Propterodactylus
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''Propterodactylus'' (meaning "before ''
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 ...
''") is an
extinct Extinction is the termination of an organism by the death of its Endling, last member. A taxon may become Functional extinction, functionally extinct before the death of its last member if it loses the capacity to Reproduction, reproduce and ...
genus of transitional
monofenestrata Monofenestrata is a clade of pterosaurs. It includes the pterosaurs in which the nasal and antorbital fenestra (openings/holes) in the skull are merged into a single fenestra. The clade includes the pterodactyloids and their close relatives. C ...
n pterosaurs from the
Late Jurassic The Late Jurassic is the third Epoch (geology), epoch of the Jurassic Period, and it spans the geologic time scale, geologic time from 161.5 ± 1.0 to 143.1 ± 0.8 million years ago (Ma), which is preserved in Upper Jurassic stratum, strata.Owen ...
Painten Formation of Germany. The genus contains a single species, ''P. frankerlae'', known from a complete articulated skeleton. Before its naming, ''Propterodactylus'' was referred to as the "Painten " in the scientific literature.


Discovery and naming

The ''Propterodactylus''
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 ...
specimen, DMA-JP-2011/006, was discovered in April 2011 in the Rygol lime works quarry of the Painten Formation (Ulmense subzone) near Painten in
Bavaria Bavaria, officially the Free State of Bavaria, is a States of Germany, state in the southeast of Germany. With an area of , it is the list of German states by area, largest German state by land area, comprising approximately 1/5 of the total l ...
, Germany. The nearly-perfect specimen consists of an
articulated An articulated vehicle is a vehicle which has a permanent or semi-permanent coupling in its construction. This coupling works as a large pivot joint, allowing it to bend and turn more sharply. There are many kinds, from heavy equipment to buse ...
complete skeleton with preserved soft tissues. In 2024, Frederik Spindler described ''Propterodactylus frankerlae'' as a new genus and species of monofenestratan pterosaur based on these fossil remains. The generic name, ''Propterodactylus'', combines the
Ancient Greek Ancient Greek (, ; ) includes the forms of the Greek language used in ancient Greece and the classical antiquity, ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following periods: Mycenaean Greek (), Greek ...
suffix "''προ-''" ("pro-"), meaning "before", with the genus name ''
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 ...
'' (derived from Greek words meaning "wing digit"). As such, the name references its position as a "forerunner" to the Pterodactyloidea, as well as its long-standing nickname as the "Painten ". 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 ...
, ''frankerlae'', honours Petra Hahn,
née The birth name is the name of the person given upon their birth. The term may be applied to the surname, the given name or to the entire name. Where births are required to be officially registered, the entire name entered onto a births registe ...
Frankerl, the deceased wife of Stephan Hahn, the discoverer of the ''Propterodactylus'' holotype.


Description

The holotype specimen of ''Propterodactylus'' belongs to either a juvenile or subadult individual. As such, it had not reached full skeletal maturity. The lack of fusion in several skeletal elements, such as the
synsacrum The synsacrum is a skeletal structure of birds and other dinosaurs, pterosaurs, as well as xenarthran mammals, in which the sacrum is extended by incorporation of additional fused or partially fused caudal or lumbar In tetrapod anatomy, lum ...
,
scapulocoracoid The scapulocoracoid is the unit of the pectoral girdle that contains the coracoid and scapula. The coracoid itself is a beak-shaped bone that is commonly found in most vertebrates with a few exceptions. The scapula is commonly known as the ''shoulde ...
,
carpal The carpal bones are the eight small bones that make up the wrist (carpus) that connects the hand to the forearm. The terms "carpus" and "carpal" are derived from the Latin carpus and the Greek καρπός (karpós), meaning "wrist". In huma ...
s, and ischiopubic plate, indicate that the animal was not fully grown. However, the general robusticity of the bones and the presence of large carpals and tarsals indicate that it was also not a young juvenile. The total length of the holotype skull is , and it has a wingspan of approximately . Despite the 'transitional' nature of the ''Propterodactylus'' holotype, the skull has several features otherwise seen only in later diverging ctenochasmatoids, such as a nearly-circular orbit, a round posterior region of the skull, and generally horizontal occipital region. In general, the skull is most proportionally similar to ''
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 ...
''.


Classification

Based on the mosaic of features observed in the ''Propterodactylus'' holotype, it can be confidently regarded as transitional between the more basal
monofenestrata Monofenestrata is a clade of pterosaurs. It includes the pterosaurs in which the nasal and antorbital fenestra (openings/holes) in the skull are merged into a single fenestra. The clade includes the pterodactyloids and their close relatives. C ...
ns such as
darwinoptera Monofenestrata is a clade of pterosaurs. It includes the pterosaurs in which the nasal and antorbital fenestra (openings/holes) in the skull are merged into a single fenestra. The clade includes the pterodactyloids and their close relatives. C ...
ns and the later diverging lophocratian
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 ...
s. Several
phylogenetic analyses In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as Computational phylogenetics, phylogenetic inference. It infers the relationship among organ ...
have been conducted that include ''Propterodactylus'' (as the "Painten "), recovering generally similar results. Most recent studies have found ''Propterodactylus'' within the Pterodactyloidea, rather than the
sister taxon In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and ...
to it. The results of Andres (2021) are displayed in the
cladogram A cladogram (from Greek language, Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an Phylogenetic tree, evolutionary tree because it does not s ...
below, with ''Propterodactylus'' as the sister taxon to the Lophocratia within Pterodactyloidea, diverging after the fragmentary ''
Kryptodrakon ''Kryptodrakon'' is an extinct genus of pterodactyloid pterosaur from the Middle to Late Jurassic with an age of approximately 162.7 million years. It is known from a single type species, ''Kryptodrakon progenitor''. The age of its fossil remai ...
''. Pêgas (2024) independently recovered the same relationships, but with
anurognathid Anurognathidae is a family (biology), family of small, short-tailed pterosaurs that lived in Europe, Asia, and possibly North America during the Jurassic and Cretaceous periods. Eight genera are definitively known: ''Anurognathus'', from the Late ...
s outside of the Monofenestrata. Similarly, Dalla Vecchia (2022) found ''Propterodactylus'' to be the basalmost pterodactyloid in an analysis with a smaller sample size, with the Darwinoptera as the sister taxon to within the Monofenestrata. Some older studies, such as Vidovic & Martill (2018), discussed the ''Propterodactylus'' fossil material as belonging to a non-pterodactyloid immediately basal to the clade's divergence. Similarly, Wang et al. (2017) recovered ''Propterodactylus'' as the sister taxon to the Pterodactyloidea, with ''
Douzhanopterus ''Douzhanopterus'' is an extinct genus of monofenestratan pterosaur from the Late Jurassic of Liaoning, China. It contains a single species, ''D. zhengi'', named by Wang ''et al.'' in 2017. In many respects, it represents a transitional fossil, t ...
'' as the sister to this clade. They further noted morphological similarities between ''Douzhanopterus'' and ''Propterodactylus'' indicating the "intermediate" status of both taxa. Later in 2024, Smyth et al. analyzed the evolution of the manus and pes in pterosaurs. They noted that in all pterodactyliforms except for ''Propterodactylus'', the ratio of the toe bones exceeds that of the finger bones. In this way, ''Propterodactylus'' is more similar to more basal taxa. They reconstructed the ancestral character-states for pterosaur hand and foot proportions. The results of their tree are displayed in the cladogram below, with ''Propterodactylus'' as the basalmost pterodactyliform:


See also

*
Monofenestrata Monofenestrata is a clade of pterosaurs. It includes the pterosaurs in which the nasal and antorbital fenestra (openings/holes) in the skull are merged into a single fenestra. The clade includes the pterodactyloids and their close relatives. C ...
* ''
Kryptodrakon ''Kryptodrakon'' is an extinct genus of pterodactyloid pterosaur from the Middle to Late Jurassic with an age of approximately 162.7 million years. It is known from a single type species, ''Kryptodrakon progenitor''. The age of its fossil remai ...
'' *
Transitional fossil A transitional fossil is any fossilized remains of a life form that exhibits traits common to both an ancestral group and its derived descendant group. This is especially important where the descendant group is sharply differentiated by gross ...
*
Paleobiota of the Solnhofen Limestone The Solnhofen Limestone or Solnhofen Plattenkalk is a collective term for multiple Late Jurassic lithographic limestones in southeastern Germany, which is famous for its well-preserved fossil flora and fauna dating to the late Jurassic (Kimmeridg ...
*
2024 in archosaur paleontology This article records new taxa of every kind of fossil archosaur that are scheduled to be Species description, described during 2024, as well as other significant discoveries and events related to the paleontology of archosaurs published in 2024. ...


References

{{Taxonbar, from1=Q127726607, from2=Q127726621 Monofenestrata Late Jurassic pterosaurs of Europe Fossils of Germany Jurassic Germany Fossil taxa described in 2024 Monotypic prehistoric reptile genera