Pterosauromorpha (meaning "pterosaur-like forms") is one of the two basic divisions of
Ornithodira
Avemetatarsalia (meaning "bird metatarsals") is a clade of diapsid reptiles containing all archosaurs more closely related to birds than to crocodilians. The two most successful groups of avemetatarsalians were the dinosaurs and pterosaurs. Di ...
that includes
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 ...
s and all
taxa
In biology, a taxon (back-formation from ''taxonomy''; : taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular name and ...
that are closer to them than to
dinosaur
Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic Geological period, period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the #Evolutio ...
s and their close relatives (i.e.
Dinosauromorpha
Dinosauromorpha is a clade of avemetatarsalians ( archosaurs closer to birds than to crocodilians) that includes the Dinosauria (dinosaurs) and some of their close relatives. It was originally defined to include dinosauriforms and lagerpetids ...
).
In addition to pterosaurs, Pterosauromorpha also includes the
basal clade
In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
Lagerpetidae
Lagerpetidae (; originally Lagerpetonidae) is a family of basal avemetatarsalians (early-diverging members of the reptile lineage leading to birds and other dinosaurs). Though traditionally considered the earliest-diverging dinosauromorphs (ar ...
and some other
Late Triassic
The Late Triassic is the third and final epoch (geology), epoch of the Triassic geologic time scale, Period in the geologic time scale, spanning the time between annum, Ma and Ma (million years ago). It is preceded by the Middle Triassic Epoch a ...
ornithodirans (such as ''
Scleromochlus
''Scleromochlus'' (from , 'hard' and , 'lever') is an extinct genus of small pterosauromorph archosaurs from the Late Triassic Lossiemouth Sandstone of Scotland. The genus contains the type and only species ''Scleromochlus taylori'', named by ...
'').
Classification

The name Pterosauromorpha was originally coined by Emil Kuhn-Schnyder and Hans Rieber (1986) for a
reptilian subclass distinct from
Archosauria which includes pterosaurs. In 1997,
Kevin Padian classified Pterosauromorpha as a clade of archosaurs and proposed
phylogenetic definition for this group: "Pterosauria and all ornithodiran archosaurs closer to them than to dinosaurs".
Brian Andres and Kevin Padian redefined Pterosauromorpha as: "The clade consisting of ''
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 ...
'' (originally ''Ornithocephalus'') ''antiquus'' (Sömmerring 1812) (Pterosauria) and all organisms or species that share a more recent
common ancestor
Common descent is a concept in evolutionary biology applicable when one species is the ancestor of two or more species later in time. According to modern evolutionary biology, all living beings could be descendants of a unique ancestor commonl ...
with it than with
''Alligator'' (originally ''Crocodilus'') ''mississippiensis'' (Daudin 1802) (
Suchia
Suchia (from Ancient Greek σούχος (''soúkhos''), meaning "crocodile") is a clade of archosaurs containing the majority of pseudosuchians (crocodilians and their extinct relatives). It was defined as the least inclusive clade containing ''A ...
) and ''
Compsognathus
''Compsognathus'' (; Ancient Greek, Greek ''kompsos''/κομψός; "elegant", "refined" or "dainty", and ''gnathos''/γνάθος; "jaw") is a genus of small, bipedalism, bipedal, carnivore, carnivorous theropoda, theropod dinosaur. Members o ...
longipes'' Wagner 1859 (Dinosauromorpha)".
Lagerpetidae was traditionally considered the earliest diverging group of dinosauromorphs. This caused no other reptiles besides the true pterosaurs to be placed in Pterosauromorpha.
The only notable exception was a small reptile named ''
Scleromochlus
''Scleromochlus'' (from , 'hard' and , 'lever') is an extinct genus of small pterosauromorph archosaurs from the Late Triassic Lossiemouth Sandstone of Scotland. The genus contains the type and only species ''Scleromochlus taylori'', named by ...
'', whose placement within the clade itself remained controversial due the poor preservation of its otherwise complete
remains. Different
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 ...
found it as a basal pterosauromorph,
a non-
aphanosauria
Aphanosauria ("hidden lizards") is an extinct group of reptiles distantly related to dinosaurs (including birds). They are at the base of a group known as Avemetatarsalia, one of two main branches of archosaurs. The other main branch, Pseudosuchi ...
n, non-
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 ...
basal avemetatarsalian, a basal dinosauromorph,
or a basal
archosauriform.
This has resulted in a large gap between the fully
aerial pterosaurs and their
terrestrial ancestors, as the earliest pterosaurs were already capable flyers.
First iteration of phylogenetic analysis produced by Kammerer et al. (2020) restored lagerpetids as a basal dinosauromorphs, which corresponds to the traditional point of view. But the second iteration, in which were added ''Scleromochlus,'' found Lagerpetidae as the most basal pterosauromorphs, and ''Scleromochlus'' as 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 ...
of pterosaurs.
In a study that used
micro-CT scans, Ezcurra et al. (2020) have found additional similarities, including large
semicircular canals
The semicircular canals are three semicircular interconnected tubes located in the innermost part of each ear, the inner ear. The three canals are the lateral, anterior and posterior semicircular canals. They are the part of the bony labyrinth, ...
within the bones of some lagerpetids that resemble that of pterosaurs. It is assumed that large semicircular canals are related to
arboreal
Arboreal locomotion is the locomotion of animals in trees. In habitats in which trees are present, animals have evolved to move in them. Some animals may scale trees only occasionally (scansorial), but others are exclusively arboreal. The hab ...
, aerial or other agile forms of terrestrial locomotion as well as rapid movements. The
flocculus, the part of the brain that aids in transmitting information, was also large in both pterosaurs and lagerpetids, though to a lesser extent. When Ezcurra et al. (2020) included ''Scleromochlus'' in their analysis, they found it to be the most basal pterosauromorph, sister to a clade including lagerpetids and pterosaurs.
Baron (2021) conducted his own analysis, which confirmed the relationship between lagerpetids and pterosaurs.
Kellner et al. (2022) described ''
Maehary
''Maehary'' (meaning "one who looks to the sky" in Guarani language, Guaraní) is an extinct genus of probable gracilisuchid pseudosuchian archosaurs from the Late Triassic (Norian) Caturrita Formation of Rio Grande do Sul, Brazil. The genus con ...
,'' a small archosaur from the Late Triassic of
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 ...
. Along with lagerpetids, it was interpreted as a basal pterosauromorph. It is noteworthy that left
maxilla
In vertebrates, the maxilla (: maxillae ) is the upper fixed (not fixed in Neopterygii) bone of the jaw formed from the fusion of two maxillary bones. In humans, the upper jaw includes the hard palate in the front of the mouth. The two maxil ...
of ''Maehary'' was previously considered to be a specimen of ''
Faxinalipterus'' that was re-classified as a lagerpetid.
Contrary to the initial classification, phylogenetic analyses conducted by Müller et al. (2023) and Müller (2024) recovered ''Maehary'' as a
gracilisuchid pseudosuchia
Pseudosuchia, from Ancient Greek ψεύδος (''pseúdos)'', meaning "false", and σούχος (''soúkhos''), meaning "crocodile" is one of two major divisions of Archosauria, including living crocodilians and all archosaurs more closely relat ...
n rather than a pterosauromorph.
In their study on ''Scleromochlus'', Foffa et al. (2023) noted that the phylogenetic position of ''Maehary'' remains uncertain due to the fragmentary nature of its material.
Nesbitt et al. (2011)
Ezcurra et al. (2020)
Kellner et al. (2022)
An analysis by Garcia & Müller (2025) found lagerpetids as
paraphyletic
Paraphyly is a taxonomic term describing a grouping that consists of the grouping's last common ancestor and some but not all of its descendant lineages. The grouping is said to be paraphyletic ''with respect to'' the excluded subgroups. In co ...
to pterosaurs, and ''Faxinalipterus'' not as a pterosauromorph but as a
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 Ornithodira, more closely related to ornithodirans than to ''
Mambachiton
''Mambachiton'' (meaning "crocodile armour") is an extinct genus of basal avemetatarsalian from the Middle/ Upper Triassic Makay Formation (Isalo II beds) of Madagascar. The genus contains a single species, ''M. fiandohana'', known from a pa ...
'' and
Aphanosauria
Aphanosauria ("hidden lizards") is an extinct group of reptiles distantly related to dinosaurs (including birds). They are at the base of a group known as Avemetatarsalia, one of two main branches of archosaurs. The other main branch, Pseudosuchi ...
.
References
{{Taxonbar, from=Q2189453
Carnian first appearances
Maastrichtian extinctions