Pterodaustro

{{short description|Genus of ctenochasmatid pterosaur from the Early Cretaceous}}

{{Italic title}}

{{speciesbox

| name = Pterodaustro

| fossil_range = Early Cretaceous, {{Fossil range|105|earliest=Aptian}}

| image = Pterodaustro guinazui.jpg

| image_caption = Cast of a fossil specimen at the Museo Argentino de Ciencias Naturales in Caballito, Buenos Aires, Argentina

| display_parents = 3

| genus = Pterodaustro

| parent_authority = Bonaparte, 1970

| species = guinazui

| authority = Bonaparte, 1970

| synonyms =

  • Puntanipterus?
    Bonaparte & Sánchez, 1975L. Codorniú and Z. Gasparini. 2007. Pterosauria. In Z. Gasparini, L. Salgado, R. A. Coria (eds.), Patagonian Mesozoic Reptiles. Indiana University Press, Bloomington, IN 143-166.

}}

Pterodaustro (from Greek {{lang|el|pteron}}, {{gloss|wing}} and Latin {{lang|la|auster}}, {{gloss|south}}{{cite journal | author = Bonaparte, J. F. | year = 1970 | title = Pterodaustro guinazui gen. et sp. nov. Pterosaurio de la Formacion Lagarcito, Provincia de San Luis, Argentina y su significado en la geologia regional (Pterodactylidae) | journal = Acta Geologica Lilloana | volume = 10 | pages = 209–225}}) is a genus of ctenochasmatid pterodactyloid pterosaur from South America. Its fossil remains dated back to the Early Cretaceous period, about 105 million years ago.

Discovery and naming

File:Pterodaustro.jpg

The first fossils, among them the holotype PVL 2571, a thigh bone, were discovered during the late 1960s by José Bonaparte in the Lagarcito Formation, situated in the San Luis Province of Argentina, and dating from the Albian. The genus was subsequently reported in Chile from the Quebrada La Carreta locality, in the Sierra da Candeleros, Segunda Región de Antofagsta, but this turned out to be erroneous; the fossils belong another pterosaur, the dsungaripterid Domeykodactylus ceciliae.{{cite journal |last1=Martill |first1=David M |last2=Frey |first2=Eberhard |last3=Diaz |first3=Guillermo Chong |last4=Bell |first4=CM |title=Reinterpretation of a Chilean pterosaur and the occurrence of Dsungaripteridae in South America |journal=Geological Magazine |date=2000 |volume=137 |issue=1 |pages=19-25 |url=https://pubs.geoscienceworld.org/cup/geolmag/article-abstract/137/1/19/65008/Reinterpretation-of-a-Chilean-pterosaur-and-the?redirectedFrom=fulltext}} At the Argentine site, the just {{convert|50|m2|sqft|sp=us}} large "Loma del Pterodaustro",{{cite journal |last1=Chiappe |first1=Luis |last2=Rivarola |first2=David |last3=Cione |first3=Alberto |last4=Fregenal-Martínez |first4=Marian |last5=Sozzi |first5=Héctor |last6=Buatois |first6=Luis |last7=Gallego |first7=Oscar |last8=Laza |first8=José |last9=Romero |first9=Edgardo |last10=López-Arbarello |first10=Adriana |last11=Buscalioni |first11=Angela |last12=Marsicano |first12=Claudia |last13=Adamonis |first13=Susana |last14=Ortega |first14=Francisco |last15=McGehee |first15=Sherri |last16=Di Iorio |first16=Osvaldo |title=Biotic association and palaeoenvironmental reconstruction of the “Loma del Pterodaustro” fossil site (Early Cretaceous, Argentina) |journal=Geobios |date=1 January 1998 |volume=31 |issue=3 |pages=349–369 |doi=10.1016/S0016-6995(98)80018-1 |url=https://doi.org/10.1016/S0016-6995(98)80018-1 |issn=0016-6995}} since then, during several expeditions, over 750 Pterodaustro specimens have been collected, 288 of them having been catalogued until 2008. This makes the species one of the best known pterosaurs, with examples from all growth stages, from egg to adult.

The genus was named in 1969 by José Bonaparte as an as yet undescribed nomen nudum. The first description followed in 1970, making the name valid, the type species being Pterodaustro guiñazui.{{cite journal | author = Bonaparte, J. F. | year = 1970 | title = Pterodaustro guinazui gen. et sp. nov. Pterosaurio de la Formacion Lagarcito, Provincia de San Luis, Argentina y su significado en la geologia regional (Pterodactylidae) | journal = Acta Geologica Lilloana | volume = 10 | pages = 209–225 }} The generic name is derived from Greek pteron, "wing" and Latin auster, "south (wind)". The elements are combined as a condensed pteron de austro, "wing from the south". The specific name honors paleontologist Román Guiñazú. It was amended in 1978 by Peter Wellnhofer into guinazui, because diacritical signs such as the tilde are not allowed in specific names.

Description

File:Pterodaustro skull.jpg

Pterodaustro has a very elongated skull, up to {{convert|29|cm|in|sp=us}} long. The portion in front of the eye sockets comprises 85 percent of skull length. The long snout and lower jaws curve strongly upwards; the tangent at the point of the snout is perpendicular to that of the jaw joint. Pterodaustro has about a thousand bristle-like modified teeth in its lower jaws that might have been used to strain crustaceans, plankton, algae, and other small creatures from the water.{{cite book |last=Wellnhofer |first=Peter |author-link=Peter Wellnhofer |title=The Illustrated Encyclopedia of Pterosaurs |orig-year=1991 |year=1996 |publisher=Barnes and Noble Books |location=New York |isbn=0-7607-0154-7 |page=132 }} These teeth stand for the most part not in separate alveoli but in two long grooves parallel to the edges of the jaw. They have a length of {{convert|3|cm|ft|sp=us}} and are oval in cross-section, with a width of just {{convert|0.2|to|0.3|mm|in|sp=us}}. At first it was suspected these structures were not true teeth at all, but later research established they were built like normal teeth, including enamel, dentine and a pulp. Despite being made of very hard material, they might still have been flexible to some extent due to their extreme length-width ratio, a bend of up to 45 degrees being possible.{{cite journal |author=John D. Currey |year=1999 |title=The design of mineralised hard tissues for their mechanical functions |journal=Journal of Experimental Biology |volume=202 |issue=23 |pages=3285–3294 |doi=10.1242/jeb.202.23.3285 |pmid=10562511}} The upper jaws also carried teeth, but these were very small with a flat conical base and a spatula-formed crown. These teeth also do not have separate tooth sockets but were apparently held by ligaments in a special tooth pad, that was also covered with small ossicles, or bone plates. It appears that they were not replaced, unlike the teeth of most other reptiles.Ignacio A Cerda, Laura Codorniú, Palaeohistology reveals an unusual periodontium and tooth implantation in a filter-feeding pterodactyloid pterosaur, Pterodaustro guinazui, from the Lower Cretaceous of Argentina, 2023 Oct;243(4):579-589. doi: 10.1111/joa.13878. Epub 2023 Apr 14.

The back of the skull was also rather elongated and in a low position; there are some indications for a low parietal crest.

File:Pterodaustro Size Comparison.svg

Pterodaustro had a maximum adult wingspan of approximately {{cvt|3|m|ft|sp=us}} and a maximum body mass of approximately {{cvt|9.2|kg|lb}}.{{cite journal |last1=Naish |first1=Darren |author1-link=Darren_Naish|last2=Witton |first2=Mark P. |author2-link=Mark_P._Witton|last3=Martin-Silverstone |first3=Elizabeth |title=Powered flight in hatchling pterosaurs: evidence from wing form and bone strength |journal=Scientific Reports |date=22 July 2021 |volume=11 |issue=1 |pages=13130 |doi=10.1038/s41598-021-92499-z |pmid=34294737 |pmc=8298463 |bibcode=2021NatSR..1113130N }} Its hindlimbs are rather robust and its feet large. Its tail is uniquely elongated for a pterodactyloid, containing twenty-two caudal vertebrae, whereas other members of this group have at most, sixteen.

Paleobiology

=Feeding=

Pterodaustro probably strained food with its tooth comb, a method called "filter feeding", also practised by modern flamingos.{{cite book |editor=Palmer, D.|year=1999 |title= The Marshall Illustrated Encyclopedia of Dinosaurs and Prehistoric Animals|publisher= Marshall Editions|location=London|page= 104|isbn= 1-84028-152-9}} Once it caught its food, Pterodaustro probably mashed it with the small, globular teeth present in its upper jaw. Like other ctenochasmatoids, Pterodaustro has a long torso and proportionally massive and splayed hindfeet, adaptations for swimming.Witton, Mark P. (2013). Pterosaurs: Natural History, Evolution, Anatomy. Princeton University Press. {{ISBN|0691150613}}. A recent study suggested that its ankle facilitated movements required for wading behavior.{{cite journal |last1=Burlot |first1=Romain |last2=Codorniú |first2=Laura |last3=Defend |first3=Lenna |last4=Laurin |first4=Michel |title=The ankle joint of Pterodaustro guinazui. |journal=Acta Palaeontologica Polonica |date=2024 |volume=69 |doi=10.4202/app.01097.2023 |url=https://doi.org/10.4202/app.01097.2023 |language=en |issn=0567-7920|hdl=11336/240037 |hdl-access=free }}

Robert Bakker suggested that, like flamingos, this pterosaur's diet may have resulted in a pink hue.{{cite book |title=Fantastic Facts About Dinosaurs |isbn=978-0-7525-3166-3 |author=Jinny Johnson |publisher=Parragon Book Service |year=2000 |url-access=registration |url=https://archive.org/details/fantasticfactsab0000john }}{{page needed|date=May 2014}}

At least two specimens of Pterodaustro have been found, MIC V263 and MIC V243, with gizzard stones in the stomach cavity, the first ever reported for any pterosaur. These clusters of small stones with angled edges support the idea that Pterodaustro ate mainly small, hard-shelled aquatic crustaceans using filter-feeding. Such invertebrates are abundant in the sediment of the fossil site.Codorniú, L., Chiappe, L.M., Arcucci, A., and Ortiz-Suarez, A. (2009). "First occurrence of gastroliths in Pterosauria (Early Cretaceous, Argentina)". XXIV Jornadas Argentinas de Paleontología de Vertebrados

=Growth and reproduction=

A study of the growth stages of Pterodaustro concluded that juveniles grew relatively fast in their first two years, attaining about half of the adult size. Then they reached sexual maturity, growing at a slower rate for four to five years until there was a determinate growth stop.{{cite journal |author=Chinsamy, A., Codorniú, L., and Chiappe, L. M. |year=2008 |title=Developmental growth patterns of the filter-feeder pterosaur, Pterodaustro guinazui |journal=Biology Letters |volume=4 |issue=3 |pages=282–285 |doi=10.1098/rsbl.2008.0004 |pmc=2610039 |pmid=18308672}}

In 2004 a Pterodaustro embryo in an egg was reported, specimen MHIN-UNSL-GEO-V246. The egg was elongated, {{convert|6|cm|in|sp=us}} long and {{convert|22|mm|in|sp=us}} across, and its mainly flexible shell was covered with a thin layer of calcite, 0.3 millimeters thick.{{cite journal |author=Chiappe, L. M., Codorniú, L., Grellet-Tinner, G. and Rivarola, D. |year=2004 |title=Argentinian unhatched pterosaur fossil |journal=Nature |volume=432 |issue=7017 |pages=571–572 |pmid=15577899 |doi=10.1038/432571a |bibcode=2004Natur.432..571C|s2cid=4396534 |url=http://doc.rero.ch/record/14928/files/PAL_E2071.pdf }} Three-dimensionally preserved eggs were reported in 2014.{{Cite journal | doi = 10.1016/j.gsf.2014.05.002| title = The first pterosaur 3-D egg: Implications for Pterodaustro guinazui nesting strategies, an Albian filter feeder pterosaur from central Argentina| journal = Geoscience Frontiers| volume = 5| issue = 6| page = 759| year = 2014| last1 = Grellet-Tinner | first1 = G. | last2 = Thompson | first2 = M. | last3 = Fiorelli | first3 = L. E. | last4 = Argañaraz | first4 = E. S. | last5 = Codorniú | first5 = L. | last6 = Hechenleitner | first6 = E. M. N. | doi-access = free | hdl = 11336/6990 | hdl-access = free }}

In 2009, possible medullary bone tissue was reported from the largest known femur of Pterodaustro (V 382). Medullary bone tissues are previously only known from modern birds that are laying eggs in reproduction, and some non-avian dinosaurs such as Tyrannosaurus rex, so this record would make it the first reported occurrence of a non-dinosaurian medullary bone.{{cite journal|author1=Chinsamy, A.|author2=Codorniú, L.|author3=Chiappe, L.|year=2009|title=Palaeobiological Implications of the Bone Histology of Pterodaustro guinazui|journal=The Anatomical Record|volume=292|issue=9|pages=1462—1477|doi=10.1002/ar.20990}}

=Comparison with waterfowls=

Comparisons between the scleral rings of Pterodaustro and modern birds and reptiles suggest that it may have been nocturnal and similar in activity patterns to modern anseriform birds that feed at night,{{cite journal|author1=Schmitz, L. |author2=Motani, R. |year=2011 |title=Nocturnality in dinosaurs inferred from scleral ring and orbit morphology |journal=Science |volume=332 |issue=6030 |pages=705–708 |doi=10.1126/science.1200043 |pmid=21493820 |bibcode=2011Sci...332..705S|s2cid=33253407 }} although method of this research is questioned by some researchers.{{Cite journal |last=Hall |first=Margaret I. |last2=Kirk |first2=E. Christopher |last3=Kamilar |first3=Jason M. |last4=Carrano |first4=Matthew T. |date=2011-12-23 |title=Comment on “Nocturnality in Dinosaurs Inferred from Scleral Ring and Orbit Morphology” |url=https://www.science.org/doi/10.1126/science.1208442 |journal=Science |language=en |volume=334 |issue=6063 |pages=1641–1641 |doi=10.1126/science.1208442 |issn=0036-8075}}

Because of its long torso and neck and comparatively short legs, Pterodaustro was unique among pterosaurs in having difficulties to launch. Even with the pterosaurian quadrupedal launching mechanism, it would have required frantic and fairly-low angled take-offs possible only in open areas, much like modern geese and swans.

Phylogeny

Bonaparte in 1970 assigned Pterodaustro to the Pterodactylidae; in 1971 to a Pterodaustriidae. However, from 1996 cladistic studies by Alexander Kellner and David Unwin have shown a position within the family Ctenochasmatidae, together with other filter feeders.

File:Pterodaustro guinazaui.jpg

In 2018, a topology by Longrich, Martill and Andres recovered Pterodaustro within the family Ctenochasmatidae, more precisely within the tribe called Pterodaustrini, in a more basal position than Beipiaopterus and Gegepterus.Longrich, N.R., Martill, D.M., and Andres, B. (2018). [http://journals.plos.org/plosbiology/article?id=10.1371/journal.pbio.2001663 Late Maastrichtian pterosaurs from North Africa and mass extinction of Pterosauria at the Cretaceous-Paleogene boundary.] PLoS Biology, 16(3): e2001663.

{{doi|10.1371/journal.pbio.2001663}}

{{clade| style=font-size:90%;line-height:90%

|label1=Ctenochasmatidae

|1={{clade

|label1=Ctenochasmatinae

|1={{clade

|1=Ctenochasma

|label2=Pterodaustrini

|2={{clade

|1=Pterodaustro

|2={{clade

|1=Beipiaopterus

|2=Gegepterus }} }} }}

|2={{clade

|1=Kepodactylus

|2={{clade

|label1=Moganopterinae

|1={{clade

|1=Moganopterus

|2=Feilongus }}

|2={{clade

|1=Ardeadactylus

|2={{clade

|1=Elanodactylus

|label2=Gnathosaurinae

|2={{clade

|1=Huanhepterus

|2={{clade

|1=Plataleorhynchus

|2=Gnathosaurus

}} }} }} }} }} }} }} }}

See also

References

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