Aerosaurus Skull
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''Aerosaurus'' (meaning "copper lizard") is an extinct
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
within
Varanopidae Varanopidae is an extinct family (biology), family of amniotes known from the Late Carboniferous to Middle Permian that resembled monitor lizards (with the name of the group deriving from the monitor lizard genus ''Varanus'') and may have filled ...
, 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 non-
mammal A mammal () is a vertebrate animal of the Class (biology), class Mammalia (). Mammals are characterised by the presence of milk-producing mammary glands for feeding their young, a broad neocortex region of the brain, fur or hair, and three ...
ian
synapsid Synapsida is a diverse group of tetrapod vertebrates that includes all mammals and their extinct relatives. It is one of the two major clades of the group Amniota, the other being the more diverse group Sauropsida (which includes all extant rept ...
s. It lived between 252-299 million years ago during the
Early Permian 01 or 01 may refer to: * The year 2001, or any year ending with 01 * The month of January * 1 (number) Music * '01 (Richard Müller album), ''01'' (Richard Müller album), 2001 * 01 (Urban Zakapa album), ''01'' (Urban Zakapa album), 2011 * ''01011 ...
in
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 ...
. The name comes from Latin ''aes'' (''aeris'') (combining stem: ''aer''-) “copper” and Greek ''sauros'' “lizard,” for El Cobre Canyon (from Spanish ''cobre'' “copper”) in northern
New Mexico New Mexico is a state in the Southwestern United States, Southwestern region of the United States. It is one of the Mountain States of the southern Rocky Mountains, sharing the Four Corners region with Utah, Colorado, and Arizona. It also ...
, where the type fossil was found and the site of former
copper Copper is a chemical element; it has symbol Cu (from Latin ) and atomic number 29. It is a soft, malleable, and ductile metal with very high thermal and electrical conductivity. A freshly exposed surface of pure copper has a pinkish-orang ...
mines. ''Aerosaurus'' was a small to medium-bodied carnivorous synapsid characterized by its recurved teeth, triangular lateral temporal fenestra, and extended teeth row. Two species are recognized: ''A. greenleeorum'' (1937) and ''A. wellesi'' (1981).


History and discovery

''Aerosaurus'' was a taxon named by American paleontologist and biologist
Alfred Romer Alfred Sherwood Romer (December 28, 1894 – November 5, 1973) was an American paleontologist and biologist and a specialist in vertebrate evolution. Biography Alfred Romer was born in White Plains, New York, the son of Harry Houston Romer an ...
in 1937. There have only been two species named: ''A. greenleeorum'' (1937) and ''A. wellesi'' (1981), both discovered in New Mexico. ''A. greenleeorum'' Romer, 1937, is based on bone fragments assumed from a single individual collected by Romer in El Cobre Canyon near the
Abo The ABO blood group system is used to denote the presence of one, both, or neither of the A and B antigens on erythrocytes (red blood cells). For human blood transfusions, it is the most important of the 47 different blood type (or group) cla ...
/
Cutler Formation The Cutler Formation or Cutler Group is a stratigraphic unit exposed across the U.S. states of Arizona, northwest New Mexico, southeast Utah and southwest Colorado. It was laid down in the Early Permian during the Wolfcampian epoch. Desc ...
of the Arroyo del Agua. Cross-examination of the fragmentary holotype of ''A. greenleeorum'' Romer with the Camp Quarry skeleton with its expanded tooth-bearing parasphenoidal plate matches it to ''Aerosaurus'' taxa. The discovery of ''A. wellesi'' came from a team led in 1935 by Dr. Samuel P. Welles, who excavated blocks of red siltstone measuring thirty square feet in the
Lower Permian The Cisuralian, also known as the Early Permian, is the first series/epoch of the Permian. The Cisuralian was preceded by the Pennsylvanian and followed by the Guadalupian. The Cisuralian Epoch is named after the western slopes of the Ural Mount ...
Abo/Cutler Formation in near the Arroyo del Aqua settlement. The blocks of
siltstone Siltstone, also known as aleurolite, is a clastic sedimentary rock that is composed mostly of silt. It is a form of mudrock with a low clay mineral content, which can be distinguished from shale by its lack of fissility. Although its permeabil ...
contained two incomplete skulls of ''Oedaleops campi'' Langston, 1965, and the two nearly complete articulated skeletons of a varanopseid
pelycosaur Pelycosaur ( ) is an older term for basal or primitive Late Paleozoic synapsids, excluding the therapsids and their descendants. Previously, the term mammal-like reptile was used, and Pelycosauria was considered an order, but this is now thoug ...
.


Description

''Aerosaurus'' has been recognized as a basal member of the
Synapsida Synapsida is a diverse group of tetrapod vertebrates that includes all mammals and their extinct relatives. It is one of the two major clades of the group Amniota, the other being the more diverse group Sauropsida (which includes all extant rep ...
, a clade that ultimately produced
mammals A mammal () is a vertebrate animal of the class Mammalia (). Mammals are characterised by the presence of milk-producing mammary glands for feeding their young, a broad neocortex region of the brain, fur or hair, and three middle e ...
. Relative to the skull, the body was much larger and the ribs were deeper. The tail was unusually long for a synapsid. Overall varanopseid pelycosaurs measure about one meter long with fewer maxillary and dentary teeth compared to other ''Varanops''. In ''Aerosaurus'', the teeth were more laterally compressed creating a strong recurved shape. ''Aerosaurus'' cranial morphology is distinguishable with its extremely long tooth row, recurved teeth, and triangular lateral temporal fenestra. The teeth were so highly curved and compressed that they may have had difficulty penetrating flesh, and the tooth row extended far behind the orbit. The lower teeth were also relatively tiny and such an arrangement suggests ''Aerosaurus'' was a
carnivore A carnivore , or meat-eater (Latin, ''caro'', genitive ''carnis'', meaning meat or "flesh" and ''vorare'' meaning "to devour"), is an animal or plant Plants are the eukaryotes that form the Kingdom (biology), kingdom Plantae; they ar ...
. Their limbs were long, and skeleton built lightly suggests they were active and agile. A long low skull and dentition extended far back to lie below the temporal fenestra. ''Aerosaurus'' had other general features found in ‘pelycosaurs’, like a slanted occiput, a lateral temporal fenestra, a septomaxilla bone that contacting the nasal and a maxilla bone contacting the quadrojugal.


Skull

The skull is 1/4 of the precaudal length of the skeleton with a jaw articulation far back seen in '' Varanops''. The snout is long and low with an lessen inclined occiput. The dermal bones are shallow, irregular with pits and grooves and below the orbit we find a saw-like serration that hasn't been seen in other 'pelycosaurs'. The skull roof has large round orbits and an undetermined laterally expanded prefrontal process. The postorbital is incomplete, but the postorbital and suborbital bars are thinly shaped. The
pineal foramen A parietal eye (third eye, pineal eye) is a part of the epithalamus in some vertebrates. The eye is at the top of the head; is photoreceptive; and is associated with the pineal gland, which regulates circadian rhythmicity and hormone production ...
is larger compared to non-varanopseid sphenacodonts and is positioned in the back of the skull. The lateral temporal fenestra of ''Aerosaurus'' is larger than other pelycosaurs with its trademark triangular shape. No septomaxilla is evident, but the premaxilla is fragile in comparison to the large naris. The maxilla contains specimens largest teeth and is extended posterior to the center of the triangular lateral temporal fenestra. The maxilla, which is longer compared to other 'pelycosaurs' contacts the quadratojugal in this specimen. The lacrimal is a sheet of bone that covers most of the anterior orbital margin and constricts the front of the orbit. The jugal has a unique lateral serration and narrow postorbital ramus similar to ''Varanops''. The quadtrojugal has a slender ramus that meets the jugal and the maxilla. The nasal is also long and slender, but has extensive fragmentation, so an exact length is undetermined. The frontal is long and slender with the posterior end not extending into the margins of the orbit. The Postfrontal is small due to the size of the frontal. The squamosal is long and narrow and even more so compared to ''Varanops''. The parietal is large with shallow grooves in the supratemporal element. The braincase is well preserved with only the cultriform process missing in all of its external surfaces. The parasphenoid is the largest part of the braincase with some overlap into the basiooccipital to form the "fenestra ovalis". The presence of the many strong, pointed, and recurved teeth is a distinguishing feature of the bottom side of the braincase. The teeth are arranged in a pattern of three groups to form a U-shaped band. There is trouble distinguishing between the basisphenoid and the parasphenoid due to the co-
ossification Ossification (also called osteogenesis or bone mineralization) in bone remodeling is the process of laying down new bone material by cells named osteoblasts. It is synonymous with bone tissue formation. There are two processes resulting in t ...
creating a vague suture line. The basisphenoid has a large basipterygoid process that extend from the base of the cultriform process. The mandible has a long and slender shape with grooves and pits like on the skull roof. The dentary is long with a posterodorsal process that extends into the lower jaw and overlapping the surangular. The angular and prearticular bones are long with the splenial almost as long as the dentary. The coronoid was unidentifiable in this specimen. A foramen intermandibularis caudalis is on the medial surface of the mandible similar to the related Varannops and ''
Ophiacodon ''Ophiacodon'' (meaning "snake tooth") is an extinct genus of synapsid belonging to the family Ophiacodontidae that lived from the Late Carboniferous to the Early Permian in North America and Europe. The genus was named along with its type specie ...
'' species. The skull of ''Aerosaurus'' had 16 marginal teeth, less than in ''Varanops'' (44) and the upper teeth were longer. Most of the teeth were recurved more than other 'pelycosaurs' with a triangular depression near the base. The size of each teeth varied with some teeth-sized gap spaces. There largest teeth (7) were on the maxilla and decreased in length posterior to the orbit. There is additional six long recurved teeth located ventral to the postorbital bar, but the function of them are a mystery. The mandibular teeth are smaller, closely spaced, uniform in its recurved shape compared to the maxilla.


Postcrania

There are twenty-six presacral vertebrae with some missing due to erosion. Around five of the anterior vertebrae could possibly be cervical and three sacral vertebrae are present in this specimen. Overall there are at least 70 caudal vertebrae making up its long slender tail, which is greater than any other 'pelycosaur'. The neural spines are shorter with a typical
atlas An atlas is a collection of maps; it is typically a bundle of world map, maps of Earth or of a continent or region of Earth. Advances in astronomy have also resulted in atlases of the celestial sphere or of other planets. Atlases have traditio ...
bearing a posterior spine to pass to the zygapophyses. The axis is about the length of the other cervical vertebrae. Each vertebra is ossified with no visible suture between the centrum and
neural arch Each vertebra (: vertebrae) is an irregular bone with a complex structure composed of bone and some hyaline cartilage, that make up the vertebral column or spine, of vertebrates. The proportions of the vertebrae differ according to their spinal ...
. The distal caudal vertebrae is slender and cylinder shaped and is poorly preserved compared to the other portion of vertebrae. Presacral vertebrae have a slender double-headed ribs, but which rib is the longest is undetermined because of the articulation of the fossil. Some dorsal ribs are distally expanded. The
Gastralia Gastralia (: 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 reptil ...
is seen in a cluster of tiny rods in some articulated ribs. Gastralia are narrow plates in other 'pelycosaurs' compared to ''Aerosaurus''. The
pectoral girdle The shoulder girdle or pectoral girdle is the set of bones in the appendicular skeleton which connects to the arm on each side. In humans, it consists of the clavicle and scapula; in those species with three bones in the shoulder, it consists o ...
is preserved in a medial surface with the left scapula partially exposed. The
clavicle The clavicle, collarbone, or keybone is a slender, S-shaped long bone approximately long that serves as a strut between the scapula, shoulder blade and the sternum (breastbone). There are two clavicles, one on each side of the body. The clavic ...
is also preserved, but no
cleithrum The cleithrum (: cleithra) is a membrane bone which first appears as part of the skeleton in primitive Osteichthyes, bony fish, where it runs vertically along the scapula. Its name is derived from Greek κλειθρον = "key (lock)", by analogy ...
is identifiable in this specimen. The
shoulder girdle The shoulder girdle or pectoral girdle is the set of bones in the appendicular skeleton which connects to the arm on each side. In humans, it consists of the clavicle and scapula; in those species with three bones in the shoulder, it consists o ...
similar to Varanops with a curved
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 ...
. The clavicle is expanded anteroposteriorly. The T-shaped
interclavicle An interclavicle is a bone which, in most tetrapods, is located between the clavicles. Therian mammals ( marsupials and placentals) are the only tetrapods which never have an interclavicle, although some members of other groups also lack one. In ...
is similar to ''Ophiacodon'' and is well preserved. The
pelvic girdle The hip bone (os coxae, innominate bone, pelvic bone or coxal bone) is a large flat bone, constricted in the center and expanded above and below. In some vertebrates (including humans before puberty) it is composed of three parts: the Ilium (bone) ...
is not entirely exposed in this specimen. The neck of the iliac blade is narrower compared to ''A. greenleeorum'', but the ischium is longer and ossified. The pubis is not exposed well enough to be described. The left side of the limbs are well preserved with the humerus and femur in similar length to the lower legs. Allowing for a high posture and long strides for a predaceous mode of life. The ulna is straight, thick with a well developed olecranon. The left manus is assumed to be complete with what is visible in the fossil. Terminal
phalanges The phalanges (: phalanx ) are digit (anatomy), digital bones in the hands and foot, feet of most vertebrates. In primates, the Thumb, thumbs and Hallux, big toes have two phalanges while the other Digit (anatomy), digits have three phalanges. ...
have short recurved claws with a phalangeal formula of 2:3:4:5:3. The femoras are straight, thick with ends that aren't enlarged. The tibia is big and bowed shaped. The astragalus is short and square-cut without a long proximal neck.


Paleobiology

''A. wellesi'' had a humerus and femur in similar length to the lower legs, which allowed it to have a high posture and long strides for a predaceous or carnivorous type of life. The ossified feet and exceedingly long tail of ''Aerosaurus'' suggests an agile terrestrial carnivore ‘Pelycosaurs’ represent first group to dominate the terrestrial fauna and achieve large body size at a time where the supercontinent
Pangea Pangaea or Pangea ( ) was a supercontinent that existed during the late Paleozoic and early Mesozoic eras. It assembled from the earlier continental units of Gondwana, Euramerica and Siberia (continent), Siberia during the Carboniferous period ...
existed. The discovery of '' Pyozia mesenensis'' from the
Middle Permian The Guadalupian is the second and middle series/epoch of the Permian. The Guadalupian was preceded by the Cisuralian and followed by the Lopingian. It is named after the Guadalupe Mountains of New Mexico and Texas, and dates between 272.95 ± 0. ...
suggests there was an adaptation to ecological and climate changes that occurs into the late Paleozoic. Varanopid tended to be small to medium-sized basal 'pelycosaurs' that were carnivores and coexisted with
herbivory A herbivore is an animal anatomically and physiologically evolved to feed on plants, especially upon vascular tissues such as foliage, fruits or seeds, as the main component of its diet. These more broadly also encompass animals that eat n ...
animals in this landscape. The ''P. mesenensis'' discovery is just an increasing trend of a “mammalness” trait during the introduction of herbivory prey that appears during the
Permian The Permian ( ) is a geologic period and System (stratigraphy), 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 251.902 Mya. It is the s ...
.


Thermoregulation

Members of 'pelycosaurs' were most likely ectothermic with some theories with
Edaphosauridae Edaphosauridae is a family of mostly large (up to or more) Late Carboniferous to Early Permian synapsids. Edaphosaur fossils are so far known only from North America and Europe. Characteristics They were the earliest known herbivorous amniotes ...
and ''
Dimetrodon ''Dimetrodon'' ( or ; ) is an extinct genus of sphenacodontid synapsid that lived during the Cisuralian (Early Permian) Epoch (geology), epoch of the Permian period, around 295–272 million years ago. With most species measuring long and ...
'' using their sail structure to preserve heat. Ectothermic animals like reptiles and 'pelycosaurs' needed to maintain a level of body temperature versus the external environment in order to survive. It is estimated that the 'pelycosaur' would need to stay within the range of 24 to 35 degrees Celsius similar to modern reptiles. This is backed up by the location of 'pelycosaur discoveries in the regions of warmer temperatures in the supercontinent of Pangea in now known localities in New Mexico, Texas and Oklahoma to name a few. A change in color, respiration, blood flow and metabolism are some of the ways reptiles today regulate their body temperature and could be some of the ways 'pelycosaurs' could evolve to expand their niches.


Paleoecology

The 'pelycosaurs' and ''Aerosaurus'' would have lived in the super continent
Pangaea Pangaea or Pangea ( ) was a supercontinent that existed during the late Paleozoic and early Mesozoic eras. It assembled from the earlier continental units of Gondwana, Euramerica and Siberia during the Carboniferous period approximately 335 mi ...
. The northern and southern hemisphere during the Early Permian was hot and arid with
evaporite An evaporite () is a water- soluble sedimentary mineral deposit that results from concentration and crystallization by evaporation from an aqueous solution. There are two types of evaporite deposits: marine, which can also be described as oce ...
deposits in North America and Europe. The southernmost and Northernmost continents of Pangaea had a cold and temperate climate. The
equator The equator is the circle of latitude that divides Earth into the Northern Hemisphere, Northern and Southern Hemisphere, Southern Hemispheres of Earth, hemispheres. It is an imaginary line located at 0 degrees latitude, about in circumferen ...
of Pangaea would have a tropical and humid climate. The
flora Flora (: floras or florae) is all the plant life present in a particular region or time, generally the naturally occurring (indigenous (ecology), indigenous) native plant, native plants. The corresponding term for animals is ''fauna'', and for f ...
would change from the tropical forests of the Carboniferous to be replaced by seed-bearing plants like conifers during the Early Permian. The most discoveries from the Late Carboniferous and Early Permian tetrapods were in the Northern Hemisphere. Discoveries of fossil tetrapods from the Middle and
Late Permian Late or LATE may refer to: Everyday usage * Tardy, or late, not being on time * Late (or the late) may refer to a person who is dead Music * ''Late'' (The 77s album), 2000 * Late (Alvin Batiste album), 1993 * Late!, a pseudonym used by Dave Groh ...
were from South Africa and Russia creating a temporal gap. A temporal gap between the fossil records of the Early and Middle Permian or the Olson's Gap, after
Everett C. Olson Everett Claire Olson (November 6, 1910 – November 27, 1993) was an American zoologist, paleontologist, and geologist noted for his seminal research of origin and evolution of vertebrate animals. Through his research studying terrestrial verte ...
, a well known vertebrate paleontologist was closed with redating Permian fossil beds and newer fossil discoveries. ''Aerosaurus'' would be found in red siltstone in the Abo/Cutler Formation in New Mexico, so it existed within an open and arid environment.


Classification

''Aerosaurus'' belongs in the extinct family
Varanopidae Varanopidae is an extinct family (biology), family of amniotes known from the Late Carboniferous to Middle Permian that resembled monitor lizards (with the name of the group deriving from the monitor lizard genus ''Varanus'') and may have filled ...
, grouped among the "
pelycosaur Pelycosaur ( ) is an older term for basal or primitive Late Paleozoic synapsids, excluding the therapsids and their descendants. Previously, the term mammal-like reptile was used, and Pelycosauria was considered an order, but this is now thoug ...
s" or "mammal-like reptiles"; they became extinct around the end of the
Permian The Permian ( ) is a geologic period and System (stratigraphy), 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 251.902 Mya. It is the s ...
. Within the Varanopidae are the
subfamilies In biological classification, a subfamily (Latin: ', plural ') is an auxiliary (intermediate) taxonomic rank, next below family but more inclusive than genus. Standard nomenclature rules end botanical subfamily names with "-oideae", and zool ...
Mycterosaurinae and Varanopinae; ''Aerosaurus'' belongs to the Varanopinae in this phylogeny. ''Elliotsmithia longiceps'' is placed in the Mycterosaurinae. The sister family of Varanopidae is Ophiacontidae.


See also

*
List of pelycosaurs This list of pelycosaurs is an attempt to create a comprehensive listing of all Genus, genera that have ever been included in the synapsida excluding therapsida and purely vernacular terms. The list includes all commonly accepted genera, but also g ...


References


External links

{{Taxonbar, from=Q3278942 Varanopidae Prehistoric synapsid genera Cisuralian synapsids of North America Taxa named by Alfred Romer Fossil taxa described in 1937 Cisuralian genus first appearances Cisuralian genus extinctions