Hesperornitheans Temporal range: Late Cretaceous, | |
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Restored skeleton of Hesperornis regalis | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Clade: | Dinosauria |
Clade: | Saurischia |
Clade: | Theropoda |
Clade: | Avialae |
Clade: | Ornithurae |
Clade: | † Hesperornithes Fürbringer, 1888 |
Subgroups [1] | |
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Synonyms | |
Hesperornithes is an extinct and highly specialized group of aquatic avialans closely related to the ancestors of modern birds. They inhabited both marine and freshwater habitats in the Northern Hemisphere, and include genera such as Hesperornis , Parahesperornis , Baptornis , Enaliornis , and Potamornis , all strong-swimming, predatory divers. Many of the species most specialized for swimming were completely flightless. The largest known hesperornithean, Canadaga arctica , may have reached a maximum adult length of 2.2 metres (7.2 ft). [5]
Hesperornitheans were the only Mesozoic avialans to colonize the oceans. They were wiped out in the Cretaceous–Paleogene extinction event, along with enantiornitheans and all other non-avian dinosaurs, and many other diverse plant and animal groups.
Most of what is known about this group rests on analyses of single species, as few provide sufficiently complete fossils for analysis. Although some of the smaller and more basal species, like those belonging to the subgroups Enaliornithidae and Brodavidae, might have been able to fly, the larger hesperornithids like Hesperornis and Baptornis had only vestigial wings. As in the case of modern foot-propelled diving birds, the femur and metatarsus of these animals were short, whereas the tibia was long. The legs were also set far back on the body, as in loons, grebes or penguins. Hesperornithids must have been powerful swimmers and divers but extremely ungainly on the land, and probably spent little time ashore except to nest. They were rather long-bodied, and measured about 6 feet (180 cm) long. [6]
Some researchers think that on land they had to slide on their bellies and push with their legs; the hip and knee joints were shaped such that these species could not move them dorsoventrally, and in a resting position the feet projected sideways from the body, which would have prevented them from walking upright. [2] The anatomy of their toes suggests that hesperornitheans had lobes of skin for propulsion underwater similar to grebes, rather than being webbed. The dense bones of these animals decreased their buoyancy, making diving easier. [7] However, morphometric comparison with modern diving birds suggests that hesperornitheans share more similarities with diving ducks and cormorants rather than with loons or grebes. [8]
The snout was long, and tipped with a slightly hooked beak. Behind the beak, the jaws were filled with a series of simple, sharp teeth which were set into a longitudinal groove. These probably helped to seize fish, like the serrated beak of mergansers. [9] [10] Unlike modern birds, they retained a joint between the lower jaw bones. It is believed that this allowed them to rotate the back portion of the mandible independently of the front, thus allowing the lower teeth to disengage. [6]
Currently, the hesperornitheans are recognized as a very specialized lineage that is not ancestral to modern birds. Still, their relationship is close enough that they probably diverged from the ancestors of modern birds as late as the earliest Cretaceous.
The earliest known hesperornithean is the Early Cretaceous Enaliornis . The majority of hesperornithean species are known from the Late Cretaceous of North America. Small hesperornithean bones are known from the freshwater deposits of the Late Cretaceous of the Judith River Group as well as the Hell Creek and Lance Formations, and in several Eurasian sites. These species were about the size of a cormorant or a loon.
The clade Hesperornithes was originally named as a subclass of Aves by Furbringer in 1888. [11] However, it was generally ignored in the scientific literature in favor of the order-level name Hesperornithiformes, coined one year later. In 2004, Clarke became the first to define the hesperornithean group in terms of phylogenetics. Clarke defined Hesperornithes as all species closer to Hesperornis regalis than to modern birds, and regarded Hesperornithiformes as a junior synonym, though she did not define the latter name. Clarke also defined the more inclusive group Hesperornithidae as all hesperornitheans closer to Hesperornis than to Baptornis. [4]
Hesperornitheans were originally combined with Ichthyornis in the paraphyletic group "Odontornithes" by Othniel Charles Marsh, in 1873. In 1875, they were separated as Odontolcae. The group was often considered to be related to loons and grebes, [12] or to the Paleognathae (based on perceived similarities in the bony palate). [13] These similarities, however, as the more recently determined fact that the osteons of their bones – at least in Hesperornis – were arranged in a pattern similar to that in Neognathae, [14] are today considered to be due to convergent evolution. [15] [16]
In 2015, a species-level phylogenetic analysis found the following relationships among hesperornitheans. [17]
Hesperornithes |
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Gaviiformes is an order of aquatic birds containing the loons or divers and their closest extinct relatives. Modern gaviiformes are found in many parts of North America and northern Eurasia, though prehistoric species were more widespread.
Ichthyornis is an extinct genus of toothy seabird-like ornithuran from the late Cretaceous period of North America. Its fossil remains are known from the chalks of Alberta, Alabama, Kansas, New Mexico, Saskatchewan, and Texas, in strata that were laid down in the Western Interior Seaway during the Turonian through Campanian ages, about 95–83.5 million years ago. Ichthyornis is a common component of the Niobrara Formation fauna, and numerous specimens have been found.
Hesperornis is a genus of cormorant-like Ornithuran that spanned throughout the Campanian age, and possibly even up to the early Maastrichtian age, of the Late Cretaceous period. One of the lesser-known discoveries of the paleontologist O. C. Marsh in the late 19th century Bone Wars, it was an early find in the history of avian paleontology. Locations for Hesperornis fossils include the Late Cretaceous marine limestones from Kansas and the marine shales from Canada. Nine species are recognised, eight of which have been recovered from rocks in North America and one from Russia.
Odontornithes is an obsolete and disused taxonomic term proposed by Othniel Charles Marsh for birds possessing teeth, notably the genera Hesperornis and Ichthyornis from the Cretaceous deposits of Kansas.
Archaeopterygidae is a group of maniraptoran dinosaurs, known from the latest Jurassic and earliest Cretaceous of Europe. In most current classifications, it contains only the genera Archaeopteryx and Wellnhoferia. As its name suggests, Protarchaeopteryx was also once referred to this group, but most paleontologists now consider it an oviraptorosaur. Other referred genera, like Jurapteryx, Wellnhoferia, and "Proornis", are probably synonymous with Archaeopteryx or do not belong into this group. Jinfengopteryx was originally described as an archaeopterygid, though it was later shown to be a troodontid. A few studies have recovered Anchiornis and Xiaotingia to also be members of the Archaeopterygidae, though most subsequent analyses have failed to arrive at the same result. Uncertainties still exist, however, and it may not be possible to confidently state whether archaeopterygids are more closely related to modern birds or to deinonychosaurs barring new and better specimens of relevant species. Teeth attributable to archaeopterygids are known from the earliest Cretaceous (Berriasian) Cherves-de-Cognac locality and the Angeac-Charente bonebed of France.
Gansus is a genus of aquatic birds that lived during the Aptian age of the Early Cretaceous (Aptian-Albian) period in what are now Gansu and Liaoning provinces, western China. The rock layers from which their fossils have been recovered are dated to 120 million years ago. It was first described in 1984 on the basis of an isolated left leg. It is the oldest-known member of the Ornithurae, the group which includes modern birds (Neornithes) and extinct related groups, such as Ichthyornis and Hesperornithes.
Ichthyornithes is an extinct group of toothed avialans very closely related to the common ancestor of all modern birds. They are known from fossil remains found throughout the late Cretaceous period of North America, though only two genera, Ichthyornis and Janavis, are represented by complete enough fossils to have been named. Ichthyornitheans became extinct at the Cretaceous–Paleogene boundary, along with enantiornitheans, all other non-avian dinosaurs, and many other animal and plant groups.
Baptornis is a genus of flightless, aquatic birds from the Late Cretaceous, some 87-80 million years ago. The fossils of Baptornis advenus, the type species, were discovered in Kansas, which at its time was mostly covered by the Western Interior Seaway, a shallow shelf sea. It is now known to have also occurred in today's Sweden, where the Turgai Strait joined the ancient North Sea; possibly, it occurred in the entire Holarctic.
Polarornis is a genus of prehistoric bird, possibly an anserimorph. It contains a single species Polarornis gregorii, known from incomplete remains of one individual found on Seymour Island, Antarctica, in rocks which are dated to the Late Cretaceous.
Neogaeornis is a controversial prehistoric genus of diving bird. The single known species, Neogaeornis wetzeli, was described from fossils found in the Campanian to Maastrichtian Quiriquina Formation of Chile. It lived about 70-67 million years ago. It remains known from the single tarsometatarsus described in 1929 by Lambrecht, and today housed in the Paläontologisches Institut und Museum in Kiel, Germany.
Parahesperornis is a genus of prehistoric flightless birds from the Late Cretaceous. Its range in space and time may have been extensive, but its remains are rather few and far between, at least compared with its contemporary relatives in Hesperornis. Remains are known from central North America, namely the former shallows of the Western Interior Seaway in Kansas. Found only in the upper Niobrara Chalk, these are from around the Coniacian-Santonian boundary, 85-82 million years ago (mya).
Limenavis is a prehistoric bird genus from the Late Cretaceous. It lived about 70 million years ago, around the Campanian-Maastrichtian boundary. Known from several broken bones, the remains of the only known species Limenavis patagonica were found in rocks of the "lower member" of the Allen Formation at Salitral Moreno, 20 km south of General Roca, Río Negro (Argentina). It is the closest relative, in the fossil record, of the modern birds.
Canadaga is a flightless bird genus from the Late Cretaceous. The single known species is Canadaga arctica. It lived in the shallow seas around what today is Bylot Island in Nunavut, Canada. Its fossils were found in rocks dated to the mid-Maastrichtian age, about 67 million years ago.
Potamornis is a prehistoric bird genus that dated back to the late Maastrichtian age of the late Cretaceous period. Its scrappy remains were found in the Lance Formation at Buck Creek, USA, and additional possible remains were found in the upper Hell Creek Formation of Montana, dated to the Danian age of the Paleogene period, though these may have been reworked. A single species was named and described in 2001: Potamornis skutchi.
Ornithurae is a natural group which includes the common ancestor of Ichthyornis, Hesperornis, and all modern birds as well as all other descendants of that common ancestor.
Chaoyangia is an extinct genus of euornithean birds, containing the single species Chaoyangia beishanensis. This species is known from a single fossil specimen consisting of a partial skeleton including vertebra, ribs, hips, and upper legs. The specimen was discovered in the Jiufotang Formation near the city of Chaoyang in Liaoning province, China. This rock formation has been dated to the Aptian age of the Early Cretaceous period, 120 million years ago.
Vegaviidae is an extinct family of ornithurines, often regarded as stem-anseriforms, which existed during the Late Cretaceous and possibly the Paleocene. Fossils attributed to the family have been found in Canada, Chile, New Zealand, and Antarctica.
Fumicollis is a genus of prehistoric flightless birds from the Late Cretaceous (Coniacian-Santonian) Niobrara Chalk of Kansas.
Chupkaornis is a genus of prehistoric flightless birds from the Late Cretaceous (Coniacian-Santonian) Kashima Formation of Hokkaido, Japan.