Ascendonanus
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''Ascendonanus'' (meaning "climbing dwarf") is an extinct
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
varanopid Varanopidae is an extinct family of amniotes that resembled monitor lizards and may have filled a similar niche, hence the name. Typically, they are considered synapsids that evolved from an ''Archaeothyris''-like synapsid in the Late Carbonifero ...
amniote Amniotes are a clade of tetrapod vertebrates that comprises sauropsids (including all reptiles and birds, and extinct parareptiles and non-avian dinosaurs) and synapsids (including pelycosaurs and therapsids such as mammals). They are disti ...
from the Early
Permian The Permian ( ) is a geologic period and stratigraphic system which spans 47 million years from the end of the Carboniferous Period million years ago (Mya), to the beginning of the Triassic Period 251.9 Mya. It is the last period of the Paleoz ...
of
Germany Germany,, officially the Federal Republic of Germany, is a country in Central Europe. It is the second most populous country in Europe after Russia, and the most populous member state of the European Union. Germany is situated betwe ...
. It is one of the earliest specialized arboreal (tree-living)
tetrapods Tetrapods (; ) are four-limbed vertebrate animals constituting the superclass Tetrapoda (). It includes extant and extinct amphibians, sauropsids (reptiles, including dinosaurs and therefore birds) and synapsids (pelycosaurs, extinct therapsids ...
currently known and outwardly resembled a small
lizard Lizards are a widespread group of squamate reptiles, with over 7,000 species, ranging across all continents except Antarctica, as well as most oceanic island chains. The group is paraphyletic since it excludes the snakes and Amphisbaenia alt ...
. The animal was about 40 cm long, with strongly curved claws, short limbs, a slender, elongated trunk, and a long tail. It would have preyed on
insects 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 of j ...
and other small
arthropods Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a Segmentation (biology), segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and Arth ...
. The taxonomic position of varanopids is currently debated between
synapsids Synapsids + (, 'arch') > () "having a fused arch"; synonymous with ''theropsids'' (Greek, "beast-face") are one of the two major groups of animals that evolved from basal amniotes, the other being the Sauropsida, sauropsids, the group that inc ...
(related to
mammals Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fur or ...
, the most widely accepted idea) and
diapsids Diapsids ("two arches") are a clade of sauropsids, distinguished from more primitive eureptiles by the presence of two holes, known as temporal fenestrae, in each side of their skulls. The group first appeared about three hundred million years ag ...
(related to
reptiles Reptiles, as most commonly defined are the animals in the Class (biology), class Reptilia ( ), a paraphyletic grouping comprising all sauropsid, sauropsids except birds. Living reptiles comprise turtles, crocodilians, Squamata, squamates (lizar ...
). The
fossil 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 ...
s of ''Ascendonanus'' are of special scientific importance because they include remains of skin, scales, scutes, bony ossicles, and soft-tissue body outlines, which could indicate that some of the oldest relatives of mammals had a scaly "
reptilia Reptiles, as most commonly defined are the animals in the class Reptilia ( ), a paraphyletic grouping comprising all sauropsids except birds. Living reptiles comprise turtles, crocodilians, squamates (lizards and snakes) and rhynchocephalians ( ...
n-type" appearance. The related small varanopid ''
Eoscansor ''Eoscansor'' (EE-oh-SKAN-sor; meaning "dawn climber") is a species of small Varanopidae, varanopid amniote that lives from the upper Pennsylvanian (geology), Pennsylvanian subperiod in northern New Mexico, New Mexico, United States 305 million y ...
'', recently described from
New Mexico ) , population_demonym = New Mexican ( es, Neomexicano, Neomejicano, Nuevo Mexicano) , seat = Santa Fe , LargestCity = Albuquerque , LargestMetro = Tiguex , OfficialLang = None , Languages = English, Spanish ( New Mexican), Navajo, Ker ...
, was also adapted to climbing, very likely in trees, but dates from 15 million years earlier during the Pennsylvanian subperiod of the
Carboniferous The Carboniferous ( ) is a geologic period and system of the Paleozoic that spans 60 million years from the end of the Devonian Period million years ago ( Mya), to the beginning of the Permian Period, million years ago. The name ''Carbonifero ...
, indicating that climbing varanopids have a longer history and were likely widespread. ''Ascendonanus'' was named and described in 2018 from remains of five individuals that were discovered in the
Chemnitz petrified forest The Chemnitz petrified forest is a petrified forest in Chemnitz, Germany. Most of the trunks are exhibited in the Museum of Natural History in Chemnitz inside of Kulturkaufhaus Tietz , including slices of trunks with polished edges. A small colle ...
, an Early Permian tropical fossil forest preserved under the city of
Chemnitz Chemnitz (; from 1953 to 1990: Karl-Marx-Stadt , ) is the third-largest city in the German state of Saxony after Leipzig and Dresden. It is the 28th largest city of Germany as well as the fourth largest city in the area of former East Germany a ...
, Germany. A
Pompeii Pompeii (, ) was an ancient city located in what is now the ''comune'' of Pompei near Naples in the Campania region of Italy. Pompeii, along with Herculaneum and many villas in the surrounding area (e.g. at Boscoreale, Stabiae), was buried ...
-like
pyroclastic Pyroclastic rocks (derived from the el, πῦρ, links=no, meaning fire; and , meaning broken) are clastic rocks composed of rock fragments produced and ejected by explosive volcanic eruptions. The individual rock fragments are known as pyroc ...
volcanic A volcano is a rupture in the crust of a planetary-mass object, such as Earth, that allows hot lava, volcanic ash, and gases to escape from a magma chamber below the surface. On Earth, volcanoes are most often found where tectonic plates a ...
eruption 291 million years ago buried the forest and created the Zeisigwald Tuff Horizon in the uppermost Leukersdorf Formation (late
Sakmarian In the geologic timescale, the Sakmarian is an age or stage of the Permian. It is a subdivision of the Cisuralian Epoch or Series. The Sakmarian lasted between 293.52 and million years ago (Ma). It was preceded by the Asselian and followed by t ...
/early
Artinskian In the geologic timescale, the Artinskian is an age or stage of the Permian. It is a subdivision of the Cisuralian Epoch or Series. The Artinskian likely lasted between and million years ago (Ma) according to the most recent revision of the ...
transition stage), preserving some of the animals that lived there in exceptional detail in a bottom layer of volcanic ash. The type species name ''Ascendonanus nestleri'' honors Knut Nestler, a long-time local supporter (deceased) of the Chemnitz Museum of Natural History (Museum für Naturkunde Chemnitz (MNC)), where the specimens of ''Ascendonanus'' are stored.


Description

''Ascendonanus'' was about 40 cm long, although the end of the tail is missing in all specimens and the full body length in life is not currently known. It is the smallest known member of the clade Varanopidae, a group of early synapsids that generally resembled the unrelated
monitor lizard Monitor lizards are lizards in the genus ''Varanus,'' the only extant genus in the family Varanidae. They are native to Africa, Asia, and Oceania, and one species is also found in the Americas as an invasive species. About 80 species are recogn ...
s. Features that identify ''Ascendonanus'' as a "pelycosaur" grade synapsid and a member of the Varanopidae include a single lateral temporal opening (fenestra) in the skull, a ridge on the underside of the centra of the vertebrae, and enlarged blades on the ilium of the pelvis.


Specimens

The five recovered fossils of ''Ascendonanus'' are strongly compacted and were split open as flattened counterslabs that revealed articulated partial or near complete skeletons with remains of soft tissue and some internal features. The bone material itself, however, often was not clearly preserved, making interpretation of some details more difficult. The specimens were CT scanned to reveal additional information. Based on the ossification of different bones, all individuals appear to be fully grown despite some differences in size. The specimen designated MNC-TA0924 was made the diagnostic holotype of ''Ascendonanus nestleri'' because it provides the clearest details of the skull. The most remarkable specimen (MNC-TA1045) preserves the clear body outline of nearly the entire animal on counterslabs, showing the thickness in life of the limbs and the neck, and the full covering of scales.Photo of ''Ascendonanus'' specimen TA1045 in Chemnitz Museum of Natural History. https://sachsen.museum-digital.de/index.php?t=objekt&oges=8453


Skull

Because of compaction over time, it is not known if the skull is relatively flat or if it has a taller profile. The tip of the snout is not well preserved in any specimens. The orbits are very large but the sclerotic rings to support the eyeball are not ossified. Some specimens preserve dozens of tiny round dermal bones or ossicles that were embedded in the skin of the upper eyelid. Such eyelid ossicles are currently not known in any other amniotes, but have been found in some
dissorophid Dissorophidae is an extinct family of medium-sized, temnospondyl amphibians that flourished during the late Carboniferous and early Permian periods. The clade is known almost exclusively from North America. History of study Dissorophidae is a ...
temnospondyl Temnospondyli (from Greek language, Greek τέμνειν, ''temnein'' 'to cut' and σπόνδυλος, ''spondylos'' 'vertebra') is a diverse order (biology), order of small to giant tetrapods—often considered Labyrinthodontia, primitive amphi ...
amphibians, a non-amniote tetrapod group that is not closely related to synapsids. The eyelid ossicles in ''Ascendonanus'' may have evolved independently or may be an ancient feature retained from the earliest tetrapods. Such eyelid ossicles are distinct from the rod-like dermal bones, called
palpebral bone The palpebral bone is a small dermal bone found in the region of the eye socket in a variety of animals, including crocodilians and ornithischian dinosaurs. It is also known as the adlacrimal or supraorbital, although the latter term may not be co ...
s, that evolved above the eye socket in a number of later non-synapsid groups, including some
ornithischia Ornithischia () is an extinct order of mainly herbivorous dinosaurs characterized by a pelvic structure superficially similar to that of birds. The name ''Ornithischia'', or "bird-hipped", reflects this similarity and is derived from the Greek s ...
n
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the evolution of dinosaurs is t ...
s. The pointed teeth are slender, without flattened cross-sections, serrations, or a cutting edge, and are moderately recurved in the upper jaw and straighter in the lower jaw.


Skeleton

''Ascendonanus'' differs notably from other varanopids in its elongated trunk section, with 34 to 37 presacral vertebrae compared to 26 in most synapsids. It has a dense set of thin belly ribs or
gastralia Gastralia (singular gastralium) are dermal bones found in the ventral body wall of modern crocodilians and tuatara, and many prehistoric tetrapods. They are found between the sternum and pelvis, and do not articulate with the vertebrae. In these ...
along the underside. The full length of its tail is not known based on current fossils, but it was longer than in other described varanopids and would have helped with balance in climbing, similar to some tree-living lizards with very long tails. It is unknown whether the distal part of the tail was prehensile.


Limbs

The forelimbs are almost as long as the hindlimbs, but both pairs of legs are very short compared to the length of the trunk. The elements of the feet are enlarged, with elongated and slender digits. Distinct from other varanopids, the claws on ''Ascendonanus'' have a very strong curvature. The curved claws and the size and shape of the feet, including a longer fourth digit on the manus and on the pes, indicate ''Ascendonanus'' would have been a "clinging" climber rather than a "grasping" climber.


Skin and scales

The specimens show a regular scale pattern over their bodies, similar to living
squamate Squamata (, Latin ''squamatus'', 'scaly, having scales') is the largest order of reptiles, comprising lizards, snakes, and amphisbaenians (worm lizards), which are collectively known as squamates or scaled reptiles. With over 10,900 species, ...
s and
archosaur Archosauria () is a clade of diapsids, with birds and crocodilians as the only living representatives. Archosaurs are broadly classified as reptiles, in the cladistic sense of the term which includes birds. Extinct archosaurs include non-avian d ...
s, suggesting dry, scaly skin was present in the earliest
amniote Amniotes are a clade of tetrapod vertebrates that comprises sauropsids (including all reptiles and birds, and extinct parareptiles and non-avian dinosaurs) and synapsids (including pelycosaurs and therapsids such as mammals). They are disti ...
s before the split into synapsids and sauropsids (reptiles) during the
Carboniferous Period The Carboniferous ( ) is a Period (geology), geologic period and System (stratigraphy), system of the Paleozoic that spans 60 million years from the end of the Devonian Period million years ago (Myr, Mya), to the beginning of the Permian Period, ...
. Some synapsid groups later developed bare, glandular skin, eventually with hair and whiskers that became characteristics of mammals. Unlike some varanopids (such as ''
Archaeovenator ''Archaeovenator'' is an extinct genus of Late Carboniferous varanopid synapsids known from Greenwood County, Kansas of the United States. It was first named by Robert R. Reisz and David W. Dilkes in 2003 and the type species is ''Archaeovenat ...
'' and ''
Microvaranops ''Microvaranops'' is a Middle Permian synapsid of the family Varanopidae from the Abrahamskraal Formation of South Africa.Frederik Spindler; Ralf Werneburg; Joerg W. Schneider; Ludwig Luthardt; Volker Annacker; Ronny Rößler (2018). "First arbor ...
''), ''Ascendonanus'' does not have dorsal osteoderms on its upper trunk section along the back. However, the middle part of the tail has a covering of small scutes that continues to where the end of the tail is missing in all current specimens. The body outlines preserved show that ''Ascendonanus'' had a plump neck and muscular thighs (although the
femur The femur (; ), or thigh bone, is the proximal bone of the hindlimb in tetrapod vertebrates. The head of the femur articulates with the acetabulum in the pelvic bone forming the hip joint, while the distal part of the femur articulates with ...
bone itself is relatively thin) on its short hindlimbs, while the trunk and the forelimbs are relatively slender.


Discovery


Hilbersdorf scientific excavation

All of the current fossils of ''Ascendonanus'' were discovered during a test dig by the Chemnitz Museum of Natural History in the Hilbersdorf district of Chemnitz between April 2008 to October 2011. The site was limited to a pit 24 m by 18 m, down to a depth of at least 5 m, and was the first systematic scientific excavation of the Chemnitz fossil forest ever conducted in the city. The museum began a second, more extensive, and ongoing, dig in the Sonnenberg district in 2014. In addition to finding fossils of trees and plants at the Hilbersdorf site, the team recovered remains of vertebrates (synapsids, temnospondyl,
aistopods Aistopoda (Greek for " avingnot-visible feet") is an order of highly specialised snake-like stegocephalians known from the Carboniferous and Early Permian of Europe and North America, ranging from tiny forms only , to nearly in length. They fi ...
) and arthropods (scorpions, ''
Arthropleura ''Arthropleura'' () is a genus of extinct millipede arthropods that lived in what is now North America and Europe around 345 to 290 million years ago, from the Viséan stage of the lower Carboniferous Period to the Sakmarian stage of the lower P ...
'', spiders), some later identified as species new to science.


Taphonomy

During an early phase of the volcanic eruption, rising
rhyolitic Rhyolite ( ) is the most silica-rich of volcanic rocks. It is generally glassy or fine-grained (aphanitic) in texture, but may be porphyritic, containing larger mineral crystals (phenocrysts) in an otherwise fine-grained groundmass. The mineral ...
magma Magma () is the molten or semi-molten natural material from which all igneous rocks are formed. Magma is found beneath the surface of the Earth, and evidence of magmatism has also been discovered on other terrestrial planets and some natural sa ...
came into contact with groundwater, exploding molten rock into tiny fragments mixed with steam, resulting in falls of wet, relatively cool, fine volcanic ash particles. An initial ash layer covered the ground and knocked leaves off trees, but left the trunks standing. All of the ''Acendonanus'' specimens were found close to the base of upright fossil trees from which the animals evidently had fallen to the ground onto the first ash fall, either after being dislodged by a volcanic blast or, more likely, after being overcome by breathing ash particles or toxic gases, or possibly by lethal temperatures. A second fall of wet ash quickly buried the ''Ascendonanus'' individuals and the other animals that were on or under the forest floor up to about 53 cm deep, preventing decay and preserving bodies largely intact (but compacted over time), with detailed impressions or traces of soft tissues. The fossil plant material in the wet ash layer shows no evidence of charring, indicating that the temperature of the ash fall was below 280° C. Later, more violent phases of the eruption covered the wet ash layer in much deeper deposits of coarser hot pyroclastic material that make up most of the Zeisigwald Tuff Horizon (total depth up to 4 m). The upper trunks and branches of embedded upright trees were snapped off above about 1 m to 3 m from the ground.


References

{{Taxonbar, from=Q55078956 Varanopids Prehistoric synapsid genera Fossil taxa described in 2018