Pelagornithidae

[1][2] Most of the common names refer to these birds' most notable trait: tooth-like points on their beak's edges, which, unlike true teeth, contained Volkmann's canals and were outgrowths of the premaxillary and mandibular bones.

They were the dominant seabirds of most oceans throughout most of the Cenozoic, and modern humans apparently missed encountering them only by a tiny measure of evolutionary time: the last known pelagornithids were contemporaries of Homo habilis and the beginning of the history of technology.

The undescribed species provisionally called "Odontoptila inexpectata"[5] – from the Paleocene-Eocene boundary of Morocco – is the smallest pseudotooth bird discovered to date and was just a bit larger than a white-chinned petrel (Procellaria aequinoctialis).

In life, the thin bones and extensive pneumatization enabled the birds to achieve large size while remaining below critical wing loadings.

The sternum had the deep and short shape typical of dynamic soarers, and bony outgrowths at the keel's forward margin securely anchored the furcula.

[7] The legs were proportionally short, the feet probably webbed and the hallux was vestigial or entirely absent; the tarsometatarsi (anklebones) resembled those of albatrosses while the arrangement of the front toes was more like in fulmars.

In February 2009, an almost-complete fossilized skull of a presumed Odontopteryx from around the Chasicoan-Huayquerian boundary c. 9 million years ago (Ma) was unveiled in Lima.

According to paleontologist Mario Urbina, who discovered the specimen, and his colleagues Rodolfo Salas, Ken Campbell and Daniel T. Ksepka, the Ocucaje skull is the best-preserved pelagornithid cranium known as of 2009.

[9] Unlike the true teeth of Mesozoic stem-birds like Archaeopteryx or Ichthyornis, the pseudoteeth of the pelagornithids do not seem to have had serrated or otherwise specialized cutting edges, and were useful to hold prey for swallowing whole rather than to tear bits off it.

Though some reconstructions show pelagornithids as diving birds in the manner of gannets, the thin-walled highly pneumatized bones which must have fractured easily judging from the state of fossil specimens make such a mode of feeding unlikely, if not outright dangerous.

Altogether, the pseudotooth birds would have filled an ecological niche almost identical to that of the larger fish-eating pteranodontian pterosaurs, whose extinction at the end of the Cretaceous may well have paved the way for the highly successful 50-million-year reign of the Pelagornithidae.

Unlike albatrosses today, which avoid the tropical equatorial currents with their doldrums, Pelagornithidae were found in all sorts of climates, and records from around 40 Ma stretch from Belgium through Togo to the Antarctic.

Secondly, pinnipeds are limited to near-shore waters while pseudotooth birds roamed the seas far and wide, like large cetaceans, and like all big carnivores all three groups were K-strategists with moderate to very low population densities.

The bony-toothed birds probably required strong updrafts for takeoff and would have preferred higher sites anyway for this reason, rendering competition with pinniped rookeries quite minimal.

As regards breeding grounds, giant eggshell fragments from the Famara mountains on Lanzarote, Canary Islands, were tentatively attributed to Late Miocene pseudotooth birds.

In that respect the presence of medullary bone in the specimens from Lee Creek Mine in North Carolina, United States, is notable, as among birds this is generally only found in laying females, indicating that the breeding grounds were probably not far away.

At least Pacific islands of volcanic origin would be eroded away in the last millions of years however, obliterating any remains of pelagornithid breeding colonies that might have once existed in the open ocean.

In that respect it may be significant that some lineages of cetaceans, like the primitive dolphins of the Kentriodontidae or the shark-toothed whales, flourished contemporary with the Pelagornithidae and became extinct at about the same time.

Also, the modern diversity of pinniped and cetacean genera evolved largely around the Mio-Pliocene boundary, suggesting that many ecological niches emerged or became vacant.

In addition, whatever caused the Middle Miocene disruption and the Messinian Salinity Crisis did affect the trophic web of Earth's oceans not insignificantly either, and the latter event led to a widespread extinction of seabirds.

From the Middle Miocene or Early Pliocene of the Lee Creek Mine, some remains of pseudotooth birds which probably fell victim to sharks while feeding are known.

Among ocean-going birds in general, the upperside tends to be much darker than the underside (including the underwings) – though some petrels are dark grey all over, a combination of more or less dark grey upperside and white underside and (usually) head is a widespread colouration found in seabirds and may either be plesiomorphic for "higher waterbirds" or, perhaps more likely, be an adaptation to provide camouflage, in particular against being silhouetted against the sky if seen by prey in the sea.

It is notable that at least the primary remiges, and often the other flight feathers too, are typically black in birds – even if the entire remaining plumage is completely white, as in some pelicans or in the Bali starling (Leucopsar rothschildi).

The bony-toothed birds' attachment of the coronoideal part of the external mandible adductor muscles was located at the midline, the rostropterygoid process had a support at its base and the mesethmoid bone had a deep depression for the caudal concha, just as in waterfowl.

The carpometacarpus of both Anseriformes and pseudotooth birds has a prominent pisiform process, which extends from the carpal trochlea far fingerwards along the bone's forward side.

Footbone traits are notoriously prone to selection forces in birds, with convergent evolution known to inhibit or even invalidate cladistic analyses; however, the apparent autapomorphies of the lower arm and hand bones are hard to explain by anything else than an actual relationship.

Such a treatment is unlikely to be completely wrong in either case, as the pseudotooth birds are well distinct from the Presbyornithidae and Scopidae, today generally regarded as the very basal divergences of, respectively, the Anseriformes and the pelican-stork group.

[35] While the authors claim it is beyond the paper's scope, the study describing Protodontopteryx suggests that the proposed pro-galloansere traits might actually be plesiomorphic in relation to Aves.

It also notes "striking" similarities between pelagornithids and Ichthyornis in terms of jaw anatomy, but still classifies them as neognaths due to the well-developed hypotarsal crests, a supratendineal bridge on the distal tibiotarsus and the caudally closed ilioischiadic foramen.

[38] Tentatively, the following genera are recognized:[39] Some other Paleogene (and in one case possibly Late Cretaceous) birds, typically taxa known only from the most fragmentary remains, might also be pelagornithids.

Skull of Pelagornis mauretanicus
Skull of Odontopteryx toliapica (white areas are restored). Unlike in other pseudotooth birds, the "teeth" in this genus were slanted forwards.
Pteranodon skeleton. A toothless Late Cretaceous pterosaur , it was similar to Pelagornis in size and proportions and possibly in feeding habits.
Pelagornis chilensis skeleton seen from below
A generic reconstruction of Osteodontornis
Eurasian teal ( Anas crecca ) skull. As typical for Galloanserae , the palatine bone ("Pa") is not expanded downwards.
Skulls of a northern gannet ( Morus bassanus , top) and various Charadriiformes (below). Note the expansion of the palatine bone visible inside the eye sockets in these Neoaves .
Note also the supraorbital salt gland impressions of the Charadriiformes.