Ludodactylus
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''Ludodactylus'' (meaning "play finger") is a
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
anhanguerid Anhangueridae (alternatively called Ornithocheiridae, 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 ...
pterodactyloid Pterodactyloidea ( ; derived from the Greek words ''πτερόν'' (''pterón'', for usual ''ptéryx'') "wing", and ''δάκτυλος'' (''dáktylos'') "finger") is one of the two traditional suborders of pterosaurs ("wing lizards"), and contai ...
pterosaur Pterosaurs are 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 million to 66 million years ago). Pterosaurs are the earli ...
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 143.1 ...
period (
Aptian The Aptian is an age (geology), age in the geologic timescale or a stage (stratigraphy), stage in the stratigraphic column. It is a subdivision of the Early Cretaceous, Early or Lower Cretaceous epoch (geology), Epoch or series (stratigraphy), S ...
stage) of what is now the
Crato Formation The Crato Formation is a geologic formation (stratigraphy), formation of Early Cretaceous (Aptian) age in northeastern Brazil's Araripe Basin. It is an important Lagerstätten, Lagerstätte (undisturbed fossil accumulation) for palaeontology, pa ...
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, south ...
in
Ceará Ceará (, ) is one of the 26 states of Brazil, located in the Northeast Region, Brazil, northeastern part of the country, on the Atlantic Ocean, Atlantic coast. It is the List of Brazilian states by population, eighth-largest Brazilian State by ...
,
Brazil Brazil, officially the Federative Republic of Brazil, is the largest country in South America. It is the world's List of countries and dependencies by area, fifth-largest country by area and the List of countries and dependencies by population ...
. 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 and ) is a genus of pterosaur that included some of the largest known flying reptiles, with ''P. longiceps'' having a wingspan of over . They lived during the late Cretaceous geological period of North America in presen ...
''-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 now 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 125 million years ago. Discovery and naming Between 1995 and 2003, bone fragments of an u ...
'' is another example.


Etymology

The genus was named by Eberhard Frey ''et al.'' in 2003 and contains one known
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), ...
, ''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 spoken by the Latins (Italic tribe), Latins in Latium (now known as Lazio), the lower Tiber area aroun ...
''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 and ) is a genus of pterosaur that included some of the largest known flying reptiles, with ''P. longiceps'' having a wingspan of over . They lived during the late Cretaceous geological period of North America in presen ...
''-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 ) is a genus of extinct 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 reptile and one of the first prehis ...
''. The
specific name Specific name may refer to: * in Database management systems, a system-assigned name that is unique within a particular database In taxonomy, either of these two meanings, each with its own set of rules: * Specific name (botany), the two-part (bino ...
"''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.#Anson, Ansón, Fernández & Ramos (2015) pp. 28–34. Works of paleoart may be r ...
John Sibbick John Sibbick is a British freelance illustrator and palaeoartist, known for his depictions of prehistoric life and for his fantasy art. Early life John Sibbick began drawing in his childhood. His earliest art involved drawing figures and co ...
.


Description

''Ludodactylus'' is based on
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 ...
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 Anhangueridae (alternatively called Ornithocheiridae, 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 ...
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 mammals h ...
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'' ( , YUCK-uh) is both the scientific name and common name for a genus native to North America from Panama to southern Canada. It contains 50 accepted species. In addition to yucca, they are also known as Adam's needle or Spanish-bayon ...
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 pecking, grasping, and holding (in probing for food, eating, manipulating and ...
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 Anhangueridae (alternatively called Ornithocheiridae, 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 E ...
. 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 na ...
'', and therefore a
junior synonym In taxonomy, the scientific classification of living organisms, a synonym is an alternative scientific name for the accepted scientific name of a taxon. The botanical and zoological codes of nomenclature treat the concept of synonymy differently. ...
. In 2014 however, Brian Andres and colleagues, in a large
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 of pterosaurs, found ''Ludodactylus'' to be slightly more closely related to ornithocheirids and
anhanguerid Anhangueridae (alternatively called Ornithocheiridae, 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 ...
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 Ornithocheiromorpha (from Ancient Greek, meaning "bird hand form") is a group of pterosaurs within the suborder Pterodactyloidea. Fossil remains of this group date back from the Early to Late Cretaceous periods (Valanginian to Turonian stages) ...
. 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 125 million years ago. Discovery and naming Between 1995 and 2003, bone fragments of an u ...
'' 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 Genus, 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 considere ...
*
Timeline of pterosaur research This timeline of pterosaur research is a chronologically ordered list of important fossil discoveries, controversies of interpretation, and Biological taxonomy, taxonomic revisions of pterosaurs, the famed flying reptiles of the Mesozoic Era (ge ...


References

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