Anurognathus Ammoni
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''Anurognathus'' 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 small
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 ...
that lived during the
Late Jurassic The Late Jurassic is the third epoch of the Jurassic Period, and it spans the geologic time from 163.5 ± 1.0 to 145.0 ± 0.8 million years ago (Ma), which is preserved in Upper Jurassic strata.Owen 1987. In European lithostratigraphy, the name ...
period Period may refer to: Common uses * Era, a length or span of time * Full stop (or period), a punctuation mark Arts, entertainment, and media * Period (music), a concept in musical composition * Periodic sentence (or rhetorical period), a concept ...
(
Tithonian In the geological timescale, the Tithonian is the latest age of the Late Jurassic Epoch and the uppermost stage of the Upper Jurassic Series. It spans the time between 152.1 ± 4 Ma and 145.0 ± 4 Ma (million years ago). It is preceded by the K ...
stage). ''Anurognathus'' was first named and described by Ludwig Döderlein in 1923.Döderlein, L. (1923). "''Anurognathus Ammoni'', ein neuer Flugsaurier". ''Sitzungsberichte der Mathematisch-Naturwissenschaftlichen Abteilung der Bayerischen Akademie der Wissenschaften zu München'', 1923, 306-307. The
type species In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specimen ...
is ''Anurognathus ammoni''. The genus name ''Anurognathus'' is derived from the
Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...
''αν''/ ''an-'' ("without"), ''оυρα''/ ''oura'' ("tail"), and ''γναθος''/ ''gnathos'' ("jaw") in reference to its unusually small tail relative to other "
rhamphorhynchoid The Rhamphorhynchoidea forms one of the two suborders of pterosaurs and represents an evolutionary grade of primitive members of flying reptiles. This suborder is paraphyletic unlike the Pterodactyloidea, which arose from within the Rhamphorhynch ...
" (i.e. basal) pterosaurs."Anurognathus." In: Cranfield, Ingrid (ed.). ''The Illustrated Directory of Dinosaurs and Other Prehistoric Creatures''. London: Salamander Books, Ltd. Pp. 292-295. 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 ...
''ammoni'' honours the Bavarian geologist Ludwig von Ammon, from whose collection Döderlein had acquired the fossil in 1922.


Description

The genus 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 ...
BSP 1922.I.42 ( Bayerische Staatssammlung für Palaeontologie und Geologie), found in the Solnhofen limestone near
Eichstätt Eichstätt () is a town in the federal state of Bavaria, Germany, and capital of the district of Eichstätt. It is located on the Altmühl river and has a population of around 13,000. Eichstätt is also the seat of the Roman Catholic Diocese ...
. It consists of a crushed, relatively complete skeleton on a slab. The counterslab is missing and with it most of the bones: much of the skeleton is only visible as an impression. ''Anurognathus'' had a short head with pin-like teeth for catching
insect Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body ( head, thorax and abdomen), three pairs ...
s and although it traditionally is ascribed to the long-tailed pterosaur group "
Rhamphorhynchoidea The Rhamphorhynchoidea forms one of the two suborders of pterosaurs and represents an evolutionary grade of primitive members of flying reptiles. This suborder is paraphyletic unlike the Pterodactyloidea, which arose from within the Rhamphorhyncho ...
", its tail was comparatively short, allowing it more maneuverability for hunting. According to Döderlein the reduced tail of ''Anurognathus'' was similar to the
pygostyle Pygostyle describes a skeletal condition in which the final few caudal vertebrae are fused into a single ossification, supporting the tail feathers and musculature. In modern birds, the rectrices attach to these. The pygostyle is the main compon ...
of modern birds. Its more typical "rhamphorhynchoid" characters include its elongated fifth toe and short metacarpals and neck. With an estimated wingspan of fifty centimetres (20 inches) and a nine centimetre long body (skull included), its weight was limited: in 2008 Mark Paul Witton estimated a mass of for a specimen with a wingspan. The holotype was redescribed by
Peter Wellnhofer Peter Wellnhofer (born Munich, 1936) is a German paleontologist at the Bayerische Staatssammlung fur Paläontologie in Munich. He is best known for his work on the various fossil specimens of ''Archaeopteryx'' or "Urvogel", the first known bird. W ...
in 1975. Later a second, smaller specimen was found, probably of a subadult individual. Its slab and counterslab are separated and both were sold to private collections; neither has an official registration. It was described by S. Christopher Bennet in 2007. This second exemplar is much more complete and better articulated. It shows impressions of a large part of the flight membrane and under UV-light remains of the muscles of the thigh and arm become visible. It provided new information on many points of the anatomy. The skull was shown to have been very short and broad, wider than long. It transpired that Wellnhofer had incorrectly reconstructed the skull in 1975, mistaking the large eye sockets for the ''fenestrae antorbitales'', skull openings that in most pterosaurs are larger than the orbits but in ''Anurognathus'' are small and together with the nostrils placed at the front of the flat snout. The eyes pointed forwards to a degree, providing some binocular vision. Most of the skull consisted of bone struts. The presumed pygostyle was absent; investigating the real nine tail vertebrae instead of impressions showed that they were unfused, though very reduced. The wing finger lacked the fourth phalanx. According to Bennett a membrane, visible near the shin, showed that the wing contacted the ankle and was thus rather short and broad. Bennett also restudied the holotype, interpreting bumps on the jaws as an indication that hairs forming a protruding bristle were present on the snout.


Classification

''Anurognathus'' was assigned by
Oskar Kuhn Oskar Kuhn (7 March 1908, Munich – 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 Munich, fr ...
to the
family Family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its ...
Anurognathid Anurognathidae is a family of small, short-tailed pterosaurs that lived in Europe, Asia, and possibly North America during the Jurassic and Cretaceous periods. Five genera are known: ''Anurognathus'', from the Late Jurassic of Germany; ''Jeholop ...
ae in 1937. In the modern
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
Anurognathidae, ''Anurognathus'' is the sister taxon of the clade
Batrachognathinae Anurognathidae is a family of small, short-tailed pterosaurs that lived in Europe, Asia, and possibly North America during the Jurassic and Cretaceous periods. Five genera are known: ''Anurognathus'', from the Late Jurassic of Germany; ''Jeholopt ...
, which contains the species ''
Batrachognathus ''Batrachognathus'' is an extinct genus of anurognathid pterosaur from the Late Jurassic ( Oxfordian to Kimmeridgian) Karabastau Formation of the central Asian republic of Kazakhstan. The genus was named in 1948 by the Russian paleontologist Ana ...
'', ''
Dendrorhynchoides ''Dendrorhynchoides'' was a genus of anurognathid pterosaur containing only the holotype species ''D. curvidentatus'' that is known from the Middle Jurassic Tiaojishan Formation of Qinglong, northern Hebei Province, China. The genus was first na ...
'' and ''
Jeholopterus ''Jeholopterus'' was a small anurognathid pterosaur from the Middle to Late Jurassic Daohugou Beds of the Tiaojishan Formation of Inner Mongolia, China, preserved with hair-like pycnofibres and skin remains. Naming The genus was named in 2002 ...
''. In 2021, a
phylogenetic analysis In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
conducted by Xuefang Wei and colleagues recovered ''Anurognathus'' within the subfamily
Anurognathinae Anurognathidae is a family of small, short-tailed pterosaurs that lived in Europe, Asia, and possibly North America during the Jurassic and Cretaceous periods. Five genera are known: ''Anurognathus'', from the Late Jurassic of Germany; ''Jeholo ...
, a subfamily within the family
Anurognathidae 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. Five genera are known: ''Anurognathus'', from the Late Jurassic of G ...
. ''Anurognathus'' was found to have been the sister taxon to ''
Vesperopterylus ''Vesperopterylus'' (meaning "dusk wing") is a genus of anurognathid pterosaur from the Early Cretaceous Jiufotang Formation of China, the geologically youngest member of its group. Notably, ''Vesperopterylus'' appears to have a reversed first to ...
'' in the study. Below is a cladogram representing their phylogenetic analysis:


Paleobiology

According to Döderlein ''Anurognathus'' was, with its long wings, a swift flyer, surprising its prey, similar to the modern
nightjar Nightjars are medium-sized nocturnal or crepuscular birds in the family Caprimulgidae and order Caprimulgiformes, characterised by long wings, short legs, and very short bills. They are sometimes called goatsuckers, due to the ancient folk ta ...
. Bennett, however, infers from the discovery of the true shorter size of the wings, combined with the short tail, that it was a slower flying predator, specialised in hunting by manoeuvrability, its large eyes adapted to a
crepuscular In zoology, a crepuscular animal is one that is active primarily during the twilight period, being matutinal, vespertine, or both. This is distinguished from diurnal and nocturnal behavior, where an animal is active during the hours of daylig ...
way of life. This would also be supported by a very large flexibility of the wing finger joints. Researcher Chris Bennett published a paper in 1995 that looked into the growth rates of pterosaurs. With the fossils found of ''Rhamphorhynchus'', he was able to see that all bones that were preserved on few-day-old species were very hard from ossifications just as would be seen with adults. This developmental stage was rapidly progressing compared to that of modern-day birds. This would lead to the young pterosaurs being less dependent on their parents or for shorter durations and be able to fly away sooner at an earlier stage compared to how we see in modern-day birds. Habib and WittonHabib, M., & Witton, M. (n.d.). Early Pterosaurs and Dimorphodontidae PTEROSAURIA > PREONDACTYLUS DIMORPHODONTIDAE. Pterosaurs, 74-76. also went in to describe how insectivory evolved with pterosaurs, just like what is seen with ''Anurognathus ammoni'' and ''Dendrorhynchoides curvidentatus'', and how that relates to their functional morphology. They explained that with the great locomotion and flexibility that their wings showed, ''Anurognathus ammoni'', ''Dendrorhynchoides curvidentatus'', and some other small species, were able to catch insects and maneuver between low areas and potentially around/under trees. When predicting the gape measurements for these small pterosaurs, they used measurements of the premaxillary tip all the way to the jaw tips and then the width between the sections as references. They predicted 1734mm^2 for the estimated gaps provided from the holotype of ''Anurognathus'', ''Dendrorhynchoides'', and ''Jeholopterus''. The research team ended up concluding that consumed prey would have to be very small in length, around 11 mm total.


See also

*
Pterosaur size Pterosaurs included the largest flying animals ever to have lived. They are a clade of prehistoric archosaurian reptiles closely related to dinosaurs. Species among pterosaurs occupied several types of environments, which ranged from aquatic to ...
*
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, Germany Late Jurassic pterosaurs of Europe Monofenestratans Solnhofen fauna Fossil taxa described in 1923