Ludodactylus
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''Ludodactylus'' (meaning "play finger") is a
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
of
anhanguerid Anhangueridae is a group of pterosaurs within the suborder Pterodactyloidea. They were among the last pterosaurs to possess teeth. A recent study discussing the group considered the Anhangueridae to be typified by a premaxillary crest and a late ...
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 to ...
from the
Early Cretaceous The Early Cretaceous ( geochronological name) or the Lower Cretaceous (chronostratigraphic name), is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 145  Ma to 100.5 Ma. Geology Pro ...
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 ago), a ...
stage) of what is now the
Crato Formation The Crato Formation is a geologic formation of Early Cretaceous ( Aptian) age in northeastern Brazil's Araripe Basin. It is an important Lagerstätte (undisturbed fossil accumulation) for palaeontologists. The strata were laid down mostly duri ...
of the
Araripe Basin The Araripe Basin () is a rift basin covering about ,Neto et al., 2013, p.1 in Ceará, Piauí and Pernambuco states of northeastern Brazil. It is bounded by the Patos and Pernambuco lineaments, and is situated east of the Parnaíba Basin, southwes ...
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 Eberhard is an old Germanic name meaning the strength or courage of a wild boar. People First name *Eberhard of Friuli (815–866), Duke and key figure in the Carolingian Empire *Eberhard of Béthune (died 1212), Flemish grammarian *Eberhard I, Du ...
''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 appropriate s ...
, ''Ludodactylus sibbicki''. The name is derived from
Latin Latin (, or , ) is a classical language belonging to the 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 the power of the ...
''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, tear ...
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 Paleoart (also spelled palaeoart, paleo-art, or paleo art) is any original artistic work that attempts to depict prehistoric life according to scientific evidence. Works of paleoart may be representations of fossil remains or imagined depiction ...
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 several ...
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 Cr ...
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 b ...
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 whitish flo ...
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 food ...
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, dea ...
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 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 Cr ...
. In 2007, Frey had reconsidered the validity of ''Ludodactylus'', suggesting that it may represent the same animal as '' Brasileodactylus'', 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, Linna ...
. 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
anhanguerid Anhangueridae is a group of pterosaurs within the suborder Pterodactyloidea. They were among the last pterosaurs to possess teeth. A recent study discussing the group considered the Anhangueridae to be typified by a premaxillary crest and a late ...
s 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 Anhangueria (or anhanguerians) is a group of pterosaurs belonging to the clade Pteranodontoidea. Fossil remains of this group date back from the Early to Late Cretaceous periods (Valanginian to Turonian stages), around 140 to 92.5 million years ...
. 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 This list of pterosaurs is a comprehensive listing of all genera that have ever been included in the order Pterosauria, excluding purely vernacular terms. The list includes all commonly accepted genera, but also genera that are now considered inval ...
*
Timeline of pterosaur research This timeline of pterosaur research is a chronologically ordered list of important fossil discoveries, controversies of interpretation, and taxonomic revisions of pterosaurs, the famed flying reptiles of the Mesozoic era. Although pterosaurs w ...


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