Helm Dictionary of Scientific Bird Names. London: Christopher Helm. p. 260. ISBN 978-1-4081-2501-4.
- ^ Wolfaardt, Leight. "Adaptations for Diving and Hunting Fact Sheet" (PDF). Birdlife.org. Retrieved August 7, 2023.
- ^ "Northern Gannet". oceanwide-expeditions.com. Oceanwide Expeditions. Retrieved 15 September 2024.
- ^ Bellincampi, Suzan (June 6, 2024). "Groovy Gannets". Vineyard Gazette. VG Media Group. Retrieved 15 September 2024.
- ^ Tucker, B.W. "Brood-patches and the physiology of incubation" (PDF). British Birds. 37 (2): 22–28. Archived from the original (PDF) on 2020-07-31. Retrieved 2017-03-07.
- ^ Munro, Alistair (18 March 2015). "Scotland's gannet population soars across country". The Scotsman. Retrieved 21 July 2017.
- ^ a b c d Miller, Loye (September–October 1961). "Birds from the Miocene of Sharktooth Hill, California" (PDF). The Condor. 63 (5): 399–402. doi:10.2307/1365299. JSTOR 1365299.
- ^ "Gannet: definition of gannet in Oxford dictionary (British & World English)". 2014-07-06. Archived from the original on July 8, 2012.
- ^ MacDonald, Fraser (2014-07-06). "The Hebridean guga hunt is 'ancient and sustainable', not a crime". The Guardian.
- ^ "BBC News – Gaga for guga: Ten things on Scottish island delicacy". BBC News. 2014-07-06.
- ^ "The Bookshop Sketch", MontyPython.net
- ^ Archived at Ghostarchive and the Wayback Machine: HarvestGod (26 November 2009). "Monty Python – Bookshop Sketch (Monty Python's Contractual Obligation Album)" – via YouTube.
- ^ William Young (2014). The Fascination of Birds: From the Albatross to the Yellowthroat. Dover Publications. p. 121. ISBN 9780486782935.
- ^ Archived at Ghostarchive and the Wayback Machine: "Gordon Ramsay's The F Word Season 1 Episode 3". YouTube. 11 June 2015.
External links
[edit]- Gannet videos on the Internet Bird Collection
Gannet
View on GrokipediaTaxonomy and etymology
Classification and species
Gannets comprise the genus Morus within the family Sulidae and order Suliformes, encompassing medium-to-large seabirds specialized for plunge-diving.[7] This placement reflects their close relation to boobies (genus Sula), from which gannets are distinguished morphologically by features such as the gular skin extending forward from the bill base, feathers projecting anterior to the eyes, and a heavier build facilitating dives from greater heights; behaviorally by breeding displays like sky-pointing; and genetically by sequence divergences confirming generic separation.[8][9][10] The genus Morus includes three extant species, whose monophyly is supported by multilocus phylogenetic analyses and control region DNA sequences showing distinct, well-supported clades with minimal interspecific gene flow despite ecological similarities.[10][11] These species exhibit subtle morphological variances, such as bill proportions and plumage nuances, alongside genetic markers indicating ancient divergence, with low population structuring within species but clear species-level boundaries.[12] The Northern gannet (Morus bassanus) represents the temperate North Atlantic lineage, characterized by genetic haplotypes clustering separately from southern congeners.[11] The Cape gannet (M. capensis) is endemic to southern African waters, distinguished by mitochondrial and nuclear markers reflecting isolation and adaptation to upwelling zones.[12] The Australasian gannet (M. serrator) occupies southern oceanic realms around Australia and New Zealand, with DNA evidence affirming its divergence and limited hybridization potential with other Morus taxa.[12][11] Historically, gannets were subsumed under Sula following Linnaeus's designation of Sula bassanus in 1758, but osteological studies revealed consistent skeletal disparities—such as cranium and sternum morphology—prompting generic reinstatement of Morus in the 20th century, later corroborated by molecular data rejecting a monotypic Sulidae genus.[13][14] This revision underscores the role of integrated evidence in avian systematics, prioritizing empirical distinctions over prior lumping based on superficial ecological convergence.[10]Etymology and nomenclature
The English word "gannet" derives from Old English ganot, denoting a sea-bird and ultimately tracing to Proto-Germanic *ganatuz or *ganutô, a term for "gander" or robust waterfowl, linked to the Indo-European root *ghans- signifying "goose" and evoking the bird's large, sturdy build.[15][16] This nomenclature reflects early Germanic observations of the bird's goose-like form and vigor, with cognates appearing in Old Frisian *ganet and Middle Dutch gent, emphasizing phonetic and semantic continuity across related languages.[15] Regional variants include the Scottish "solan goose," where "solan" stems from Old Norse súla (gannet) combined with ǫnd (duck), a designation recorded as early as 1450 in Older Scots texts and tied to North Atlantic breeding sites like the Bass Rock.[17][18] In scientific nomenclature, Carl Linnaeus formalized the northern gannet's binomial as Pelecanus bassanus in his 1758 Systema Naturae, with "bassanus" likely referencing the Bass Rock colony in Scotland as the type locality; subsequent reclassifications placed it under Sula and then Morus (from Greek môros, "foolish," alluding to terrestrial clumsiness), reflecting evolving ornithological understanding without altering the species descriptor.[13]Physical description
Morphology and adaptations for diving
Gannets of the genus Morus exhibit a large body size optimized for aerial spotting and plunge-diving, with adults typically measuring 80–110 cm in length, possessing a wingspan of 170–200 cm, and weighing 1–2.2 kg depending on species and sex.[19] This robust build includes a streamlined fuselage with a tapered profile that minimizes hydrodynamic drag upon water entry, complemented by a dagger-like bill averaging 10–12 cm long, reinforced for piercing fish at high velocities without fracturing.[20] The skeletal structure features a reinforced cranium and a specialized bone plate at the bill's base to distribute impact forces, while the humerus and coracoid bones in the shoulder girdle are proportionally robust to withstand deceleration stresses exceeding 10 g-forces during dives.[21] Powerful neck musculature, comprising hypertrophied sternocleidomastoid and splenius capitis muscles, enables rapid head tucking to align with the body's trajectory and absorb shocks up to 24 m/s entry speeds, preventing vertebral injury as modeled in biomechanical analyses of Morus bassanus.[19] Subcutaneous air diverticula—extensive, intercommunicating sacs extending from the cervical and thoracic regions—function as pneumatic cushions, compressing on impact to mitigate trauma from dives reaching depths of 10–22 m, with rare excursions to 35 m.[22] These adaptations reflect selective pressures for efficient energy transfer in piscivorous pursuits, prioritizing structural integrity over flight economy in Sulidae evolution.[20] Respiratory modifications include valvular nostrils that seal prior to immersion, preventing water ingress and facilitating rapid recovery, alongside expanded facial and pectoral air sacs that further buffer deceleration.[23] Ocular features encompass a prominent nictitating membrane for corneal protection and lens accommodation mechanisms enabling swift focus shifts from aerial to aqueous media, sustaining visual acuity underwater immediately post-entry.[24] Such traits collectively enable gannets to execute precision-oriented dives, with the absence of dense marrow in select long bones reducing overall mass while maintaining rigidity against compressive loads.[19]Plumage, dimorphism, and identification
Adult gannets of the genus Morus exhibit predominantly white plumage with black primaries forming distinct wingtip patches, a feature shared across the three species: Northern (M. bassanus), Cape (M. capensis), and Australasian (M. serrator).[25][26] The head and nape display a yellowish-buff wash, most pronounced in the Northern Gannet, subtler in the Cape Gannet, and minimal or absent in the Australasian Gannet.[27][28] Bills are pale blue to bluish-gray with black cutting edges, and eyes are pale blue surrounded by black bare skin.[25] Sexual dimorphism is minimal, with males averaging slightly larger in body mass, wing length, and bill dimensions than females, but no differences in plumage coloration or pattern.[26][29] This contrasts with some Sulidae relatives, such as certain boobies, where males may display ornamental plumes or brighter facial skin during breeding.[30] Juveniles fledge with mottled gray-brown plumage, featuring sooty brown upperparts speckled with white and paler underparts, gradually molting toward adult whiteness over 3–5 years through intermediate subadult stages.[25][31] First-year birds are largely dark with fine white flecks, while second- and third-year individuals show increasing white on the body and head but retain more extensive black on wings and tail.[26][32] Full adult plumage is typically attained by the fourth or fifth year.[33] Field identification relies on subtle species-specific traits: Northern and Australasian gannets have white secondaries and tails white with a black terminal band, whereas Cape gannets show black secondaries (except innermost) and an all-black tail.[27] Australasian gannets additionally feature a longer black gular stripe and darker tail feathers compared to Northern.[28] Age determination in the field uses progression from uniform juvenile brown to patchy subadult mosaics, with diagnostic bill color gradients (darker in juveniles) and tail shape (wedge-shaped in adults).[25][34] Banding studies confirm regional plumage variations are minimal, supporting these morphological markers for accurate identification.[30]Distribution and habitat
Global range across species
The Northern gannet (Morus bassanus) breeds exclusively in the North Atlantic Ocean across approximately 48 colonies, distributed from the British Isles and Iceland southward to about 47°N and northward to 74°N, including sites in Scandinavia and eastern Canada such as Newfoundland and the Gulf of St. Lawrence.[35] Post-breeding dispersals, tracked via geolocators and satellite tags, reveal oriented chain migrations southward to wintering areas off northwest Africa, the Mediterranean, and western Atlantic regions, with birds exhibiting fidelity to continental shelf waters rather than random vagrancy.[36] [37] The Cape gannet (Morus capensis) is endemic to southern Africa, breeding on six islands off Namibia and South Africa: Mercury, Ichaboe, and Possession Islands off Namibia; and Malgas Island, Bird Island (Lambert's Bay guano islands), and Bird Island (Algoa Bay).[38] [39] Juveniles undertake long-distance dispersals during the non-breeding season, extending from waters off the western Sahara eastward to near Tanzania in the southwest Indian Ocean.[40] The Australasian gannet (Morus serrator) maintains breeding colonies numbering over 30 along the southeastern coasts of Australia, Tasmania, and New Zealand, predominantly on offshore islands but with some mainland sites; a minor colony exists on Norfolk Island further north.[41] [28] Non-breeding ranges encompass adjacent shelf waters, with northward extensions along western and eastern Australian coasts.[42] These species exhibit largely non-overlapping distributions, segregated by ocean basins: M. bassanus in the northern Atlantic, M. capensis in southern Atlantic and western Indian Ocean waters, and M. serrator in the southwestern Pacific and southern oceans. Vagrancy remains exceptional, with M. bassanus documented in central and eastern European countries, the Black Sea, Bermuda, and Cuba, while M. capensis occurs rarely in KwaZulu-Natal, Mozambique, and Tanzania.[3] Tracking studies confirm post-breeding movements follow structured patterns, reinforcing geographic isolation and oceanic specialization across the genus.[36] In the Northern gannet, historical range expansions over the past century have included eastward shifts in Europe and the North Atlantic, driven by population increases, with notable colonization of previously unoccupied sites in the North Sea region.[43]Habitat requirements and environmental preferences
Gannets require breeding sites characterized by steep cliffs, sea stacks, or offshore islands that provide protection from terrestrial predators and facilitate aerial launch into prevailing winds. These locations typically feature bare rock ledges positioned above the splash zone and up to elevations exceeding 200 meters, minimizing ground-based threats while optimizing takeoff conditions. Such habitats correlate empirically with reduced mammalian predation pressure, as observed across species colonies where nesting density is high on exposed, inaccessible substrates.[44] For foraging, gannets preferentially exploit continental shelf waters where upwelling or nutrient-rich currents concentrate schooling fish prey, enabling plunge-diving from heights up to 30 meters. Northern gannets (Morus bassanus) favor temperate to subarctic shelf regions with cold-water productivity, avoiding deep pelagic zones beyond 200 meters depth except during non-breeding migrations. Cape gannets (Morus capensis) target subtropical Benguela Current upwelling zones off southern Africa, where sardine and anchovy schools aggregate seasonally, with foraging radii extending up to 350 kilometers from colonies but centered on shelf edges. Australasian gannets (Morus serrator) similarly prioritize shelf habitats in temperate to subtropical southern waters, adapting dives to detect prey via visual cues in productive nearshore environments.[45][1][3][46] Gannets demonstrate tolerance for moderate environmental variability, including temperature ranges from subtropical to subarctic latitudes tied to prey distribution rather than absolute thermal limits. They exhibit resilience by occasionally utilizing human-modified sites, such as disused structures on islands, provided these mimic natural cliff exposures and support nest construction without excessive disturbance. This adaptability underscores a primary dependence on prey availability over strict climatic niches, with foraging efficiency declining in depleted shelf waters.[2]Behavior and ecology
Foraging strategies and diet
Gannets forage primarily through plunge-diving, in which they spot prey from flight altitudes of up to 30 meters before folding their wings and impacting the water at speeds exceeding 100 km/h to pursue targets in shoals.[21] This technique targets small- to medium-sized pelagic fish aggregated near the surface, with dives typically lasting 5–10 seconds and reaching depths of 5–15 meters, supplemented occasionally by pursuit diving or surface pecking for accessible items.[20] Dive frequency averages 20–100 per foraging trip, adjusted to prey patch quality rather than fixed patterns.[20] The diet is overwhelmingly piscivorous, with over 90% consisting of forage fish such as Atlantic mackerel (Scomber scombrus), capelin (Mallotus villosus), herring (Clupea harengus), and sardines (Sardinops sagax), varying regionally by local abundance; for instance, in Newfoundland waters, mackerel comprised 41%, capelin 20%, and herring 11% of regurgitated items.[47] Short-finned squid (Illex illecebrosus) and other cephalopods constitute an opportunistic 5–17% of intake, exploited during fish scarcity or in overlapping distributions.[47] Prey selection favors energy-dense species forming dense schools, as evidenced by stable isotope analysis and chick-provisioning studies across North Atlantic colonies.[48] Foraging ranges from breeding colonies typically span 35–540 km, with mean maximum distances of 100–250 km per trip, determined by GPS tracking of hundreds of individuals; trip durations and extents correlate positively with colony size due to resource depletion around large aggregations but show inter-annual fluctuations linked directly to prey biomass rather than broad climatic trends.[49] [50] Northern colonies exhibit shorter commutes (e.g., 70–85 km) owing to higher polar-front productivity, minimizing flight costs estimated at 10–15 times basal metabolic rates during extended bouts.[51] Colony placement near upwelling zones or shelf edges optimizes net energy intake by balancing high plunge-dive expenditures—up to 20–30 kJ per dive inclusive of aerial search—against caloric returns from lipid-rich prey, consistent with models predicting habitat selection for maximal profitability over random search.[52] This strategy yields daily intakes of 500–1000 g for breeding adults, sufficient to offset 2–3 times field metabolic rates during chick-rearing, as quantified via doubly labeled water techniques.[53]Social structure, migration, and interactions
Gannets are highly gregarious seabirds that form large, dense colonies, or gannetries, supporting tens of thousands of pairs in close proximity; the Northern Gannet's Bass Rock colony, the world's largest, historically held approximately 75,000 pairs.[54] Spatial organization within colonies relies on ritualized displays and agonistic behaviors to defend nest sites and maintain minimal inter-nest distances of about 0.7–1 meter.[45] These structures facilitate collective vigilance but also intraspecific competition for limited space on cliff ledges or rocky islands.[1] Migration is partial and age-dependent across species. Northern Gannet juveniles disperse southward post-fledging via oriented chain migration along the Atlantic at 24–32 km/day, reaching tropical wintering grounds, while many adults remain nearer breeding sites.[55][56] Cape and Australasian Gannets exhibit more sedentary behavior, with adults foraging locally year-round and immatures undertaking regional dispersals, such as New Zealand-origin birds crossing the Tasman Sea to Australian waters.[42] Tracking studies confirm high individual consistency in non-breeding routes over years.[57] Conspecific interactions emphasize social foraging benefits over conflict. Gannets exploit local enhancement by aerially following successful divers to prey patches, improving capture efficiency through transferred information without cooperative hunting.[58] Intraspecific kleptoparasitism remains rare, with isolated observations during plunge dives but no widespread impact on energy budgets.[59] Predation on adults is infrequent, supported by annual survival rates of 91–92%, bolstered by aggressive defense, group vigilance in colonies, and low vulnerability at sea due to rapid flight and diving escapes from threats like sharks.[60][1] Alloparental care is absent, limiting cooperative rearing to monogamous pairs.[45]Reproduction and life history
Mating behaviors and pair bonding
Gannets are socially monogamous, forming long-term pair bonds that often persist for life, with pairs typically returning to the same nest site annually; studies of northern gannets (Morus bassanus) report 88–94% of individuals exhibiting this nest-site fidelity.[1] [61] Pair retention enhances breeding success, as established partners show higher reproductive output compared to newly formed pairs, though divorce rates vary widely (13–46% annually, averaging 22%) and correlate negatively with prior-year breeding performance and prey availability.[62] [63] Divorce appears adaptive under poor conditions, allowing birds to seek more compatible mates, but divorced individuals often experience reduced success in subsequent seasons until re-pairing stabilizes.[62] Courtship rituals reinforce pair bonds through synchronized displays, including sky-pointing—where one or both partners extend the neck, point the bill skyward, and spread wings—and fencing, involving rapid bill-thrusting and mutual touching to establish or reaffirm compatibility.[1] These behaviors, observed primarily upon site return in spring, minimize aggression and facilitate synchronization before egg-laying.[61] No evidence supports polygyny across gannet species; breeding sex ratios approach 1:1, consistent with genetic and observational data on monomorphic adults pairing monogamously without multiple mates.[61] Northern gannets typically first breed at 3–5 years of age, after acquiring experience in subadult attendance at colonies, with males often securing sites slightly earlier than females to attract partners.[64] Initial pairings may dissolve more readily due to inexperience, but surviving bonds contribute to lifetime fitness by reducing energetic costs of mate-searching and improving chick-rearing efficiency.[62] Similar patterns hold for Cape (M. capensis) and Australasian (M. serrator) gannets, though data are sparser.[61]Nesting colonies and parental investment
Gannets form large, dense nesting colonies on coastal cliffs, sea stacks, or islands, with northern gannets (Morus bassanus) occupying sites such as those in the Gulf of St. Lawrence or Bass Rock, where tens of thousands of pairs breed in close proximity, often within pecking distance of neighboring nests.[65][66] Nests are constructed primarily by males using materials like seaweed, grass, mud, and debris, forming a low, conical or cup-shaped mound compacted by guano and droppings, which accumulates over multiple seasons to create a pyramid-like structure on narrow ledges.[31][66] This dense packing defends against predators but intensifies intraspecific competition for space and resources. Females lay a single egg per clutch, rarely two, with incubation lasting 42-46 days and shared approximately equally between both parents, who take turns covering the egg with their brood patch to maintain temperature.[45] Parental investment during this phase involves continuous attendance to prevent chilling or predation, with no evidence of significant sex bias in effort allocation. Similar patterns occur in Cape (M. capensis) and Australasian (M. serrator) gannets, though colony-specific environmental factors influence exact durations.[67] Post-hatching, both parents provide biparental care through alternating foraging trips, regurgitating partially digested fish directly into the chick's mouth during frequent nest visits, which sustains rapid growth over the 82-99 day nestling period.[45][68] Provisioning rates vary with prey availability, but chicks face high mortality from starvation, with breeding success often ranging from 50-70% in average years, dropping lower during food shortages due to density-dependent depletion around large colonies.[69][70] Colony dynamics exhibit density dependence, where edge nests experience elevated failure rates from increased predation and exposure, while central nests benefit from collective vigilance but suffer from intensified competition for nearby forage, leading to longer parental trips in larger aggregations.[71][72] Small colonies often achieve higher per-nest success due to reduced intraspecific pressure, contributing to variable recruitment across sites.[73]Development and fledging
Gannet chicks exhibit rapid post-hatch growth, reaching near-adult body size within about 10 weeks through continuous parental provisioning of lipid-rich fish meals that meet their high energetic demands.[74] In Northern Gannets (Morus bassanus), nestling mass increases over 40-fold during the approximately 13-week nestling period, with energy density rising due to lipid accumulation that eventually comprises about 60% of tissue energy.[74] Chicks are brooded and fed frequently by at least one parent throughout rearing, supporting physiological milestones such as feather development from initial downy coverage to full plumage by fledging.[75] Fledging typically occurs at 82–99 days of age, with Northern Gannet chicks departing nests around 90–95 days by leaping from cliffs or platforms, often remaining nearby for additional days before sustained flight.[45][75] Towards the end of this period, parents progressively reduce feeding frequency to induce independence, prompting chicks to forage autonomously at sea.[76] Cape Gannet (M. capensis) chicks follow a comparable timeline, achieving fledging masses after 12 weeks of growth, though rates can vary with local prey availability.[77] Post-fledging survival remains challenging, with juvenile Northern Gannets experiencing high mortality; approximately 30% of fledged young survive to breeding age around 5 years, amid a potential lifespan exceeding 20 years supported by high adult annual survival rates of 92%.[64][75] Variations across species are minimal, though Northern Gannet chicks in colder northern climates may exhibit slightly extended nestling durations compared to subtropical Cape Gannets, reflecting adaptive responses to environmental provisioning constraints.[75][77]Conservation and threats
Population trends and status assessments
The Northern gannet (Morus bassanus) is classified as Least Concern by IUCN criteria, with an estimated global population of 1,500,000–1,800,000 mature individuals derived from colony censuses across the North Atlantic.[75] Overall trends indicate sustained increase since early 20th-century protections curtailed egg harvesting, with breeding colonies expanding from fewer than 20 sites to 54 by the 2020s and annual growth rates averaging 3% in major strongholds like British waters.[2][72] Metrics from aerial photography, ground counts, and color-banding returns confirm this rebound, though short-term fluctuations occur, such as a 6.7% decline in apparently occupied sites at Bass Rock from 2023 to 2024 and 51% at Funk Island in 2023.[78][79] These patterns reflect localized prey dynamics rather than overarching declines. The Cape gannet (Morus capensis) holds Endangered status under IUCN assessments, based on a global breeding population of 123,080 pairs equating to roughly 246,000 mature individuals from synchronized colony surveys in Namibia and South Africa.[3] Trends show ongoing decrease, with rates accelerating in recent generations to exceed 1% annually in regional subsets, resulting in over 50% loss at key sites like Namibia's colonies since the 1970s.[80][38] Annual aerial and boat-based censuses, supplemented by Bayesian state-space modeling for generational trends, underpin these estimates, highlighting vulnerability without implying uniform causation across populations.[81] The Australasian gannet (Morus serrator) is assessed as Least Concern globally, with breeding pairs totaling around 80,000–100,000 individuals across Australia and New Zealand based on periodic island surveys. Populations remain stable overall, featuring increases in Australian colonies from 6,600 pairs in 1980 to approximately 20,000 by 2000, though some New Zealand sites exhibit declines tied to episodic food shortages.[82] Banding recoveries and at-sea counts provide longevity data supporting resilience, with no evidence of rapid contraction despite localized variability from prey cycles.[83]| Species | IUCN Status | Estimated Mature Individuals | Trend | Key Data Sources |
|---|---|---|---|---|
| Northern gannet | Least Concern | 1,500,000–1,800,000 | Increasing overall | Aerial surveys, banding[75][2] |
| Cape gannet | Endangered | ~246,000 | Decreasing | Colony censuses, modeling[3] |
| Australasian gannet | Least Concern | ~160,000–200,000 | Stable | Island counts, recoveries[82][83] |
Anthropogenic impacts including bycatch
Bycatch in commercial fisheries represents a significant anthropogenic mortality factor for gannets, particularly in trawl and gillnet operations targeting pelagic fish. For the Northern Gannet (Morus bassanus), entanglement in midwater trawls accounts for the majority of incidents in the North Atlantic, with estimates indicating hundreds to thousands of individuals annually across monitored fisheries; for instance, U.S. east coast waters report fewer than 500 birds per year, deemed insufficient to drive population-level declines in this region. Globally, trawl bycatch affects over 44,000 seabirds yearly across 27 fisheries, with Northern Gannets among the most frequently captured species due to their foraging overlap with fishing grounds. Cape Gannets (Morus capensis) face similar risks in southern African purse-seine and trawl fisheries, where incidental capture contributes to overall mortality alongside prey depletion, though quantitative estimates remain limited compared to northern populations.[2][85][86] Historical exploitation exacerbated these pressures; prior to early 20th-century protections, intensive egg collection and adult harvesting—such as on remote islands like St. Kilda—severely depleted Northern Gannet colonies, reducing breeding pairs to historic lows by the late 1800s. Subsequent legal safeguards enabled marked recoveries, underscoring population resilience to targeted removal when ceased. Modern fisheries management shows variable integration of seabird data; while some quotas overlook bycatch thresholds, others mandate monitoring, yet inconsistent enforcement persists across jurisdictions.[87] Mitigation strategies, including bird-scaring lines (streamers) on trawls and discard management protocols, have demonstrated reductions in gannet interactions by 47-80% in tested scenarios, primarily by deterring foraging near nets and minimizing haul-back attraction. For longlines, weighted branch lines sink bait faster, curbing access, though gannets' plunge-diving behavior renders them less vulnerable than albatrosses to this gear. Empirical evidence from protected colonies indicates that, despite ongoing bycatch, Northern Gannet numbers have expanded substantially since the mid-20th century—exceeding 500,000 pairs—suggesting fishing mortality alone does not overwhelm intrinsic growth rates where habitat and prey remain viable, though cumulative effects warrant scrutiny in declining subpopulations like Cape Gannets.[88][80]Natural threats, disease, and resilience factors
Northern gannets face predation primarily on eggs and chicks from avian predators such as great black-backed gulls (Larus marinus), herring gulls (Larus argentatus), and skuas (Stercorarius spp.), which target vulnerable nests in dense colonies.[61] [89] Nestling losses also occur to mammals like red foxes (Vulpes vulpes) and short-tailed weasels (Mustela erminea) at accessible sites, though adults experience low predation risk due to aggressive defense and size.[61] At sea, sharks and large predatory fish occasionally take fledglings or water-landed adults, but such events remain infrequent relative to colony-based threats.[1] Severe weather, including storms, contributes to nest failures by dislodging eggs or chicks from cliff ledges, though gannet colonies are typically established on exposed, wind-resistant rock stacks and islands that minimize widespread structural collapse.[64] Highly pathogenic avian influenza (HPAI) H5N1 emerged as a significant pathogen in 2022, affecting 41 of 53 Northern gannet colonies worldwide and causing elevated adult and nestling mortality, with estimates of over 13,000 deaths in Newfoundland alone and up to 11% of the Canadian breeding population lost.[90] [91] [92] Parasitic infections, such as renal coccidiosis (Eimeria spp.) and Sarcocystis meningoencephalitis, occur sporadically but lack evidence of population-level impacts or clinical significance in most cases.[93] [94] [64] Gannet populations exhibit resilience through high reproductive output, with breeding success often reaching 75% in stable colonies, enabling numerical recovery via density-dependent regulation where reduced competition post-mortality boosts subsequent fledging rates.[64] Survivors of HPAI events have demonstrated capacity for renesting, though short-term declines of 20-40% in affected areas underscore limits to immediate rebound without broader ecological buffers.[95] [96]Evolutionary history
Fossil record and phylogenetic relationships
The fossil record of gannets (genus Morus) primarily dates to the Miocene epoch, with the family's main evolutionary radiation occurring between 15 and 25 million years ago.[13] A key specimen, representing an extinct sulid species closely allied to Morus, consists of a partial humerus from the Langhian stage (middle Miocene, approximately 15 million years ago) at Penedo beach, Setúbal Peninsula, southwestern Portugal; this fossil exhibits osteological features indicative of plunge-diving adaptations akin to those in extant gannets, such as a robust deltopectoral crest for enhanced flight muscle attachment during aerial pursuits of prey.[97] Miocene sulid fossils from other regions, including Peru along the Pacific coast of South America, further document early diversification of the group, with morphologies emphasizing specialized aquatic foraging. Subfossil and archaeological evidence extends prehistoric records of Morus into more recent periods. In the Adriatic Sea, the earliest confirmed remains from Palagruža Island, dated to the Holocene, mark the first fossil occurrence of the genus in the region and imply potential breeding activity in a warmer paleoenvironment than today.[98] Around the British Isles, prehistoric coastal sites yield bones of the northern gannet (Morus bassanus), often alongside those of extirpated seabirds like the great auk, reflecting exploitation or natural deposition in now-abandoned nesting areas.[99] Such finds, spanning the late Pleistocene to historic times, indicate persistence in North Atlantic localities without evidence of genus-wide extinction events, unlike some booby lineages.[100] Phylogenetically, Morus comprises a monophyletic clade within Sulidae, sister to the boobies (Sula and Papasula), with divergence from this lineage estimated at approximately 23 million years ago based on fossil-calibrated molecular clocks applied to multilocus data.[10] These analyses, incorporating strict and relaxed clock models, align the split with early Miocene marine transgressions that expanded piscivorous niches, preserving core sulid traits like streamlined fusiform bodies and precise plunge-diving via conserved skeletal proportions.[101] Unlike Sula, which shows greater species turnover in the fossil record, Morus lacks documented post-Miocene extinctions, suggesting relative stability tied to temperate-cold water foraging specialization.[13]Human interactions
Historical exploitation and management
Northern gannets (Morus bassanus) faced significant exploitation prior to the 20th century, primarily through egg collection, hunting for meat and feathers, and localized trade in Scotland. Colonies such as Bass Rock and Ailsa Craig supplied "solan geese" to Edinburgh markets, where birds fetched prices around 20 pence each in the 18th and 19th centuries, with tenants on Ailsa Craig historically paying rents in harvested gannets.[102] Egging practices involved systematic removal during breeding seasons, disrupting reproduction, while limited shooting targeted adults for food and plumage.[103] These activities contributed to drastic global population declines in the 19th century, reducing breeding pairs to critically low levels across North Atlantic sites.[103][104] Regulatory responses emerged in the early 20th century, with formal protections accelerating post-World War II through the establishment of bird reserves and legal safeguards in the UK and Canada. In the UK, sites like the Bass Rock became managed reserves under organizations such as the National Trust for Scotland, prohibiting unauthorized harvesting while allowing monitored access; similar measures in Canada protected key colonies like Bonaventure Island starting in the mid-1900s.[104][105] Management also included targeted culling of invasive predators, such as rats or feral mammals on peripheral islands, to safeguard nest sites, though Northern gannet colonies on remote sea stacks often required minimal intervention due to natural isolation.[106] These measures proved empirically effective, enabling steady population recovery throughout the 20th century from historic lows to over 263,000 breeding pairs by the 1980s, with adult survival rates stabilizing at approximately 92%.[104][107] The rebound, driven by cessation of direct persecution rather than intensive modern techniques like habitat engineering, underscores the sufficiency of core protections in restoring viability without expansive ongoing interventions.[104][108]Cultural and symbolic references
In Scottish folklore, the northern gannet (Morus bassanus) is traditionally known as the "solan goose," a term derived from Old Norse súla, evoking its pillar-like stance or keen visual acuity for spotting prey through water surfaces.[109] This nomenclature appears in historical accounts of Hebridean island life, where the bird's colonies on sites like Sula Sgeir influenced local place names and seasonal harvesting practices, though without documented prophetic or supernatural omens beyond practical associations with maritime abundance.[110] In heraldry, the gannet serves as a symbol of the Prophet, linked to its dramatic plunging dives interpreted as emblematic of divine insight or descent.[111] Among Māori, the Australasian gannet (Morus serrator), termed tākapu, holds cultural recognition as a coastal seabird integral to traditional observations of marine ecosystems, though lacking elevated mythological status.[28] Gannets feature peripherally in Anglo-Saxon literature, referenced alongside other seabirds in glosses and riddles denoting maritime peril or vigilance, as in Exeter Book entries equating ganot with predatory sea fowl. The gannet's high-velocity plunge-diving biomechanics have inspired modern engineering, particularly in biomimetic designs for aerial-to-aquatic vehicles; studies replicate its streamlined head and skeletal adaptations to mitigate impact forces during water entry, informing submersible prototypes and safer crash structures.[112][113] No major religious iconography attaches to gannets across traditions, distinguishing them from more symbolically laden birds like eagles or doves.References
- https://en.wiktionary.org/wiki/gannet
