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''Anatalavis'' is
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
of prehistoric
bird Birds are a group of warm-blooded vertebrates constituting the class (biology), class Aves (), characterised by feathers, toothless beaked jaws, the Oviparity, laying of Eggshell, hard-shelled eggs, a high Metabolism, metabolic rate, a fou ...
s related to
duck Duck is the common name for numerous species of waterfowl in the family (biology), family Anatidae. Ducks are generally smaller and shorter-necked than swans and goose, geese, which are members of the same family. Divided among several subfam ...
s and geese, perhaps to the magpie-goose (''Anseranas semipalmata'') in particular. Alternatively, it might have been a more basal lineage of Anserimorphae distinct from the living waterfowl, similar or even related to the roughly contemporary '' Conflicto antarcticus'' from the
Danian The Danian is the oldest age or lowest stage of the Paleocene Epoch or Series, of the Paleogene Period or System, and of the Cenozoic Era or Erathem. The beginning of the Danian (and the end of the preceding Maastrichtian) is at the Cretac ...
of
Antarctica Antarctica () is Earth's southernmost and least-populated continent. Situated almost entirely south of the Antarctic Circle and surrounded by the Southern Ocean (also known as the Antarctic Ocean), it contains the geographic South Pole. ...
.Tambussi ''et al.'' 2019


Species

The
type species In International_Code_of_Zoological_Nomenclature, zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the spe ...
''Anatalavis rex'' is known from Hornerstown, New Jersey, and its remains were collected by J.G.Meirs in 1869 and W.Ross in 1878. They are probably from the Hornerstown Formation (
Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the more recent of two epochs into which the Cretaceous Period is divided in the geologic time scale. Rock strata from this epoch form the Upper Cretaceous Series. The Cretaceous is named after ''cre ...
or
Early Paleocene The Danian is the oldest age (geology), age or lowest stage (stratigraphy), stage of the Paleocene Epoch or series (stratigraphy), Series, of the Paleogene Period or system (stratigraphy), System, and of the Cenozoic Era or Erathem. The beginnin ...
, some 66 
million years ago Million years ago, abbreviated as Mya, Myr (megayear) or Ma (megaannum), is a unit of time equal to (i.e. years), or approximately 31.6 teraseconds. Usage Myr is in common use in fields such as Earth science and cosmology. Myr is also used w ...
) – usually, they are assumed to be from the lower
Maastrichtian The Maastrichtian ( ) is, in the International Commission on Stratigraphy (ICS) geologic timescale, the latest age (geology), age (uppermost stage (stratigraphy), stage) of the Late Cretaceous epoch (geology), Epoch or Upper Cretaceous series (s ...
layer of the Hornerstown Formation, but they might be younger and postdate the
Mesozoic The Mesozoic Era is the Era (geology), era of Earth's Geologic time scale, geological history, lasting from about , comprising the Triassic, Jurassic and Cretaceous Period (geology), Periods. It is characterized by the dominance of archosaurian r ...
, or even from the slightly older Navesink Formation, as these deposits are reworked and locally mixed across the K-Pg boundary, and no exact locality data was recorded for the ''A. rex'' fossils. The remains were recognized as a new species by Robert Wilson Shufeldt in the early 20th century already, but for a long time they remained in the putative shorebird genus '' Telmatornis''. As this species was larger than its supposed congeners ''T. priscus'' and ''T. affinis'' (which are today considered a single species), Shufeldt gave it the
species name In taxonomy, binomial nomenclature ("two-term naming system"), also called binary nomenclature, is a formal system of naming species of living things by giving each a name composed of two parts, both of which use Latin grammatical forms, altho ...
''rex'',
Latin Latin ( or ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally spoken by the Latins (Italic tribe), Latins in Latium (now known as Lazio), the lower Tiber area aroun ...
for "king". Its affinity to waterfowl was only recognized in 1987 by Storrs L. Olson and David C. Parris, who consequently established the genus ''Anatalavis'', meaning "duck-winged bird", from Latin ''anas'' "duck", ''ala'' "wing" and ''avis'' "bird". ''A. rex'' is only known from two humeri (
holotype A holotype (Latin: ''holotypus'') is a single physical example (or illustration) of an organism used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of s ...
YPM 902 of a right wing, and
paratype In zoology and botany, a paratype is a specimen of an organism that helps define what the scientific name of a species and other taxon actually represents, but it is not the holotype (and in botany is also neither an isotype (biology), isotype ...
YPM 948 of a left wing) which are distinct and characteristic even though they lack the proximal end. They are not particularly large, but quite stout, and the living bird was probably similar to an average-sized dabbling duck of genus '' Anas'' in bulk.As per Olson & Parris (1987) pp.6 & 11, ''A. rex'' was somewhat smaller than '' Graculavus velox'', which in turn was slightly larger than a Beach stone-curlew which weighs about 1 kilogram and is around 55 cm in length. Thus, ''A. rex'' may have been about the size of a Gadwall, Northern Shoveler or Lesser Scaup but shorter-winged and probably quite long-legged, measuring roughly 45-50 cm from billtip to tail and weighing 700-800 grams. Considering the heaviness of its humerus, in size and general shape it was thus probably much similar to a Double-striped Thick-knee, but with a duck's filter-feeding bill. Notably, the shaft of the humerus has a pronounced curve like in living ducks; in the original humerus fossils of ''Telmatornis'' little of the shaft was preserved, but eventually a more complete specimen was discovered, and its straight shaft is quite unlike that of ''Anatalavis''.Shufeldt (1915)Olson & Parris (1987) Some 80 years after ''A. rex'', a second
species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
''Anatalavis oxfordi'' was described by Olson, based on
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserve ...
s found in the earliest
Eocene The Eocene ( ) is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (Ma). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene'' comes ...
(
Ypresian In the geologic timescale the Ypresian is the oldest age (geology), age or lowest stage (stratigraphy), stratigraphic stage of the Eocene. It spans the time between , is preceded by the Thanetian Age (part of the Paleocene) and is followed by th ...
age, about 55 mya)
London Clay The London Clay Formation is a Sediment#Shores and shallow seas, marine formation (geology), geological formation of Ypresian (early Eocene Epoch, c. 54-50 million years ago) age which outcrop, crops out in the southeast of England. The London C ...
Bed A at Walton-on-the-Naze,
England England is a Countries of the United Kingdom, country that is part of the United Kingdom. It is located on the island of Great Britain, of which it covers about 62%, and List of islands of England, more than 100 smaller adjacent islands. It ...
, and named in honor of the collector Andrew Oxford of Great Mongeham, Kent, who donated the fossil for scientific research. This species is known from a partial skeleton, one of the best-preserved London Clay fossils at the time it was found, but in a delicate state of preservation and much broken. Even so, the close similarity of its humerus to the remains of the ''A. rex'' was noted. Olson at that time also erected the
subfamily In biological classification, a subfamily (Latin: ', plural ') is an auxiliary (intermediate) taxonomic rank, next below family but more inclusive than genus. Standard nomenclature rules end botanical subfamily names with "-oideae", and zo ...
Anatalavinae for the genus. Jiří Mlíkovský in 2002 disagreed with uniting ''A. oxfordi'' with the considerably older ''A. rex'' and pointed out that the distal end of the humerus seems to be set at a slightly different angle relative to the bone's shaft in the two species. He thus established the genus ''Nettapterornis'' – with the same meaning as ''Anatalavis'' but in
Ancient Greek Ancient Greek (, ; ) includes the forms of the Greek language used in ancient Greece and the classical antiquity, ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following periods: Mycenaean Greek (), Greek ...
, from ''netta'' (νεττα), ''pteron'' (πτερον) and ''ornis'' (ορνις) – for ''A. oxfordi'', but this remains controversial.Olson (1999)Mlíkovský (2002): pp.107-108


Systematics

A preliminary
cladistic Cladistics ( ; from Ancient Greek 'branch') is an approach to biological classification in which organisms are categorized in groups ("clades") based on hypotheses of most recent common ancestry. The evidence for hypothesized relationships is ...
analysis in 2001Dyke (2001) resolved ''A. oxfordi'' as a member of the group of living waterfowl, and indeed more advanced than the magpie-goose, while still retaining more ancestral traits than the slightly younger '' Presbyornis pervetus''; the latter species, however, was the only fossil with sufficient remains for a quantitative comparison at that time. The subsequent discovery of ''Conflicto antarcticus'', described in 2019 and also known from a wide range of fossilized bones – thus well-comparable to ''A. oxfordi'' –, only made ''Anatalavis'' more enigmatic, as it proved that archaic waterfowl whose beak was already more like that of modern ducks than that of the magpie-goose were already present soon after the end of the
Mesozoic The Mesozoic Era is the Era (geology), era of Earth's Geologic time scale, geological history, lasting from about , comprising the Triassic, Jurassic and Cretaceous Period (geology), Periods. It is characterized by the dominance of archosaurian r ...
. A more comprehensive cladistic analysis conducted at that time, comparing a wide range of living and fossil waterfowl and their closest relatives, found ''C. antarcticus'' and ''A. oxfordi'' to be possibly (but with very low confidence) very closely related and together forming one distinct lineage outside the group of living waterfowl (including the magpie-goose). Thus, whether ''Anatalavis'' was indeed a member of the modern waterfowl, or belongs to a basal lineage – possibly family Conflictonidae – that evolved a filter-feeding beak in parallel to today's ducks and geese, is as unresolved as ever. The hypothesis that it is even closer to living ducks than the magpie-goose, however, seems to be an artifact of the small sample of
taxa In biology, a taxon (back-formation from ''taxonomy''; : taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular name and ...
analyzed by Dyke in 1999: the 2019 analysis rejects it with some confidence, even though it cannot exclude it at a high level of certainty.


Footnotes


References

* (2000). The fossil waterfowl (Aves: Anseriformes) from the Tertiary of England. '' J. Vertebr. Paleontol.'' 20(Supplement 3): 39A. * (2001). The fossil waterfowl (Aves: Anseriformes) from the Eocene of England. '' Am. Mus. Novit.'' 3354: 1-15.
Fulltext
* (2002). ''Cenozoic Birds of the World, Part 1: Europe'']. Ninox Press, Prague
PDF fulltext
!-- This should be treated with extreme caution as regards merging of species. Splits are usually good though. See also critical review in Auk121:623–627 here http://www.findarticles.com/p/articles/mi_qa3793/is_200404/ai_n9396879 --> * (1999). The Anseriform Relationships of ''Anatalavis'' Olson and Parris (Anseranatidae), with a New Species from the Lower Eocene London Clay. ''In:'' (ed.): ''Avian Paleontology at the Close of the 20th Century. Proceedings of the 4th International Meeting of the Society of Avian Paleontology and Evolution, Washington, DC, 4–7 June 1996. Smithsonian Contributions and Studies Series, Smithsonian Contrib. Paleobiol.'' 89: 231–243
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* (1987). The Cretaceous Birds of New Jersey. '' Smithsonian Contributions and Studies Series, Smithsonian Contrib. Paleobiol.'' 63: 1-22
Fulltext
* (1915). Fossil Birds in the Marsh Collection of Yale University. '' Trans. Connectic. Acad. Arts Sci.'' 19: 1-110
PDF fulltext
* (2019). A stem anseriform from the early Palaeocene of Antarctica provides new key evidence in the early evolution of waterfowl. '' Zool. J. Linn. Soc.'' 186(3): 673–700. Bird genera Anseriformes Anseranatidae Paleogene birds Prehistoric birds of North America Fossil taxa described in 1915 Fossil taxa described in 1987 Fossil taxa described in 1999 Fossil taxa described in 2002 {{Anseriformes-stub