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The family Sulidae comprises the and . Collectively called sulids, they are medium-large coastal that plunge-dive for and similar . The 10 in this family are often considered in older sources, placing all in the Sula. However, Sula (true boobies) and Morus (gannets) can be distinguished via morphological, , and characters. Abbott's booby ( Papasula) is given its own genus, as it stands apart from both in these respects. It appears to be a distinct and ancient lineage, maybe closer to the gannets than to the true boobies.Kennedy, Martyn; Spencer, Hamish G. & Gray, Russell D. (1996): Hop, step and gape: do the social displays of the Pelecaniformes reflect phylogeny? Animal Behaviour 51(2): 273-291. (HTML abstract) Erratum: Animal Behaviour 51(5): 1197. Friesen, V.L.; Anderson, D.J.; Steeves, T.E.; Jones, H. & Schreiber, E.A. (2002): Molecular Support for Species Status of the Nazca Booby ( Sula granti). Auk 119(3): 820–826. English DOI: 10.1642/0004-8038(2002)1190820:MSFSSO2.0.CO;2 PDF fulltext


Description
Sulids measure about in length and have a wingspan around . They have long, narrow, and pointed wings, and a quite long, graduated, and rather lozenge-shaped tail whose outer feathers are shorter than the central ones. Their flight muscles are rather small to allow for the small cross-section required for plunge-diving, as an adaptive trade-off relative to some sacrifice in flight performance. Consequently, they are very , reducing drag, so their bodies are "-shaped" and somewhat flat.Nelson, J. Bryan (2003): Gannets and Boobies. In: (ed.): The Firefly Encyclopedia of Birds: 82–87. Firefly Books, Oxford.

They have stout legs and , with the web connecting all four toes. In some species, the webs are brightly colored and used in courtship displays. The bill is usually conspicuously colored, long, deep at the base, and pointed, with saw-like edges. The curves down slightly at the tip and can be moved upward to accept large prey. To keep water out during plunges, the nostrils enter into the bill rather than opening to the outside directly. The eyes are angled forward, and provide a wider field of than in most other birds.

Their is either all-white (or light brownish or greyish) with dark wingtips and (usually) tail, or at least some dark brown or black above with white underparts; gannets have a yellowish hue to their heads. The face usually has some sort of black markings, typically on the lores. Unlike their relatives (the and ), sulids have a well-developed whose waxy secretions they spread on their feathers for waterproofing and pest control. They their tail feathers irregularly and the of their wings in stages, so that starting at the first moult, they always have some old feathers, some new ones, and some partly grown ones. Moult as a response to periods of stress has been recorded.


Distribution and ecology
The sulids are distributed mainly in and waters, but they, particularly gannets, are found in regions, too. These birds are not truly seabirds like the related Procellariiformes, and usually stay rather close to the coasts, but the abundant colonies of sulids that exist on many islands suggest that they are not infrequently blown away from their home range by , and can wander for long distances in search of a safe place to land if need be.

All feed entirely at sea, mostly on mid-sized and similarly sized marine (e.g. ). Many species feed communally, and some species follow to discarded and . The typical hunting behavior is a dive from midair, taking the bird a 1–2 m under water. If prey manages to escape the diving birds at first, they may give chase using their legs and wings for underwater swimming.

As noted above, the traits of gannets and boobies differ considerably, but the Sulidae as a whole are characterized by several behavioral : Before taking off, they point their bills upwards (gannets) or forward (boobies). After landing again, they point downwards with their bills. In response to a threat, they do not attack, but shake their heads and point their bills towards the intruders.


Reproduction
All sulids breed in . Males examine the colony area in flight and then pick a nest site, which they defend by fighting and territorial displays. Males then advertise to females by a special display and call. Their display behavior is characteristic, though not as diverse as the numerous variations found among the ; it typically includes the male shaking his head. Females search the colony in flight and on foot for a mate. Once they select males, pairs maintain their bonds by preening each other and by frequent copulation. The clutch is typically two . The eggs are unmarked (but may become stained by in the nest), whitish, pale blue, green, or pink, and have a coating that resembles lime. Egg weight ranges from 3.3 to 8.0% of the female's weight. Incubation lasts 42 to 55 days, depending on the species. Both sexes ; like their relatives, they do not have , but their feet become and hot, and the birds place the eggs under the webs. Eggs lost during the first half of incubation are replaced.

At hatching, parents move the eggs and then the hatchlings to the tops of their webs. The young hatch naked, but soon develop white down. They beg by touching the parent's bill and take regurgitated food straight from its . At first, at least one parent is always in attendance of the young; after two weeks, both parents leave the nest unguarded at times while they go fishing. The times for the chicks to fledge and become independent of their parents depend greatly on the food supply. Rarely does more than one chick survive to maturity, except in the ( Sula variegata), which has the biggest clutch (two to four eggs), and less often in the blue-footed booby ( S. nebouxii). by the stronger of two chicks is frequent.


Systematics and evolution
Sulids are related to a number of other birds, which all lack external and a , but have all four toes webbed and a . The closest living relatives of the Sulidae are the Phalacrocoracidae (cormorants and shags) and the (darters). The latter are somewhat intermediate between sulids and cormorants, but (like many cormorants) they are birds in a containing otherwise , and also with sulids but with cormorants in some other respects. Thus, the Sulidae seem to be the oldest and most distinct lineage of those three, which are united in a . Therein, the Sulidae are typically placed simply as a family; sometimes, a superfamily Suloidea is recognized, wherein some of the primitive prehistoric forms (e.g. , , and ) are placed as basal lineages distinct from the living Sulidae. However, the proposed family Pseudosulidae (or Enkurosulidae) is almost certainly invalid. (1985): Section X.G.5.a. Sulidae. In: The Fossil Record of Birds. Avian Biology 8: 203-204. PDF fulltextChristidis, Les & Boles, Walter E. (2008): Systematics and Taxonomy of Australian Birds. CSIRO Publishing, CollingwoodVictoria, Australia. Excerpt at p.100Mayr, Gerald (2009): 7.1.3 Sulidae (Gannets and Boobies). In: Paleogene Fossil Birds: 64-65. Springer-Verlag, Heidelberg & New York. Excerpt at

The Sulae were traditionally included in the in its obsolete circumscription, but , the namesake family of the Pelecaniformes, are actually more closely related to , ibises and spoonbills, the , and the than to the sulids and allies. In recognition of this, the Sulae have been proposed for separation in a new order , which also includes the (Fregatidae), as well as one or more lineages that are entirely today. The IOC World Bird List uses Suliformes as the proposed order name.

Within the family itself, three living genera— Sula (boobies, six species), (Abbott's booby), and Morus (gannets, three species)—are recognized. A 2011 study of multiple genes found Abbott's booby to be basal to all other gannets and boobies, and likely to have diverged from them around 22 million years ago, and the ancestors of the gannets and remaining boobies split around 17 million years ago. The most recent common ancestor of all boobies lived in the late Miocene around 6 million years ago, after which time the boobies steadily diverged. The gannets split more recently, only around 2.5 million years ago.


( Morus bassanus)
Western Europe and North America

( Morus capensis)
West to Southwest African coast
Australasian gannet
( Morus serrator or Sula bassana)
Australia and New Zealand
Abbott's booby
( Papasula abbotti)
Christmas Island
Blue-footed booby
( Sula nebouxii)
Eastern Pacific Ocean from California to the Galápagos Islands south into Peru

( Sula dactylatra)
Tropical oceans between the 30th parallel north and 30th parallel south. In the Indian Ocean it ranges from the coastlines of the Arabian Peninsula and East Africa across to Sumatra and Western Australia

( Sula granti)
Eastern Pacific from the islands in Baja California to the Galápagos Islands and the Isla de la Plata in Ecuador and Malpelo in Colombia

( Sula leucogaster)
Pantropical areas of the Atlantic and Pacific Oceans

( Sula sula)
Most tropical areas of oceans, main exceptions being tropical areas of E Atlantic and SE Pacific, breeding on islands

( Sula variegata)
Coast of South America from Peru to Chile

The record of sulids is quite extensive due to the many / forms that have been recovered, but the lineage of sulids extends back to the , and all things (such as the ) considered, the sulids seem to have diverged from the lineage leading to cormorants and darters around 50 million years ago (Mya), perhaps a bit earlier. The initial evolutionary radiation formed a number of that are now completely extinct, such as the (which, as noted above, might not have been a modern-type sulid) or the bizarre (which had a bill shaped like an 's). The modern genera (like many other living genera of birds) around the -Miocene boundary about 23 Mya. , which lived during that time, seems to have been a primitive booby that still had many with gannets. Like the other Phalacrocoraciformes, the sulids originated probably in the general region of the or western – probably the latter rather than the former, given that their earliest fossils are abundant in , but absent from the well-studied contemporary deposits.

Prehistoric sulids (or suloids) only known from are:

  • (Early Eocene of Messel, Germany) – may belong in Eostega
  • (Late Eocene of Cluj-Manastur, Romania) – may include Masillastega
  • Sulidae gen. et sp. indet. (Thalberg Late Oligocene of Germany) – Empheresula?A distal fragment; larger than Microsula: Göhlich (2003), Mayr (2009: p.65)
  • Sulidae gen. et sp. indet. (Late Oligocene of South Carolina, United States) – Microsula?Some fossils that "do not differ substantially from modern sulid genera"; no further details given: Olson (1985: p.203)
  • (Late Oligocene of Gannat, France – Middle Miocene of Steinheimer Becken, Germany) – including "Sula" arvernensis, "Parasula"C.J.O. Harrison's "Parasula" of 1975 is a of as established by in 1913: Mlíkovský (2002: p.66)
  • (Late Oligocene of South Carolina, United States – Grund Middle Miocene of Austria) – may belong in Morus or Sula, includes "Sula" avita, "S." pygmaea, Enkurosula, "Pseudosula"
  • (Middle Miocene of Credinţa, Romania)
  • (Late Miocene of California)
  • (Early Pliocene? of California)
  • (Pisco Early Pliocene of SC Peru)
  • (middle Pliocene of South Carolina)
  • Sulidae gen. et sp. indet. (Late Pliocene of Valle di Fine, Italy) – Morus?A proximal fragment somewhat similar to a gannet's: Lambrecht (1933: p.286)

For prehistoric of the extant , see the genus articles.

The of Ronzon, France, was variously placed in the genus , in Sula, and after a distinct genus was established for it, in the Phalacrocoracidae. While it is quite likely to belong in the Sulae and may have been an ancient sulid (or suloid), of the three placements explicitly proposed, none seems to be correct.Göhlich, Ursula B. (2003): The avifauna of the Grund Beds (Middle Miocene, Early Badenian, northern Austria). Annalen des Naturhistorischen Museums Wien A 104: 237-249 English. PDF fulltext


Further reading
  • (1933): Familia Sulidae. In: Handbuch der Palaeornithologie: 284-287 German. Gebrüder Bornträger, Berlin.
  • Mlíkovský, Jirí (2007): Taxonomic identity of Eostega lebedinskyi LAMBRECHT, 1929 (Aves) from the middle Eocene of Romania. Annalen des Naturhistorischen Museums in Wien A 109: 19-27 English. PDF fulltext


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