Thursday, June 16, 2016

THE ISLAND FOX

The island fox (Urocyon littoralis) is a small fox that is native to six of the eight Channel Islands of California. There are six subspecies of the fox, each unique to the island it lives on, reflecting its evolutionary history. Other names for the island fox include coast fox, short-tailed fox, island gray fox, Channel Islands fox, Channel Islands gray fox, California Channel Island fox and insular gray fox.
Urocyon littoralis pair.jpg

 

 

Taxonomy and evolution




The skull of an island fox (right) compared with a skull of the related gray fox (left).
 The island fox shares the Urocyon genus with the mainland gray fox, the species from which it is descended. Its small size is a result of insular dwarfism, a form of allopatric speciation. Because the island fox is geographically isolated, it has no immunity to parasites and diseases brought in from the mainland and is especially vulnerable to those that the domestic dog may carry. In addition, predation by the golden eagle and human activities devastated fox numbers on several of the Channel Islands in the 1990s. Four island fox subspecies were federally protected as an endangered species in 2004, and efforts to rebuild fox populations and restore the ecosystems of the Channel Islands are being undertaken. Radio collars are being attached to foxes in an effort to track and locate the young foxes. To date these efforts have been largely successful.
There are six subspecies of the island fox,[1] each of which is native to a specific Channel Island, and which evolved there independently of the others. The subspecies are:[1]
Foxes from each island are capable of interbreeding, but have genetic and phenotypic distinctions that make them unique; for example, the subspecies have differing numbers of tail vertebrae.
The small size of the island fox is an adaptation to the limited resources available in the island environment. The foxes are believed to have "rafted" to the northern islands between 10,400 and 16,000 years ago.[3][4] Initially, fox populations were located on the three northern islands, which were likely easier to access during the last ice age—when lowered sea levels united four of the northernmost islands into a single mega-island (Santa Rosae) and the distance between the islands and the mainland was reduced—it is likely that Native Americans brought the foxes to the southern islands of the archipelago, perhaps as pets or hunting dogs.[5][6]
Engraving of the Island Fox from the Pacific Railroad survey of 1855

The fox did not persist on Anacapa Island because it has no reliable source of fresh water; Santa Barbara Island is too small to support the food demands of the fox.
Rene Vellanoweth, an archaeologist, believes that inbreeding depression can be managed by mixing the different island fox subspecies populations much as the indigenous peoples did, by moving them from island to island, creating a higher genetic diversity and assisting them in recovery.[7]

 

 

 

Description


 


The island fox is significantly smaller than the gray fox and is probably the smallest fox in North America, averaging slightly smaller than the swift and kit foxes. Typically, the head-and-body length is 48–50 cm (19–19.5 in), shoulder height 12–15 cm (4.5–6 in), and the tail is 11–29 cm (4.5–11.5 in) long, which is notably shorter than the 27–44 cm (10.5–17.5 in) tail of the gray fox. This is due to the fact that the island fox generally has two fewer tail vertebrae than the gray fox.[8] The island fox weighs between 1 and 2.8 kg (2.2 and 6.2 lb). The male is always larger than the female.[9] The largest of the subspecies occurs on Santa Catalina Island and the smallest on Santa Cruz Island.[9]
The island fox has gray fur on its head, a ruddy red coloring on its sides, white fur on its belly, throat and the lower half of its face, and a black stripe on the dorsal surface of its tail.[9] In general the coat is darker and duller hued than that of the gray fox. The island fox molts once a year between August and November. Before the first molt pups are woolly and have a generally darker coat than adult foxes. A brown phase, with the grey and black fur of the body replaced by a sandy brown and a deeper brown, may occur in the San Clemente Island and San Nicolas Island populations.[10] It is unclear if this is a true color phase, a change that occurs with age, or possibly a change that occurs because of interactions with Opuntia cactus spines that become embedded in the pelt.[10]
An island fox kit nestled in the brush

Reproduction

The island fox typically forms monogamous breeding pairs, which are frequently seen together beginning in January and through the breeding season, from late February to early March. The gestation period is 50–63 days. The female island fox gives birth in a den, a typical litter having one to five pups, with an average of two or three. Pups are born in the spring and emerge from the den in early summer; the mother lactates for 7–9 weeks. Sexual maturity is reached at 10 months, and the females usually breed within the first year. Island foxes live for 4–6 years in the wild and for up to 8 years in captivity.[9]

Ecology and behavior

A nighttime shot of an island fox with three mice in its jaws.
Its preferred habitat is complex layer vegetation with a high density of woody, perennially fruiting shrubs. The fox lives in all of the island biomes including temperate forest, temperate grassland and chaparral, with no island supporting more than 1,000 foxes. The island fox eats fruits, insects, birds, eggs, crabs, lizards, and small mammals, including deer mice. The fox tends to move around by itself, rather than in packs. It is generally nocturnal, albeit with peaks of activity at dawn and dusk. Activity also fluctuates with the season; it is more active during the day in summer than it is in winter.[9]
The island fox is not intimidated by humans, although at first may show aggression. It is quite easy to tame and is generally docile.[9] The island fox communicates using auditory, olfactory and visual signals. A dominant fox uses vocalizations, staring, and ear flattening to cause another fox to submit. Signs of dominance and submission are visual, such as facial expression and body posture.[11] Its main vocalizations are barking and growling.[11] The island fox marks territory with urine and feces.

Conservation status and Federal Protection

 




In March 2004, four subspecies of the island fox were classified as a federally protected endangered species: the Santa Cruz island fox, Santa Rosa island fox, San Miguel island fox and the Santa Catalina island fox.[12] As of 2013, the IUCN lists the entire species as near threatened, an improvement from its previous status of "critically endangered".[2] A decline in island fox populations was identified in the 1990s. On San Miguel Island, the decline began in 1994, with the adult population falling from 450 to 15 in 1999. Similar population declines were discovered on Santa Cruz Island, where the population decreased from 2,000 adults in 1994 to less than 135 in 2000, and on Santa Rosa Island where foxes may have numbered more than 1,500 in 1994, but were reduced to 14 animals by 2000.[13][14] In 2004, there were 38 San Miguel island foxes, all in captivity; 46 foxes in captivity on Santa Rosa Island and 7 in the wild (golden eagle predation prevented the release of captive foxes into the wild); Santa Cruz Island had 25 captive foxes and a stable wild population of around 100 foxes.[14]
Golden eagle predation, discovered when foxes were radio-collared and monitored, proved to be the cause of the high mortality rates.[15] The golden eagle was an uncommon visitor to the Channel Islands before the 1990s according to data gathered by Dr. Lyndal Laughrin of the University of California Santa Cruz Island Reserve, and the first golden eagle nest was recorded on Santa Cruz Island in 1999.[16] Biologists propose that the eagle may have been attracted to the islands in the 1960s after the decline of the bald eagle. The golden eagle replaced the bald eagle and began to feed on feral pigs following the decimation of the local bald eagle population due to DDT exposure in the 1950s—the bald eagle would have deterred the golden eagle from settling on the islands while it subsisted on fish.[15]



 The feral pigs on Santa Rosa were exterminated by the National Park Service in the early 1990s, which removed one of the golden eagle's food sources. The golden eagle then began to prey on the island fox population. Feral pigs on Santa Cruz Island and deer and elk on Santa Rosa Island were introduced almost 70 years prior to island fox decline, therefore, the golden eagle most likely did not seek these animals as alternative prey.[17] This has occurred most likely as a result of a process known as 'apparent competition'. In this process, a predator, like the golden eagle, feeds on at least two prey, for example, the island fox and feral pigs. One prey item is adapted to high predation pressure and supports the predator population (i.e. pigs), whereas the other prey item (i.e. the island fox) is poorly adapted to predation and declines as a consequence of the predation pressure. It has also been proposed that the complete removal of golden eagles may be the only action that could save three subspecies of the island fox from extinction.[18] However, the pigs on Santa Cruz Island were killed by the Nature Conservancy under the idea that they brought the eagles to the foxes.[19]
Introduced diseases or parasites can devastate island fox populations. Because the island fox is isolated, it has no immunity to parasites and diseases brought in from the mainland and are especially vulnerable to those the domestic dog may carry. A canine distemper outbreak in 1998 killed approximately 90% of Santa Catalina Island's foxes, reducing the population from 1,300 to 103 in 2000.[16] A vaccination program has been initiated to protect Catalina Island foxes from canine distemper.[20] After several years of carefully trapping the foxes and vaccinating them against distemper and rabies, their population has reached 1,717 in 2015, surpassing the pre-disease population of about 1,300.[21] Scientists believe the distemper virus was introduced by a pet dog or a raccoon from the mainland that hitched a ride on a boat or a barge.[22] To eliminate the risk of disease, pets are not permitted in Channel Islands National Park.
Diminished food supply and general degradation of the habitat due to introduced mammal species, including feral cats, pigs, sheep, goats, and American bison, the latter having been introduced to Catalina Island in the 1920s by a Hollywood film crew shooting a Western,[23] also has had a negative effect on fox populations.
The foxes threaten a population of the severely endangered San Clemente Island loggerhead shrike in residence on San Clemente Island. The island fox population has been negatively affected by trapping and removal or euthanasia of foxes by the United States Navy. Since 2000, the Navy has employed different management strategies: trapping and holding foxes during the shrike breeding season, the installation of an electric fence system around shrike habitats, and the use of shock collar systems.[24] With the gradual recovery of the shrike population on San Clemente Island, the Navy no longer controls the foxes.
The populations of Santa Cruz island foxes, San Miguel island foxes, and Santa Rosa island foxes have dramatically rebounded from lows in 2000 of 70 for the Santa Cruz foxes and 15 each on San Miguel and Santa Rosa Islands.[25] The Catalina Island Conservancy runs a captive breeding program on Catalina Island.[26] On September 14, 2012, the US Fish and Wildlife Service released a draft recovery plan for the San Miguel island fox, Santa Rosa island fox, Santa Cruz island fox, and the Santa Catalina island fox.[27] By 2012, the Catalina Island Conservancy determined that there were 1,500 Santa Catalina island foxes and the population was stabilized.[28] As of 2015, there were 520 native foxes on San Miguel and 874 on Santa Rosa, according to the group Friends of the Island Fox. The number of foxes on Santa Cruz Island had risen to 1,750. The U. S Fish and Wildlife Service recommended delisting Santa Cruz, San Miguel and Santa Rosa island foxes in a major success of the Endangered Species Act. However, they are recommending that the Santa Catalina island be reclassified from endangered to threatened, because of the threat of diseases on this heavily visited island.[21]
Two other subspecies on San Nicolas and San Clemente aren't endangered. There were 263 foxes on San Nicolas and 1,230 on San Clemente.
Because the Channel Islands are almost entirely owned and controlled by either the Catalina Island Conservancy, The Nature Conservancy, or the federal government, the fox has a chance to receive the protection it needs, including constant supervision by interested officials without the ongoing threat of human encroachment on its habitat.



 Subspecies of island fox.png

THE GRAY FOX

The gray fox (Urocyon cinereoargenteus), or grey fox, is a carnivorous mammal of the family Canidae ranging throughout most of[3] the southern half of North America from southern Canada to the northern part of South America (Venezuela and Colombia).[1] No other canid's natural range spans both North and South America and it is the only American canid that can climb trees.[4] This species and its only congener, the diminutive Channel Island fox (Urocyon littoralis), are the only living members of the genus Urocyon, which is considered to be the most basal of the living canids.[5] Though it was once the most common fox in the eastern United States, and still is found there,[3][6][7] human advancement and deforestation allowed the red fox to become more dominant. The Pacific States still have the gray fox as a dominant. Its specific epithet cinereoargenteus means 'ashen silver'.



Urocyon cinereoargenteus.jpg

Origin and genetics

The gray fox appeared in North America during the mid-Pliocene epoch 3.6 million years ago (AEO) with the first fossil evidence found at the lower 111 Ranch site, Graham County, Arizona with contemporary mammals like the giant sloth, the elephant-like Cuvieronius, the large-headed llama, and the early small horses of Nannippus and Equus.[8] Genetic analyses of the fox-like canids confirmed that the gray fox is a distinct genus from the red foxes (Vulpes spp.). Genetically, the gray fox often clusters with two other ancient lineages, the east Asian raccoon dog (Nyctereutes procyonoides) and the African bat-eared fox (Otocyon megalotis).[9] Chromosome number is 2n=66.[10] Faunal remains at two northern California cave sites confirm the presence of the gray fox during the late Pleistocene.[11] Genetic analysis has shown that the gray fox migrated into the northeastern United States post-Pleistocene in association with the Medieval Climate Anomaly warming trend.[12] Recent mitochondrial genetic studies suggests divergence of North American eastern and western gray foxes in the Irvingtonian mid-Pleistocene into separate sister taxa.[13]
The gray fox's dwarf relative, the Channel Island fox, is likely descended from mainland gray foxes.[14] These foxes apparently were transported by humans to the islands and from island to island, and are descended from a minimum of 3-4 matrilineal founders.[13]

Description and behavior

Gray fox kit at the Palo Alto Baylands in California
A yawning gray fox, northern Florida
The gray fox is mainly distinguished from most other canids by its grizzled upper parts, black-tipped tail and strong neck, while the skull can be easily distinguished from all other North American canids by its widely separated temporal ridges that form a U-shape. There is little sexual dimorphism, save for the females being slightly smaller than males. The gray fox ranges from 76 to 112.5 cm (29.9 to 44.3 in) in total length. The tail measures 27.5 to 44.3 cm (10.8 to 17.4 in) of that length and its hind feet measure 100 to 150 mm (3.9 to 5.9 in). The gray fox typically weighs 3.6 to 7 kg (7.9 to 15.4 lb), though exceptionally can weigh as much as 9 kg (20 lb).[15][16][17] It is readily differentiated from the red fox by the lack of "black stockings" that stand out on the latter and the stripe of black hair that runs along the middle of the tail. In contrast to all Vulpes and related (Arctic and fennec) foxes, the gray fox has oval (instead of slit-like) pupils.[18]
The gray fox's ability to climb trees is shared only with the Asian raccoon dog among canids. Its strong, hooked claws allow it to scramble up trees to escape many predators, such as the domestic dog or the coyote,[19] or to reach tree-bound or arboreal food sources. It can climb branchless, vertical trunks to heights of 18 meters and jump from branch to branch.[20] It descends primarily by jumping from branch to branch, or by descending slowly backwards as a domestic cat would do. The gray fox is nocturnal or crepuscular and makes its den in hollow trees, stumps or appropriated burrows during the day. Such gray fox tree dens may be located 30 ft above the ground.[18] Prior to European colonization of North America, the red fox was found primarily in boreal forest and the gray fox in deciduous forest, but now the red fox is dominant in most of the eastern United States since they are the more adaptable species to development and urbanization.[21] In areas where both red and gray foxes exist, the gray fox is dominant.[22]

Reproduction

Gray fox, showing black tail stripe, Sierra Nevada
The gray fox is monogamous. The breeding season of the gray fox varies geographically; in Michigan, the gray fox mates in early March, in Alabama, breeding peaks occur in February. The gestation period lasts approximately 53 days. Litter size ranges from 1 to 7. Kits begin to hunt with their parents at the age of 3 months. By the time that they are four months old, the kits will have developed their permanent dentition and can now easily forage on their own. The family group remains together until the autumn, when the young reach sexual maturity, then they disperse.

Diet

A gray fox at night
Adult male and female gray fox
The gray fox is an omnivorous, solitary hunter. It frequently preys on the eastern cottontail (Sylvilagus floridanus) in the eastern U.S., though it will readily catch voles, shrews, and birds. In California, the gray fox primarily eats rodents, followed by lagomorphs, e.g. jackrabbit, brush rabbit, etc.[19] In some parts of the Western United States (such as in the Zion National Park in Utah), the gray fox is primarily insectivorous and herbivorous.[22] Fruit is an important component of the diet of the gray fox and they seek whatever fruits are readily available, generally eating more vegetable matter than does the red fox (Vulpes vulpes).[15]



 Leefgebied grijze vos.JPG

Wednesday, June 8, 2016

THE NORTHERN GANNET

The Northern gannet (Morus bassanus) is a seabird and the largest member of the gannet family, Sulidae. "Gannet" is derived from Old English ganot "strong or masculine", ultimately from the same Old Germanic root as "gander".[2] Morus is derived from Ancient Greek moros, "foolish" due to the lack of fear shown by breeding gannets and boobies allowing them to be easily killed. The specific bassanus is from the Bass Rock in the Firth of Forth, which holds the world's largest colony of northern gannets.[3]
The northern gannet has the same colours as the Australasian gannet and is similar in appearance. Nesting in colonies as large as 60,000 pairs on both sides of the north Atlantic this bird undertakes seasonal migrations and is a spectacular high-speed diver.
Old names for the northern gannet include solan and solan goose.

     
     
Morus bassanus adu.jpg
     
     

Description





Northern gannet breeding pairs

Northern gannet flying over the English Channel, in the 7 Islands Nature Reserve, northern France
Adults are 81–110 cm (32–43 in) long, weigh 2.2–3.6 kg (4.9–7.9 lb) and have a 165–180 cm (65–71 in) wingspan.[4][5] Before fledging, the immature birds (at about 10 weeks of age) can weigh more than 4 kg (8.8 lb). Each wing measures between 47 and 53 cm (19 and 21 in) when outstretched and the beak measures between 9 and 11 cm (3.5 and 4.3 in) (measured from the head). The two sexes are a similar size.
The plumage of the adults is white with dark wing tips, with colours that range from brown to black. The colour of the head, cheeks and side of the neck depends on the season and the individual; during breeding, the head and neck are brushed in a delicate yellow, although this colouring may not be evident in some individuals.[6] The feathers are waterproof, which allows the birds to spend long periods in water. A water-impermeable secretion produced by a sebaceous gland covers the feathers and the birds spread it across their body using their beak or their head.[7] The eye is light blue, and it is surrounded by bare, black skin, which gives the birds their characteristic facial expression.

Young northern gannet. The front part of its body shows adult plumage.

Young birds are dark brown
Fledglings are brown with white wing tips. They have white spots on their head and on their back and a v-shaped white area underneath.[8] The plumage of one-year-olds can be almost completely brown. In the second year the birds’ appearance changes depending on the different phases of moulting: they can have adult plumage at the front and continue to be brown at the rear.[9] They gradually acquire more white in subsequent seasons until they reach maturity after five years.
Newborn chicks are featherless and are dark blue or black in colour. In the second week of life they are covered in white down.[10] From the fifth week they are covered in dark brown feathers flecked with white.[6]
Their beak is long, strong and conical with a slight downward curve at the end. The front part has a sharp edge. In adults, the beak is blue-grey with dark grey or black edges. It is brownish in immature birds.
The northern gannet’s eyes are large and point forwards, and they have a light blue to light grey iris surrounded by a thin black ring. The four toes of their feet are joined by a membrane that can vary from dark grey to dark brown. There are yellow lines running along the toes that continue along their legs; these lines probably have a role in mating.[7] The rear toe is strong and faces inwards allowing the birds to firmly grip onto vertical cliff faces.[11]

Distinguishing anatomical features


Northern gannet on Bonaventure Island, in Quebec

Selecting a dive target

Plunge-diving with wings retracted
Northern gannets dive vertically into the sea at velocities of up to 100 km/h (62 mph) and the structure of their bodies is adapted for this practice. They do not have external nostrils and their secondary nostrils can be closed when they are in water. The opening of their auditory canal is very small and is covered with feathers; the openings can also be closed in water using a system that is similar to that used for the nostrils. The sternum is very strong and sufficiently long to provide protection for the internal organs from impacts with water.[12]
The lungs are highly developed and probably also play a role in reducing the effects of hitting water at high speeds and protecting the body from these effects. There are subcutaneous air sacs in the lower body and along the sides. Other air sacs are located between the sternum and the pectoral muscles and between the ribs and the intercostal muscles. These sacs are connected to the lungs and are filled with air when the bird breathes in. The air can be expelled by muscle contractions.[12]
Individuals have a subcutaneous fat layer, dense down and tightly overlapping feathers that help them withstand low temperatures. A reduced blood flow in the webbing on their feet outside of the breeding season also helps to maintain body temperature when they swim.[13]

By air, land and sea


Silhouette in flight
The wings of the northern gannet are long and narrow and are positioned towards the front of the body, allowing efficient use of air currents when flying. Even in calm weather they can attain velocities of between 55 and 65 km/h (34 and 40 mph)[14] even though their flying muscles are not highly developed: in other birds flying muscles make up around 20% of total weight, while in northern gannets the flying muscles are less than 13%.[15] The consequence of this is that northern gannets need to warm up before they begin flying. They also walk with difficulty and this means that they have difficulty taking off from a flat area. They take off from water by facing into the wind and strongly beating their wings. In light winds and high waves they are sometimes unable to take off and they can become beached.[14] They take advantage of the wind produced by the front of a wave in the same way as the albatross does. They are only seen inland when they have been blown off-course by storms.
They alight on water with their feet retracted. They rarely land on water with their feet stretched forward like pelicans or cormorants. When they are on the water their body is rather low in the water with their tail pointing diagonally upwards. They alight with difficulty on land and often with a bump as their narrow wings do not allow them to turn easily and they have to use their feet and tail to aid in these manoeuvres. Individuals often suffer damage to their legs or feet when they land on the ground if there is not sufficient wind.[16] Damaged or broken wings are a frequent cause of death in adults.[17] The position of the legs towards the rear of the body means that they walk in a similar way to ducks.

 

 

Northern gannet colony on Bonaventure Island near to Percé, Quebec, Canada

  Call

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Northern Gannet calls from Grassholm. Wales.
The northern gannet does not have a very characteristic acoustic repertory. Its typical call is rab-rab-rab, which is emitted when fishing and also when on the nest.[18] They have a special call when they approach the colony: this call is often heard because there is usually a lot of toing and froing in a colony.[19] Males and females make similar calls.
According to the ornithologist Bryan Nelson northern gannets can recognize the call of their breeding partner, chicks and birds in neighbouring nests. Individuals from outside this sphere are treated with more aggression.[20]

Distribution


Breeding colonies in the north Atlantic

Panoramic view of the Seven Island Nature Reserve that supports a northern gannet colony, in Brittany (France)
Their breeding range is the North Atlantic on coasts influenced by the Gulf Stream,[21] the exception being the colonies of the Gulf of Saint Lawrence and the islands off the east coast of Canada. They normally nest in large colonies, on cliffs overlooking the ocean or on small rocky islands. The waters near to these cliffs have a summer temperature at the surface of between 10 and 15 °C (50 and 59 °F).[22] The water temperature determines the distribution of Atlantic mackerel and herring, which are the main food source for the northern gannet. For this reason there is a close relationship between the location of northern gannet breeding colonies and the distribution of these fish. Northern gannet colonies can be found in the far north in regions that are very cold and stormy. The ornithologist Dr Bryan Nelson has suggested that they can survive in these regions due to a number of factors including: the combination of body weight and a strong beak that allows them to capture strong muscly fish and the ability to dive to great depths and capture prey far from the cliffs. In addition they are able to stand long periods without eating owing to their large fat reserves.[23]
The northern limit of their breeding area depends on the presence of waters that are free of sea ice during the breeding season. Therefore, while Greenland and Spitsbergen offer suitable breeding sites, the arctic regions have summers that are too short to allow the northern gannets to lay their eggs and raise a brood, which requires between 26 and 30 weeks.[24] The southern limit of their distribution mainly depends on the presence of sufficient prey.[25]
The species is a rare visitor in the Black Sea region.[26]

Breeding colonies


Northern gannet colony on Bonaventure Island near to Percé, Quebec, Canada
Some breeding colonies have been recorded as being located in the same place for hundreds of years. The cliffs containing the colonies appear to be covered in snow when seen from a distance, due to the number of nests present on them. There is a written record of a colony on the island of Lundy from 1274. It noted that the population was declining due to hunting and the theft of eggs. The colony finally disappeared in 1909.[27] 68% of the world population breeds around the coasts of Great Britain and Ireland. The biggest colonies include:
  • Bass Rock off the east coast of Scotland, first recorded in 1448.[28] In 2004, it contained more than 48 000 nests.[29] This is where part of the species’ Latin name comes from. The Bass Rock is now the world's largest colony of Northern gannets. Visitors to the Scottish Seabird Centre have the opportunity to see the Bass Rock via interactive live cameras in the Centre and on seasonal boat trips.
  • Saint Kilda and Sula Sgeir, in the Hebrides. Saint Kilda is the largest colony in Europe with more than 60 000 nests.[29]
  • Sceilg Bheag one of the Skellig Islands located off the South West Coast is the largest colony in Ireland hosting around 30,000 pairs.
  • Eldey off Iceland, where between 14 000 and 15 000 pairs breed.[30][31]
Other European colonies are found in the south west of Ireland, and off the west (Runde Island) and north of Norway (Syltefjord, Hovflesa and Storstappen). The most southerly European colony is on the island of Rouzic off the French Atlantic coast. There are breeding colonies along the coast of Newfoundland and on the islands in the Gulf of Saint Lawrence. The largest colony has 32 000 nests and is on Bonaventure Island off the south coast of Quebec.[29]

Population

A 2004 survey counted 45 breeding colonies and some 361 000 nests.[32] The population is apparently growing between 3% and 5% a year, although this growth is concentrated in just a few colonies.[32] Although northern gannet populations are now stable, their numbers were once greatly reduced due to loss of habitat, removal of eggs and killing of adults for their meat and feathers. In 1939, there were 22 colonies and some 83 000 nests, which means that the populations have increased fourfold since that time.[24] This increase in numbers could also be due to northern gannets benefiting from the growing activities of deep sea fishing.
In 1992, the International Union for the Conservation of Nature (IUCN) estimated the bird’s population to be some 526 000.[33] However, taking into account an estimate produced for BirdLife International in 2004 of the European population, the IUCN revised its global population to between 950 000 and 1 200 000 individuals.[34]
A count was undertaken on the Bass Rock population in 2014 - this was an aerial count and confirmed the Bass Rock as the world's largest colony with over 150,000 pairs.

Conservation status


Nests among the rocks. The population of this species appears to be increasing.
The IUCN lists northern gannets as a species of least concern, as they are widely distributed, do not fulfil many of the criteria set for vulnerable species, and as there is a large population that appears to be growing.[1]
In the United Kingdom, gannets are a protected species. However, a legal exception is made for the inhabitants of the district of Ness (also known as Nis) of the Isle of Lewis in Scotland's Outer Hebrides, who are allowed to cull up to 2,000 gannets (locally known as guga) annually to serve as a traditional local delicacy—the taste is described as fishy.[35] Many of these gannets are taken from Sula Sgeir, which is itself named after them.

Depredation

This species is not heavily predated. The only known habitual natural predators of adults are the bald and white-tailed eagles.[36]
Predators of eggs and nestlings include the great black-backed gull and American herring gull, common ravens, ermine, and red fox. Predation at sea is insignificant though large sharks and seals may rarely snatch a gannet out at sea.[36]

Diet and foraging

Northern gannets forage for food during the day, generally by diving into the sea. They search for food both near to their nesting sites but also further out to sea. Birds that are feeding young have been recorded searching for food up to 320 km (200 mi) from their nest. It has been found that 2% of birds nesting in the colony on Bass Rock search for fish at Dogger Bank, between 280 and 320 km (170 and 200 mi) away. It is likely that they fly greater distances than this while searching for food, possibly up to double this distance; however, they normally fly less than 150 km (93 mi).[37]

Diving

When feeding, these birds are spectacular high-speed divers. They can locate their prey from heights of up to 45 m (148 ft), but they normally search from a height of between 10 and 20 m (33 and 66 ft).[38] When they see a fish they will dive into the water. They dive with their bodies straight and rigid, wings tucked close to the body but reaching back, extending beyond the tail, before piercing the water like an arrow. They control the direction of the dive using their wings.[39] Just as it is going to hit the water a bird will fold its wings against its body. A bird’s head and neck are stretched out in front of the body and the beak is shut.[39] Birds can hit the water at speeds of up to 100 km/h (62 mph). This allows them to penetrate 3–5 m (10–16 ft) below the surface, and occasionally they will swim down to 12–15 m (40–50 ft).

Northern gannet searching for fish
They usually push their prey deeper into the water and capture it as they return to the surface. When a dive is successful, gannets swallow the fish underwater before surfacing, and never fly with the fish in their bill. Larger fish are swallowed headfirst, smaller fish are swallowed sideways or tail first. The bird’s subcutaneous air bags aid their rapid return to the surface.
Their white colour helps other gannets to identify one of their kind and they can deduce the presence of a shoal of fish by this diving behaviour; this in turn facilitates group foraging, which makes capturing their prey easier.[40] Northern gannets also forage for fish while swimming with their head under water.
Some studies have found that the duration and direction of flights made while foraging for food are similar for both sexes. However, there are significant differences in the search behaviour of males and females. Female northern gannets are not only more selective than males in choosing a search area: they also make longer and deeper dives and spend more time floating on the surface than males.[41]

Diet

They eat mainly fish 2.5–30.5 cm (0.98–12.01 in) in length which shoal near the surface. Virtually any small fish (roughly 80–90% of their diet) or other small pelagic species (largely squid) will be taken opportunistically. Sardines, anchovies, haddock, smelt, Atlantic cod and other shoal-forming species are eaten.[42][43] In the case of the larger fish species northern gannets will only eat the young fish.
They will also follow fishing boats with the hope of finding food in the same way as gulls do. They fly around the boats to take fish from the fishing nets or pick up the remains thrown into the sea.[44]

Reproduction

The oldest birds are the first to return to the breeding colonies. The exact duration of the breeding season depends on the colony’s geographic location: the breeding season on Bass Rock starts in the middle of January, that of Iceland at the end of March or in April.[45] The birds that are not of breeding age arrive a few weeks later. In general, birds first return to a colony (not necessarily the one they were born in) when they are two or three years old.[46] It is not unusual for birds to change colony before they reach breeding age, but once an individual has successfully bred in a colony it will not change to another.[47]
Immature birds stay on the edges of the colony. They may even make a nest but they will not breed until they are four or five years old.[45] Some birds of this age will occupy empty nests that they will aggressively defend if they have sat on them for two or three days. If an apparently empty nest has an owner the immature bird will abandon it without putting up a struggle when the owner arrives to claim the nest.[48]

Nest construction


Northern gannet transporting material for its nest.\
The preferred nesting sites are on coastal hillsides or cliffs. If these sites are not available northern gannets will nest in groups on islands or flat surfaces. As they find it more difficult to take off from these locations they will often cross the area occupied by an adjacent nest causing an aggressive reaction from the pair occupying that nest; this means that the stress levels are higher in this type of colony than in those on more vertical surfaces. Notwithstanding this, nests are always built close together and ideal nesting sites will not be used if they are some distance from a colony.[49] On average there are 2.3 nests per square metre.[50]
Nests are made from seaweed, plants, earth and all types of object that float on the sea. The males usually collect the materials. Nests measure between 50 and 70 cm (20 and 28 in) in diameter and are some 30 cm (12 in) in height; during the course of a breeding season they will sustain damage from the wind and other causes and they require frequent maintenance. The area which a nest occupies grows throughout the breeding season as the breeding pairs throw their excrement outside the nest.

Aggressive behaviour on the nest


Female will not react if a male approaches her nest, but she will defend it fiercely if another female approaches
Northern gannets exhibit many types of aggressive behaviour while they are nesting. Confrontations normally only take place between birds of the same sex. Females will lower their heads before an aggressive male that is defending its nest: this will expose the back of the female’s neck and the male will take it in its beak and expel the female from the nest. A female will not react if a male approaches a nest but it will react fiercely if another female approaches.[51] The fights between males that occupy nests for the first time are particularly intense. Such fights can lead to serious injuries. The fights are preceded by threatening gestures, which are also seen outside the breeding season. Males will demonstrate ownership of a nest by gesturing towards their neighbours with their head with the beak pointing down and the wings slightly outstretched.[52]

Mating


"Billing", a mutual greeting gesture[53]

Mating
Once males have found a place to breed they try to attract an available female. The females will fly over the colony a number of times before landing. Their posture, with the neck stretched out, tells the male that they are available for courtship. The male will then shake their heads in a similar way to when they are guarding their nest but with their wings closed.
Gannet pairs are monogamous and may remain together over several seasons, if not for all their lives. The pairs separate when their chicks leave the nest but they pair up again the following year. Should one of the pair die the other bird will leave the breeding ground and pair up with another single bird.
They fiercely defend the area around their nest. Where space allows, the distance between nests is double the reach of an individual.

Eggs and chicks

Northern gannets only lay one egg that on average weighs 104.5 grams.[54] This is lighter than for other seabirds.[55] Where two eggs are found in a nest this is the result of two females laying an egg in the same nest or one of the eggs has been stolen from another nest. Northern gannets will lay another egg if the first one is lost. Incubation takes 42 to 46 days. During incubation the egg is surrounded by the brooding bird’s webbed feet that are flooded with warming blood. The process of breaking the eggshell can take up to 36 hours. When this is about to take place the brooding bird will release the egg from its webbed feet to prevent the egg from breaking under the adult's weight as the chick breaks it open. This is a frequent cause of death for chicks of birds that are breeding for the first time.[56] The webbed feet are also used to cover the chicks, which are only rarely left alone by their parents. Chicks that are left unattended are often attacked and killed by other northern gannets.[57]
Young chicks are fed regurgitated semi-digested fish by their parents. Older chicks receive whole fish. Unlike the chicks of other species, northern gannet chicks do not move about the nest or flap their wings to ask for food: this reduces the likelihood that they will fall from the nest.[58]
The adults feed their offspring for 11 or 12 weeks, until they are strong enough to leave the nest for good. A chick will glide from the nest down to the sea after 75 days, which will mark the point at which it separates permanently from its parents.[46] Young birds that weigh 4 kg (8.8 lb) at this point will not be able to fly so they are unable to return to the cliff. Their fat reserves allow them to pass two or three weeks without eating. If the young birds leave the nest in bad weather they can be mortally wounded as they can be blown against the rocks.[57]
The young birds are attacked by adults if they enter the breeding ground, so they stay at sea learning to fish and fly. A high proportion of the young birds can die if storms occur at this time.[59]

Migration

The young birds migrate southwards for great distances and have even been recorded as far south as Ecuador. In their second year a number of birds return to the colony they were born in, where they arrive after the mature birds, they will then migrate south again at the end of the breeding season. They travel shorter distances in this second migration.
After the breeding season the adults spread out over a wide area although they travel no more than 800 to 1,600 km (500 to 990 mi) from the breeding colony. It is not known if all birds from one colony migrate to the same over-wintering area. Many adults migrate to the west of the Mediterranean, passing over the Strait of Gibraltar and flying over land as little as possible. Other birds follow Africa's Atlantic coastline to arrive in the Gulf of Guinea. Immature northern gannets from colonies in Canada fly to the Gulf of Mexico while the adults do not fly that far.[37]

Gallery