Batrachognathinae
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Anurognathidae is a
family Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...
of small, short-tailed
pterosaurs Pterosaurs are an extinct clade of flying reptiles in the Order (biology), 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). Pterosau ...
that lived in
Europe Europe is a continent located entirely in the Northern Hemisphere and mostly in the Eastern Hemisphere. It is bordered by the Arctic Ocean to the north, the Atlantic Ocean to the west, the Mediterranean Sea to the south, and Asia to the east ...
,
Asia Asia ( , ) is the largest continent in the world by both land area and population. It covers an area of more than 44 million square kilometres, about 30% of Earth's total land area and 8% of Earth's total surface area. The continent, which ...
, and possibly
North America North America is a continent in the Northern Hemisphere, Northern and Western Hemisphere, Western hemispheres. North America is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South Ameri ...
during the
Jurassic The Jurassic ( ) is a Geological period, geologic period and System (stratigraphy), stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately 143.1 Mya. ...
and
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 143.1 to 66 mya (unit), million years ago (Mya). It is the third and final period of the Mesozoic Era (geology), Era, as well as the longest. At around 77.1 million years, it is the ...
periods. Eight genera are definitively known: ''
Anurognathus ''Anurognathus'' (from the Greek ''ανоυρα γναθος'' "frog jaw") is an extinct genus of small pterosaur from the Late Jurassic Altmühltal Formation of Germany. ''Anurognathus'' was first named and described by Ludwig Döderlein in 19 ...
'', 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 ...
of Germany; ''
Jeholopterus ''Jeholopterus'' was a small anurognathid pterosaur known from the Middle to Late Jurassic Daohugou Beds of the Tiaojishan Formation of Inner Mongolia, China, and possibly the Early Cretaceous Sinuiju Formation of North Korea. Naming The gen ...
'', '' Luopterus'', ''
Cascocauda ''Cascocauda'' (meaning "ancient tail") is an extinct genus of anurognathid pterosaurs from the Late Jurassic, Late–⁠Middle Jurassic, Middle Jurassic Tiaojishan Formation of Hebei, Hebei Province, China. The genus contains a single species, '' ...
'' and '' Sinomacrops'', from the
Middle Middle or The Middle may refer to: * Centre (geometry), the point equally distant from the outer limits. Places * Middle (sheading), a subdivision of the Isle of Man * Middle Bay (disambiguation) * Middle Brook (disambiguation) * Middle Creek ...
to Late Jurassic of China; '' Batrachognathus'', from the Late Jurassic of
Kazakhstan Kazakhstan, officially the Republic of Kazakhstan, is a landlocked country primarily in Central Asia, with a European Kazakhstan, small portion in Eastern Europe. It borders Russia to the Kazakhstan–Russia border, north and west, China to th ...
; and '' Dendrorhynchoides'' and ''
Vesperopterylus ''Vesperopterylus'' (meaning "dusk wing") is a genus of anurognathidae, anurognathid pterosaur from the Early Cretaceous Jiufotang Formation of China, the geologically youngest member of its group. Notably, ''Vesperopterylus'' appears to have a r ...
'', 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 ...
of China. Bennett (2007) suggested that the
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 ...
of '' Mesadactylus'', BYU 2024, a
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 ...
, belonged to an anurognathid, though this affinity has been questioned by other authors. ''Mesadactylus'' is from the Late Jurassic
Morrison Formation The Morrison Formation is a distinctive sequence of Upper Jurassic sedimentary rock found in the western United States which has been the most fertile source of dinosaur fossils in North America. It is composed of mudstone, sandstone, siltston ...
of the United States. Indeterminate anurognathid remains have also been reported from the Middle Jurassic Bakhar Svita of
Mongolia Mongolia is a landlocked country in East Asia, bordered by Russia to the north and China to the south and southeast. It covers an area of , with a population of 3.5 million, making it the world's List of countries and dependencies by po ...
and the Early Cretaceous of
North Korea North Korea, officially the Democratic People's Republic of Korea (DPRK), is a country in East Asia. It constitutes the northern half of the Korea, Korean Peninsula and borders China and Russia to the north at the Yalu River, Yalu (Amnok) an ...
. Anurognathids had wingspans of up to 900 mm.


Classification

A family Anurognathidae was named in 1928 by
Franz Nopcsa von Felső-Szilvás Baron Franz Nopcsa von Felső-Szilvás (also Baron Nopcsa von Felső-Szilvás, Baron Nopcsa, Ferenc Nopcsa, báró felsőszilvási Nopcsa Ferenc, Baron Franz Nopcsa, and Franz Baron Nopcsa; May 3, 1877 – April 25, 1933) was a Hungarian Aristoc ...
(as the subfamily Anurognathinae) with ''
Anurognathus ''Anurognathus'' (from the Greek ''ανоυρα γναθος'' "frog jaw") is an extinct genus of small pterosaur from the Late Jurassic Altmühltal Formation of Germany. ''Anurognathus'' was first named and described by Ludwig Döderlein in 19 ...
'' as the
type genus In biological taxonomy, the type genus (''genus typica'') is the genus which defines a biological family and the root of the family name. Zoological nomenclature According to the International Code of Zoological Nomenclature, "The name-bearin ...
. The family name Anurognathidae was first used by
Oskar Kuhn Oskar Kuhn (7 March 1908, Munich – 1 May 1990) was a German palaeontologist. Life and career Kuhn was educated in Dinkelsbühl and Bamberg and then studied natural science, specialising in geology and paleontology, at the University of Mun ...
in 1967. The
phylogeny A phylogenetic tree or phylogeny is a graphical representation which shows the evolutionary history between a set of species or Taxon, taxa during a specific time.Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, M ...
of Anurognathidae is disputed. Both
Alexander Kellner Alexander Wilhelm Armin Kellner (born September 26, 1961) is a Brazilian geologist and paleontologist who is a leading expert in the field of studying pterosaurs. His research has focused mainly on fossil reptiles from the Cretaceous Period, i ...
and David Unwin in 2003 defined the group as a node clade: the
last common ancestor A most recent common ancestor (MRCA), also known as a last common ancestor (LCA), is the most recent individual from which all organisms of a set are inferred to have descended. The most recent common ancestor of a higher taxon is generally assu ...
of ''Anurognathus'' and ''Batrachognathus'' and all its descendants. Some analyses, such as that of Kellner (2003), place them as the most basal group in the pterosaur tree. Unwin also recovered the group as very basal, falling between Dimorphodontidae and Campylognathoididae. However, anurognathids have some characteristics in common with the derived Pterodactyloidea, such as short and fused tail bones. More recent analyses, which include more fossils and taxa, support this observation and recover the group as substantially more derived than previously thought, but still basal to pterodactyloids. In 2010 an analysis by Brian Andres indicated the Anurognathidae and Pterodactyloidea were sister taxa. This conformed better to the
fossil record A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
because no early anurognathids were known at the time, and being the basalmost pterosaur clade would require a
ghost lineage A ghost lineage is a hypothesized ancestor in a species lineage that has left no fossil evidence, but can still be inferred to exist or have existed because of gaps in the fossil record or genomic evidence. The process of determining a ghost line ...
of over sixty million years. However, the reassignment of "''
Dimorphodon ''Dimorphodon'' ( ) is a genus of medium-sized pterosaur from Europe during the early Jurassic Period (about 201-191 million years ago).Müller R.T., Ezcurra M.D., Garcia M.S., Agnolín F.L., Stocker M.R., Novas F.E., Soares M.B., Kellner A.W.A. ...
'' ''weintraubi"'' to a basal position within Anurognathidae helps fill this gap and suggests this group appeared earlier than previously thought, possibly in the
Early Jurassic The Early Jurassic Epoch (geology), Epoch (in chronostratigraphy corresponding to the Lower Jurassic series (stratigraphy), Series) is the earliest of three epochs of the Jurassic Period. The Early Jurassic starts immediately after the Triassic ...
Period. Depending on where Anurognathidae falls within the Pterosauria, the existence of "''Dimorphodon'' ''weintraubi"'' may have important implications for the timing of the evolution of major pterosaur clades, making further study of this specimen critical for pterosaur research. In 2022, a phylogenetic analysis accompanying the description of ''
Cascocauda ''Cascocauda'' (meaning "ancient tail") is an extinct genus of anurognathid pterosaurs from the Late Jurassic, Late–⁠Middle Jurassic, Middle Jurassic Tiaojishan Formation of Hebei, Hebei Province, China. The genus contains a single species, '' ...
'' recovered Anurognathidae as a sister clade to Breviquartossa.


Lifestyle

Anurognathids are widely believed to have been
nocturnal Nocturnality is a ethology, behavior in some non-human animals characterized by being active during the night and sleeping during the day. The common adjective is "nocturnal", versus diurnality, diurnal meaning the opposite. Nocturnal creatur ...
or
crepuscular In zoology, a crepuscular animal is one that is active primarily during the twilight period, being matutinal (active during dawn), vespertine (biology), vespertine/vespertinal (active during dusk), or both. This is distinguished from diurnalit ...
akin to
bat Bats are flying mammals of the order Chiroptera (). With their forelimbs adapted as wings, they are the only mammals capable of true and sustained flight. Bats are more agile in flight than most birds, flying with their very long spread-out ...
s. The fact that many anurognathids have large eye sockets supports the theory of operating in low-light environments. Anurognathid teeth suggest they were largely insectivorous, though some may have had more prey choices, such as ''Batrachognathus'' and ''Jeholopterus,'' which have been hypothesized to have been
piscivorous A piscivore () is a carnivorous animal that primarily eats fish. Fish were the diet of early tetrapod evolution (via water-bound amphibians during the Devonian period); insectivory came next; then in time, the more terrestrially adapted rept ...
. At least some, such as ''Vesperopterylus'', were
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 ...
, with claws suited for gripping tree branches.


Feathers

A 2018 study of the remains of two small
Jurassic The Jurassic ( ) is a Geological period, geologic period and System (stratigraphy), stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately 143.1 Mya. ...
-age pterosaurs from
Inner Mongolia Inner Mongolia, officially the Inner Mongolia Autonomous Region, is an Autonomous regions of China, autonomous region of China. Its border includes two-thirds of the length of China's China–Mongolia border, border with the country of Mongolia. ...
,
China China, officially the People's Republic of China (PRC), is a country in East Asia. With population of China, a population exceeding 1.4 billion, it is the list of countries by population (United Nations), second-most populous country after ...
, named as the genus ''
Cascocauda ''Cascocauda'' (meaning "ancient tail") is an extinct genus of anurognathid pterosaurs from the Late Jurassic, Late–⁠Middle Jurassic, Middle Jurassic Tiaojishan Formation of Hebei, Hebei Province, China. The genus contains a single species, '' ...
'' in 2022, found that pterosaurs had a wide array of pycnofiber shapes and structures, as opposed to the homogeneous structures that had generally been assumed to cover them. Some of these had frayed ends, very similar in structure to four different feather types known from birds or other dinosaurs but almost never known from pterosaurs prior to the study, suggesting homology. A response to this study was published in 2020, where it was suggested that the structures seen on the anurognathids were actually a result of the decomposition of aktinofibrils: a type of fibre used to strengthen and stiffen the wing. However, in a response to this, the authors of the 2018 paper point to the fact that the presence of the structures extend past the
patagium The patagium (: patagia) is a membranous body part that assists an animal in obtaining lift when gliding or flying. The structure is found in extant and extinct groups of flying and gliding animals including bats, theropod dinosaurs (inclu ...
, and the presence of both aktinofibrils and filaments on '' Jeholopterus ningchengensis'' and ''
Sordes pilosus ''Sordes'' is a genus of small pterosaur from the late Jurassic ( Oxfordian/ Kimmeridgian) Karabastau Svita of Kazakhstan. This genus was named in 1971 by Aleksandr Grigorevich Sharov. The type species is ''Sordes pilosus''. The genus name is L ...
''. The various forms of filament structure present on the anurognathids in the 2018 study would also require a form of decomposition that would cause the different 'filament' forms seen. They therefore conclude that the most parsimonious interpretation of the structures is that they are filamentous proto-feathers. But Liliana D’Alba points out that the description of the preserved integumentary structures on the two anurognathid specimens is still based upon gross morphology. She also points out that ''Pterorhynchus'' was described to have feathers to support the claim that feathers had a common origin with ornithodirans but was argued against by several authors. The only method to assure if it was homologous to feathers is to use a scanning electron microscope.


Notes

{{Taxonbar, from=Q137354 Anurognathidae Callovian first appearances Pterosaur families