Aphrosaurus Furlongi
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''Aphrosaurus'' was an extinct genus of
plesiosaur The Plesiosauria or plesiosaurs are an Order (biology), order or clade of extinct Mesozoic marine reptiles, belonging to the Sauropterygia. Plesiosaurs first appeared in the latest Triassic Period (geology), Period, possibly in the Rhaetian st ...
from the
Maastrichtian The Maastrichtian ( ) is, in the International Commission on Stratigraphy (ICS) geologic timescale, the latest age (geology), age (uppermost stage (stratigraphy), stage) of the Late Cretaceous epoch (geology), Epoch or Upper Cretaceous series (s ...
. The
type species In International_Code_of_Zoological_Nomenclature, 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 spe ...
is ''Aphrosaurus furlongi'' (LACM 2748), named by Welles in 1943. 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 ...
specimen was discovered in the
Moreno Formation The Moreno Formation is a Mesozoic geologic Formation (geology), formation located in San Joaquin Valley (California).Dinosaur remains diagnostic to the genus level are among the fossils that have been recovered from the formation.Weishampel, et ...
in
Fresno County Fresno County (), officially the County of Fresno, is a county located in the central portion of the U.S. state of California. As of the 2020 United States census, the population was 1,008,654. The county seat is Fresno, the fifth-most populo ...
,
California California () is a U.S. state, state in the Western United States that lies on the West Coast of the United States, Pacific Coast. It borders Oregon to the north, Nevada and Arizona to the east, and shares Mexico–United States border, an ...
in 1939 by rancher Frank C. Piava. A second specimen - LACM 2832 - was also found in the same formation and initially diagnosed as a juvenile of the same species, but has since been removed from the genus.


Discovery

In 1943, Samuel Welles described ''Aphrosaurus furlongi'' along with several other plesiosaurs from the same fossil assemblage in Fresno County, California. ''Aphrosaurus'' was found below a different juvenile species, '' Morenosaurus stocki'', within the Tierra Loma Member of the Panoche Hills. The Moreno Formation dates back to the early Maastrichtian, and is composed of depositional layers of
turbidite A turbidite is the geologic Deposition (geology), deposit of a turbidity current, which is a type of amalgamation of fluidal and sediment gravity flow responsible for distributing vast amounts of clastic sediment into the deep ocean. Sequencing ...
,
sandstone Sandstone is a Clastic rock#Sedimentary clastic rocks, clastic sedimentary rock composed mainly of grain size, sand-sized (0.0625 to 2 mm) silicate mineral, silicate grains, Cementation (geology), cemented together by another mineral. Sand ...
, and
shale Shale is a fine-grained, clastic sedimentary rock formed from mud that is a mix of flakes of Clay mineral, clay minerals (hydrous aluminium phyllosilicates, e.g., Kaolinite, kaolin, aluminium, Al2Silicon, Si2Oxygen, O5(hydroxide, OH)4) and tiny f ...
. It is part of the larger Chico Formation, which contacts the Panoche Formation and, during the Cretaceous, composed a sea shelf along the coast of California and the
Pacific Ocean The Pacific Ocean is the largest and deepest of Earth's five Borders of the oceans, oceanic divisions. It extends from the Arctic Ocean in the north to the Southern Ocean, or, depending on the definition, to Antarctica in the south, and is ...
. The diagnosis of ''Aphrosaurus'' as a unique species was initially determined by the presence of a deepened ventral notch on the centra of the cervical vertebrae, which was determined to be an
autapomorphy In phylogenetics, an autapomorphy is a distinctive feature, known as a Synapomorphy, derived trait, that is unique to a given taxon. That is, it is found only in one taxon, but not found in any others or Outgroup (cladistics), outgroup taxa, not ...
of the species. LACM 2832, found in the same formation, was also initially determined to be ''Aphrosaurus'' with ontogenic features, but that classification was rejected in a reappraisal on the basis of systemic differences between the two specimens that likely could not be explained by ontogeny. Welles took the name ''Aphrosaurus'' from the Greek for "sea foam" + "lizard", and ''furlongi'' in honor of University of California Berkeley field assistant and specimen preparator Eustace Furlong. The
axial skeleton The axial skeleton is the core part of the endoskeleton made of the bones of the head and trunk of vertebrates. In the human skeleton, it consists of 80 bones and is composed of the skull (28 bones, including the cranium, mandible and the midd ...
of the holotype fossil is composed of 18 cervical, three pectoral, and 15 dorsal vertebrae, though Welles initially described "10 posterior cervical preceded by 11 indeterminate cervical and followed by 17 crushed dorsals." The diagnostic ventral groove is visible on the 14th to 6th prepectorals, and the prezygapophyses project anteriorly and meet. Only a few of the vertebrae retain the diapophyses, which slant ventrally in the more posterior half, though that may be an artifact of being crushed. Only three dorsal ribs are preserved, each different from each other, suggesting that they are each from a different section of the skeleton. In addition to the axial skeleton, a good portion of the
appendicular skeleton The appendicular skeleton is the portion of the vertebrate endoskeleton consisting of the bones, cartilages and ligaments that support the paired appendages ( fins, flippers or limbs). In most terrestrial vertebrates (except snakes, legless li ...
was also preserved. Both the
pectoral Pectoral may refer to: * The chest region and anything relating to it. * Pectoral cross, a cross worn on the chest * a decorative, usually jeweled version of a gorget * Pectoral (Ancient Egypt), a type of jewelry worn in ancient Egypt * Pectora ...
and
pelvic girdles The pelvis (: pelves or pelvises) is the lower part of an anatomical trunk, between the abdomen and the thighs (sometimes also called pelvic region), together with its embedded skeleton (sometimes also called bony pelvis or pelvic skeleton). ...
are mostly complete, with a smooth clavicle-interclavicle complex lacking the keel and suture lines. The
scapula The scapula (: scapulae or scapulas), also known as the shoulder blade, is the bone that connects the humerus (upper arm bone) with the clavicle (collar bone). Like their connected bones, the scapulae are paired, with each scapula on either side ...
e shape is indeterminate due to damage, but likely met in the midline. Two elements were initially labeled by Welles as the ilium, which was rectified when O’Gorman articulated both with the
ischium The ischium (; : is ...
and was able to classify one as a sacral rib instead. Both fore and hind limbs were preserved, though both
humeri The humerus (; : humeri) is a long bone in the arm that runs from the shoulder to the elbow. It connects the scapula and the two bones of the lower arm, the radius and ulna, and consists of three sections. The humeral upper extremity consists of ...
and the left
femur The femur (; : femurs or femora ), or thigh bone is the only long bone, bone in the thigh — the region of the lower limb between the hip and the knee. In many quadrupeds, four-legged animals the femur is the upper bone of the hindleg. The Femo ...
are very badly damaged. The left humerus has a large muscle scar on the ventral surface. The
radius In classical geometry, a radius (: radii or radiuses) of a circle or sphere is any of the line segments from its Centre (geometry), center to its perimeter, and in more modern usage, it is also their length. The radius of a regular polygon is th ...
and
ulna The ulna or ulnar bone (: ulnae or ulnas) is a long bone in the forearm stretching from the elbow to the wrist. It is on the same side of the forearm as the little finger, running parallel to the Radius (bone), radius, the forearm's other long ...
in both limbs are wider than long, a typical feature seen in plesiosaurus, and an epipodial foramen is present on the right forelimb despite Welles’ initial analysis indicating none present. Welles noted that the right forelimb is severely distorted, and suggested that it was the result of a pathology sustained to the animal when it was alive. The hind limb description is based entirely on the right hind limb due to the damage to the left, and has no separation between the
trochanter A trochanter is a tubercle of the femur near its joint with the hip bone. In humans and most mammals, the trochanters serve as important muscle attachment sites. Humans have two, sometimes three, trochanters. Etymology The anatomical term ' ...
and
capitulum capitulum (plural capitula) may refer to: *the Latin word for chapter ** an index or list of chapters at the head of a gospel manuscript ** a short reading in the Liturgy of the Hours *** derived from which, it is the Latin for the assembly known ...
. There is a raised muscle scar on the ventral surface. No skull exists.


Description

''Aphrosaurus'' was a large, highly derived plesiosaur. A 2005 study by O’Keefe et al. comparing the body size and proportions of plesiosaurs across time show that ''Aphrosaurus'', like other late-Cretaceous plesiosaurs, was comparatively large, and a 2001 study also by O’Keefe comparing flipper
aspect ratio The aspect ratio of a geometry, geometric shape is the ratio of its sizes in different dimensions. For example, the aspect ratio of a rectangle is the ratio of its longer side to its shorter side—the ratio of width to height, when the rectangl ...
with equivalent wing proportions on birds and planes indicate a gliding, long-distance type of travel with low agility. Welles described it as "likely less active than '' Morenosaurus''." Like most plesiosauromorphs, ''Aphrosaurus'' probably ate a diet of relatively small prey that were then ground up by
gastrolith A gastrolith, also called a stomach stone or gizzard stone, is a rock held inside a gastrointestinal tract. Gastroliths in some species are retained in the muscular gizzard and used to grind food in animals lacking suitable grinding teeth. In ...
s. A 2008 study by Zammit et al. using a model of the complete vertebral column and neck indicated that while ''Aphrosaurus'' and similar plesiosaurs would not have been able to create a "swan curve" of the neck to strike, other feeding methods, including benthic grazing, shallow horizontal curving for ambush, and horizontal or vertical shearing during active pursuit would have been possible. Based on the fossil assemblage of the Moreno Formation, ''Aphrosaurus'' shared the waters of the Late-Cretaceous Pacific with a diverse spread of
mosasaur Mosasaurs (from Latin ''Mosa'' meaning the 'Meuse', and Ancient Greek, Greek ' meaning 'lizard') are an extinct group of large aquatic reptiles within the family Mosasauridae that lived during the Late Cretaceous. Their first fossil remains wer ...
s, turtles, and a variety of other plesiosaurs: ''Morenosaurus'', '' Fresnosaurus,'' and ''
Hydrotherosaurus ''Hydrotherosaurus'' (meaning "fisherman lizard") is an extinct genus of elasmosauridae, elasmosaurid plesiosaur from the Upper Cretaceous (Maastrichtian stage) Moreno Formation of Fresno County, California, USA. The only known species, ''H. ale ...
'', the latter two of which were larger than ''Aphrosaurus.'' Notably, five different genera of mosasaurs, mostly mosasaurines, have been discovered in the Moreno Formation. The lack of plioplatocarpines was hypothesized by Lindgren and Schulp in 2010 to indicate an environment as an open ocean with high piscivore competition for resources. Similar genera of mosasaurs in both the Moreno Formation and
Western Interior Seaway The Western Interior Seaway (also called the Cretaceous Seaway, the Niobraran Sea, the North American Inland Sea, or the Western Interior Sea) was a large inland sea (geology), inland sea that existed roughly over the present-day Great Plains of ...
indicates free exchange of species between both environments. Within the mostly morphologically similar
elasmosaurids Elasmosauridae, often called elasmosaurs or elasmosaurids, is an extinct family (biology), family of plesiosaurs that lived from the Hauterivian Stage (stratigraphy), stage of the Early Cretaceous to the Maastrichtian stage of the Late Cretaceou ...
, ''Aphrosaurus'' can be differentiated by the presence of a deepened ventral notch along the centra of the
cervical vertebrae In tetrapods, cervical vertebrae (: vertebra) are the vertebrae of the neck, immediately below the skull. Truncal vertebrae (divided into thoracic and lumbar vertebrae in mammals) lie caudal (toward the tail) of cervical vertebrae. In saurop ...
, and a wide, dorsal-ventrally compressed interclavicle-clavicle complex that lacks a ridge along the sternum.


Classification

The Elasmosaurid clade is highly unstable. However, current phylogenetic studies have recovered ''Aphrosaurus'' in a highly nested position within the
Weddellonectia Elasmosauridae, often called elasmosaurs or elasmosaurids, is an extinct family (biology), family of plesiosaurs that lived from the Hauterivian Stage (stratigraphy), stage of the Early Cretaceous to the Maastrichtian stage of the Late Cretaceou ...
clade, but outside of the Aristonectinae with the filter-feeding ''
Aristonectes ''Aristonectes'' (meaning "best swimmer") is an extinct genus of large elasmosaurid plesiosaurs that lived during the Maastrichtian stage of the Late Cretaceous. Two species are known, ''A. parvidens'' and ''A. quiriquinensis'', whose fossil ...
.''


See also

*
Timeline of plesiosaur research This timeline of plesiosaur research is a chronologically ordered list of important fossil discoveries, controversies of interpretation, Taxonomy (biology), taxonomic revisions, and cultural portrayals of plesiosaurs, an order of marine reptiles ...
*
List of plesiosaur genera This list of plesiosaurs is a comprehensive listing of all genera that have ever been included in the order Plesiosauria, excluding purely vernacular terms. The list includes all commonly accepted genera, but also genera that are now considered inv ...


Notes


External links


''Aphrosaurus''
in the Paleobiology Database
''Aphrosaurus''
in the Plesiosaur Site {{Taxonbar, from=Q4180945 Late Cretaceous plesiosaurs of North America Fossil taxa described in 1943 Elasmosauridae Taxa named by Samuel Paul Welles Sauropterygian genera