Bambolinetta Lignitifila
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''Bambolinetta lignitifila'' is a
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 preserved ...
species of
waterfowl Anseriformes is an order of birds also known as waterfowl that comprises about 180 living species of birds in three families: Anhimidae (three species of screamers), Anseranatidae (the magpie goose), and Anatidae, the largest family, which i ...
from the Late
Miocene The Miocene ( ) is the first epoch (geology), geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and mea ...
of
Italy Italy ( it, Italia ), officially the Italian Republic, ) or the Republic of Italy, is a country in Southern Europe. It is located in the middle of the Mediterranean Sea, and its territory largely coincides with the homonymous geographical ...
, now classified as the sole member of the
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...
''Bambolinetta''. First described in 1884 as a typical dabbling duck, it was not revisited until 2014, when a study showed it to be a highly unusual duck species, probably a flightless, wing-propelled diver similar to a penguin.


History

The species is known from partial skeleton collected in
Montebamboli Montebamboli is a hamlet and località in the comune of Massa Marittima, Tuscany, Italy. The settlement was first mentioned in a parchment from the year 754, and it is located in the hills of what is now the centre of Massa Marittima, within the P ...
,
Tuscany it, Toscano (man) it, Toscana (woman) , population_note = , population_blank1_title = , population_blank1 = , demographics_type1 = Citizenship , demographics1_footnotes = , demographics1_title1 = Italian , demogra ...
, placed in the Museum of Geology and Palaeontology at the
University of Turin The University of Turin (Italian language, Italian: ''Università degli Studi di Torino'', UNITO) is a public university, public research university in the city of Turin, in the Piedmont (Italy), Piedmont region of Italy. It is one of the List ...
. Among the other fossils collected from the same locality are some of the hominid ''
Oreopithecus ''Oreopithecus'' (from the Greek , and , , meaning "hill-ape") is an extinct genus of hominoid primate from the Miocene epoch whose fossils have been found in today's Tuscany and Sardinia in Italy. It existed nine to seven million years ago in ...
''.
Tommaso Salvadori Count Adelardo Tommaso Salvadori Paleotti (30 September 1835 – 9 October 1923) was an Italian zoologist and ornithologist. Biography Salvadori was born in Porto San Giorgio, son of Count Luigi Salvadori and Ethelyn Welby, who was English. His ...
was the first to study these fossils, publishing an account as part of an 1868 paper by B. Gastaldi, in which he pointed to similarities with both waterfowl and
auk An auk or alcid is a bird of the family Alcidae in the order Charadriiformes. The alcid family includes the murres, guillemots, auklets, puffins, and murrelets. The word "auk" is derived from Icelandic ''álka'', from Old Norse ''alka'' (a ...
s. The species was formally described by Alessandro Portis in 1884, with the name ''Anas lignitifila'', a member of the genus ''
Anas ''Anas'' is a genus of dabbling ducks. It includes the pintails, most teals, and the mallard and its close relatives. It formerly included additional species but following the publication of a molecular phylogenetic study in 2009 the genus was s ...
'' that contains common duck species such as the mallard. Portis cited correspondence with Salvadori, who by then was convinced that the fossil was an
anatid The Anatidae are the biological family of water birds that includes ducks, geese, and swans. The family has a cosmopolitan distribution, occurring on all the world's continents except Antarctica. These birds are adapted for swimming, floating ...
; at the time, essentially all fossil ducks were placed in ''Anas''. After Portis, no studies were made and the only mentions of the species in scientific literature were listings in catalogues; most recently,
Jiří Mlíkovský Jiří (; ''YI-RZHEE''), the Czech is a masculine given name, equivalent to English George, may refer to: Given name B * Jiří Antonín Benda * Jiří Baborovský *Jiří Barta *Jiří Bartoška * Jiří Bicek * Jiří Bobok *Jiří Bubla *Jiř ...
placed the species as ''
incertae sedis ' () or ''problematica'' is a term used for a taxonomic group where its broader relationships are unknown or undefined. Alternatively, such groups are frequently referred to as "enigmatic taxa". In the system of open nomenclature, uncertain ...
'' (position uncertain) among birds in 2002. The only illustration of the fossil was the lithograph from Gastaldi's 1868 paper, which remains an important documentation of the fossil since much of the distal part of the wing has been lost. The first reexamination of the species was made in a paper by
Gerald Mayr Gerald Mayr is a German palaeontologist who is Curator of Ornithology at the Senckenberg Research Institute in Frankfurt am Main, Hesse. He has published extensively on fossil birds, especially the Paleogene avifauna of Europe. He is an expert on ...
and Marco Pavia published in 2014. Mayr and Pavia showed the species to have morphological features not present in any other anatid. According to them, it most likely is a highly unusual anatine, falling outside the three main
tribes The term tribe is used in many different contexts to refer to a category of human social group. The predominant worldwide usage of the term in English is in the discipline of anthropology. This definition is contested, in part due to confli ...
(Anatini, Mergini and Aythini). Accordingly, they placed the species in its own genus, taking the name ''Bambolinetta'' from its type locality.


Description

The sole specimen of the species consists of a partial skeleton on a slab, that includes an incomplete skull, most of the trunk, right wing, parts of the
tibiotarsus The tibiotarsus is the large bone between the femur and the tarsometatarsus in the leg of a bird. It is the fusion of the proximal part of the tarsus with the tibia. A similar structure also occurred in the Mesozoic Heterodontosauridae. These s ...
and tarsometatarsus. This species was probably mid-sized for a duck in life, and had arm bones, in particular the humerus, that are much stouter than those of any other anatids. The
ulna The ulna (''pl''. ulnae or ulnas) is a long bone found in the forearm that stretches from the elbow to the smallest finger, and when in anatomical position, is found on the medial side of the forearm. That is, the ulna is on the same side of t ...
is short relative to the humerus and hand, as in auks and penguins. As in the crested auklet, the processus extensorius of the
carpometacarpus The carpometacarpus is a bone found in the hands of birds. It results from the fusion of the carpal and metacarpal bone, and is essentially a single fused bone between the wrist and the knuckles. It is a smallish bone in most birds, generally flat ...
is well-developed and extends less than it does in other diving birds.


Paleobiology

''Bambolinetta'' appears to have had limited, if any, flight capacities. Mayr and Pavia tentatively suggest that the best explanation for its wing anatomy is that it was a wing-propelled diver, similar to penguins and plotopterids. That would make it the only waterfowl species ever to use wings rather than feet for propulsion in water. ''Bambolinetta'' is known from the MN12
European land mammal age The European Land Mammal Mega Zones (abbreviation: ELMMZ, more commonly known as European land mammal ages or ELMA) are zones in rock layers that have a specific assemblage of fossils (biozones) based on occurrences of fossil assemblages of Europe ...
(corresponding to the Middle
Turolian The Turolian age is a period of geologic time (9.0–5.3 Ma) within the Miocene used more specifically with European Land Mammal Ages. It precedes the Ruscinian age and follows the Vallesian age. The Turolian overlaps the Tortonian The Tort ...
period) of the Late Miocene. During this time period, the region that it lived in formed the Tusco-Sardinian island, where in its presumed freshwater habitats, the only predators were
crocodilians Crocodilia (or Crocodylia, both ) is an order of mostly large, predatory, semiaquatic reptiles, known as crocodilians. They first appeared 95 million years ago in the Late Cretaceous period (Cenomanian stage) and are the closest living ...
and otters. The species evolved in the relative absence of terrestrial predators, allowing it to pursue an unorthodox ecological niche. Similar specialisation and loss of flight ability were seen in a number of island waterfowl during the Miocene and Holocene, notably the
moa-nalo The moa-nalo are a group of extinct aberrant, goose-like ducks that lived on the larger Hawaiian Islands, except Hawaii itself, in the Pacific. They were the major herbivores on most of these islands until they became extinct after human settle ...
s of Hawaii, '' Chendytes'' in modern California, and ''
Cnemiornis The New Zealand goose is the common name given to the extinct genus ''Cnemiornis'' of the family Anatidae, subfamily Anserinae. The genus, endemic to New Zealand, consisted of two species: the North Island goose, ''C. gracilis'' and the ...
'' in New Zealand.


References

{{Taxonbar, from=Q18356377 Fossils of Italy Extinct flightless birds Bird genera Miocene birds Fossil taxa described in 1884 Anatinae