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''Ludodactylus'' (meaning "play finger") is a
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 ...
of anhanguerid
pterodactyloid Pterodactyloidea (derived from the Greek words ''πτερόν'' (''pterón'', for usual ''ptéryx'') "wing", and ''δάκτυλος'' (''dáktylos'') "finger" meaning "winged finger", "wing-finger" or "finger-wing") is one of the two traditional ...
pterosaur Pterosaurs (; from Greek ''pteron'' and ''sauros'', meaning "wing lizard") is an extinct clade of flying reptiles in the order, Pterosauria. They existed during most of the Mesozoic: from the Late Triassic to the end of the Cretaceous (228 ...
from the
Early Cretaceous The Early Cretaceous (geochronology, geochronological name) or the Lower Cretaceous (chronostratigraphy, chronostratigraphic name), is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 145& ...
period (
Aptian The Aptian is an age in the geologic timescale or a stage in the stratigraphic column. It is a subdivision of the Early or Lower Cretaceous Epoch or Series and encompasses the time from 121.4 ± 1.0 Ma to 113.0 ± 1.0 Ma (million years ag ...
stage) of what is now the Crato Formation of the Araripe Basin in
Ceará Ceará (, pronounced locally as or ) is one of the 26 states of Brazil, located in the northeastern part of the country, on the Atlantic coast. It is the eighth-largest Brazilian State by population and the 17th by area. It is also one of the ...
,
Brazil Brazil ( pt, Brasil; ), officially the Federative Republic of Brazil (Portuguese: ), is the largest country in both South America and Latin America. At and with over 217 million people, Brazil is the world's fifth-largest country by area ...
. The type and only species is ''L. sibbicki''. The generic name ''Ludodactylus'' refers to the fact that the animal had the combination of teeth and a ''
Pteranodon ''Pteranodon'' (); from Ancient Greek (''pteron'', "wing") and (''anodon'', "toothless") is a genus of pterosaur that included some of the largest known flying reptiles, with ''P. longiceps'' having a wingspan of . They lived during the late Cr ...
''-like head crest, similar to many toy pterosaurs, and no such creature was known to exist until the discovery of ''Ludodactylus''. However, ''Ludodactylus'' is not the only pterosaur known to possess these features, its very close relative ''
Caulkicephalus ''Caulkicephalus'' is a genus of anhanguerid pterosaur from the Isle of Wight off the coast of England. It lived during the Early Cretaceous period, about 130 million years ago. Discovery and naming Between 1995 and 2003, bone fragments of an ...
'' is another example.


Etymology

The genus was named by Eberhard Frey ''et al.'' in 2003 and contains one known
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriat ...
, ''Ludodactylus sibbicki''. The name is derived from
Latin Latin (, or , ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through ...
''ludus'', "play" and Greek ''daktylos'', "finger". ''Ludus'' refers to the fact, long lamented by paleontologists, that many toy pterosaurs combined
teeth A tooth ( : teeth) is a hard, calcified structure found in the jaws (or mouths) of many vertebrates and used to break down food. Some animals, particularly carnivores and omnivores, also use teeth to help with capturing or wounding prey, ...
with a ''
Pteranodon ''Pteranodon'' (); from Ancient Greek (''pteron'', "wing") and (''anodon'', "toothless") is a genus of pterosaur that included some of the largest known flying reptiles, with ''P. longiceps'' having a wingspan of . They lived during the late Cr ...
''-like head crest, while no such creature was known to exist — however, ''Ludodactylus'' shows exactly this combination of features. "Dactylus", in reference to the characteristic long wing finger, has been a common element in the names of pterosaurs, since the first known genus was named ''
Pterodactylus ''Pterodactylus'' (from Greek () meaning 'winged finger') is an extinct genus of pterosaurs. It is thought to contain only a single species, ''Pterodactylus antiquus'', which was the first pterosaur to be named and identified as a flying rept ...
''. The specific name "''sibbicki''" is an homage to the paleoartist
John Sibbick John Sibbick is a British freelance illustrator and paleoartist best known for his fantasy art and reconstructions of prehistoric life in several media. Sibbick studied Graphics and Illustration at Guildford School of Art (now University for ...
.


Description

''Ludodactylus'' is based on
holotype A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of seve ...
SMNK PAL 3828, a skull missing part of the head crest, that was removed from the plate before the fossil was illegally sold. Unlike other
ornithocheirid Ornithocheiridae (or ornithocheirids, meaning "bird hands") is a group of pterosaurs within the suborder Pterodactyloidea. These pterosaurs were among the last to possess teeth. Members that belong to this group lived from the Early to Late Cre ...
s (the group of which ''Ludodactylus'' was originally assigned to), it had no
premaxilla The premaxilla (or praemaxilla) is one of a pair of small cranial bones at the very tip of the upper jaw of many animals, usually, but not always, bearing teeth. In humans, they are fused with the maxilla. The "premaxilla" of therian mammal has ...
ry crest on the snout, but did have a crest at the back of the skull. Frey ''et al.'' interpreted the deep mandible as a crest on the lower jaws. Trapped between the rami of the mandible is a
yucca ''Yucca'' is a genus of perennial shrubs and trees in the family Asparagaceae, subfamily Agavoideae. Its 40–50 species are notable for their rosettes of evergreen, tough, sword-shaped leaves and large terminal panicles of white or whitis ...
leaf; Frey suggested that the animal got it caught in its
beak The beak, bill, or rostrum is an external anatomical structure found mostly in birds, but also in turtles, non-avian dinosaurs and a few mammals. A beak is used for eating, preening, manipulating objects, killing prey, fighting, probing for fo ...
and unsuccessfully tried to dislodge it (the edge of the leaf is frayed), and then possibly died from
starvation Starvation is a severe deficiency in caloric energy intake, below the level needed to maintain an organism's life. It is the most extreme form of malnutrition. In humans, prolonged starvation can cause permanent organ damage and eventually, de ...
or a complication of starving. The skull would have been more than 66 centimeters (26 in) long.Frey, E., Martill, D., and Buchy, M. (2003). A new crested ornithocheirid from the Lower Cretaceous of northeastern Brazil and the unusual death of an unusual pterosaur. In: Buffetaut, E., and Mazin, J.-M. (eds.). ''Evolution and Palaeobiology of Pterosaurs''. ''Geological Society Special Publication'' 217:56–63. .


Classification

In 2003, Frey ''et al.'' classified ''Ludodactylus'' in the family Ornithocheiridae. In 2007, Frey had reconsidered the validity of ''Ludodactylus'', suggesting that it may represent the same animal as ''
Brasileodactylus ''Brasileodactylus'' a genus of pterosaur from the Aptian-age (Early Cretaceous period) lower Santana formation of Chapada do Araripe, Ceará, Brazil. The genus was named by paleontologist Alexander Wilhelm Armin Kellner in 1984. The genus nam ...
'', and therefore a
junior synonym The Botanical and Zoological Codes of nomenclature treat the concept of synonymy differently. * In botanical nomenclature, a synonym is a scientific name that applies to a taxon that (now) goes by a different scientific name. For example, Linn ...
. In 2014 however, Brian Andres and colleagues, in a large
cladistic Cladistics (; ) 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 typically shared derived char ...
analysis of pterosaurs, found ''Ludodactylus'' to be slightly more closely related to ornithocheirids and anhanguerids than to ''Brasileodactylus''. In the analysis of Andres and colleagues, ''Ludodactylus'' is classified just outside Ornithocheiridae and Anhangueridae as a derived member of the more inclusive group Anhangueria. Their cladogram is shown on the left. In 2020, a different topology by Borja Holgado and Rodrigo Pêgas recovered ''Ludodactylus'' within the family Anhangueridae, more specifically to the subfamily Anhanguerinae, sister taxon to both ''
Caulkicephalus ''Caulkicephalus'' is a genus of anhanguerid pterosaur from the Isle of Wight off the coast of England. It lived during the Early Cretaceous period, about 130 million years ago. Discovery and naming Between 1995 and 2003, bone fragments of an ...
'' and ''
Guidraco ''Guidraco'' (Chin. ''gui'' (鬼) "malicious ghost" + Lat. ''draco'' "dragon") is an extinct genus of toothed pterodactyloid pterosaur known from the Early Cretaceous of Liaoning Province, northeast China. According to many recent studies, ''Gu ...
''. This proves that it is indeed very closely related to the anhanguerids. Their cladogram is shown on the right. Topology 1: Andres ''et al.'' (2014). Topology 2: Holgado & Pêgas (2020).


See also

* List of pterosaur genera * Timeline of pterosaur research


References

{{Portal bar, Paleontology, Cretaceous, Brazil Pteranodontoids Aptian life Early Cretaceous pterosaurs of South America Cretaceous Brazil Fossils of Brazil Crato Formation Fossil taxa described in 2003