''Europasaurus'' (meaning 'Europe lizard') is a basal
macronaria
Macronaria is a clade of sauropod dinosaurs. Macronarians are named after the large diameter of the nasal opening of their skull, known as the external naris, which exceeded the size of the orbit, the skull opening where the eye is located (hence ...
n
sauropod
Sauropoda (), whose members are known as sauropods (; from '' sauro-'' + '' -pod'', 'lizard-footed'), is a clade of saurischian ('lizard-hipped') dinosaurs. Sauropods had very long necks, long tails, small heads (relative to the rest of their b ...
, a form of
quadruped
Quadrupedalism is a form of locomotion in which animals have four legs that are used to bear weight and move around. An animal or machine that usually maintains a four-legged posture and moves using all four legs is said to be a quadruped (fr ...
al
herbivorous
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 ...
dinosaur
Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic Geological period, period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the #Evolutio ...
. It lived during the Late
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. ...
(middle
Kimmeridgian
In the geologic timescale, the Kimmeridgian is an age in the Late Jurassic Epoch and a stage in the Upper Jurassic Series. It spans the time between 154.8 ±0.8 Ma and 149.2 ±0.7 Ma (million years ago). The Kimmeridgian follows the Oxfordian ...
, from about 154 to 151 million years ago) of northern
Germany
Germany, officially the Federal Republic of Germany, is a country in Central Europe. It lies between the Baltic Sea and the North Sea to the north and the Alps to the south. Its sixteen States of Germany, constituent states have a total popu ...
, and has been identified as an example of
insular dwarfism
Insular dwarfism, a form of phyletic dwarfism, is the process and condition of large animals evolving or having a reduced body size when their population's range is limited to a small environment, primarily islands. This natural process is disti ...
resulting from the isolation of a sauropod population on an island within the
Lower Saxony
Lower Saxony is a States of Germany, German state (') in Northern Germany, northwestern Germany. It is the second-largest state by land area, with , and fourth-largest in population (8 million in 2021) among the 16 ' of the Germany, Federal Re ...
basin.
Discovery and naming

In 1998, a single sauropod tooth was discovered by private fossil collector Holger Lüdtke in an active quarry at
Langenberg Mountain, between the communities of
Oker
The Oker () is a river in Lower Saxony, Germany, that has historically formed an important political boundary. It is a left tributary of the River Aller (Germany), Aller, in length and runs in a generally northerly direction.
Origin and meanin ...
,
Harlingerode and
Göttingerode
Göttingerode (; ) is a village in Germany and district of Bad Harzburg in the district of Goslar in Lower Saxony. As of 2020, Göttingerode had a population of 902. It gained international relevance owing to the discovery of the Europasaurus in ...
in Germany. The
Langenberg chalk quarry had been active for more than a century; rocks are quarried using blasting and are mostly processed into fertilisers. The quarry exposes a nearly continuous, thick succession of
carbonate rock
Carbonate rocks are a class of sedimentary rocks composed primarily of carbonate minerals. The two major types are limestone, which is composed of calcite or aragonite (different crystal forms of CaCO3), and Dolomite (rock), dolomite rock (also kn ...
s
belonging to the
Süntel Formation, that ranges in age from the early
Oxfordian to late
Kimmeridgian
In the geologic timescale, the Kimmeridgian is an age in the Late Jurassic Epoch and a stage in the Upper Jurassic Series. It spans the time between 154.8 ±0.8 Ma and 149.2 ±0.7 Ma (million years ago). The Kimmeridgian follows the Oxfordian ...
stages and have been deposited in a shallow sea with a water depth of less than . The layers exposed in the quarry are oriented nearly vertically and slightly overturned, which is a result of the ascent of the adjacent
Harz mountains
The Harz (), also called the Harz Mountains, is a Mittelgebirge, highland area in northern Germany. It has the highest elevations for that region, and its rugged terrain extends across parts of Lower Saxony, Saxony-Anhalt, and Thuringia. The nam ...
during the
Lower Cretaceous
Lower may refer to:
* ''Lower'' (album), 2025 album by Benjamin Booker
* Lower (surname)
* Lower Township, New Jersey
*Lower Receiver (firearms)
* Lower Wick Gloucestershire, England
See also
* Nizhny
{{Disambiguation ...
. Widely known as a classical exposure among geologists, the quarry had been extensively studied, and visited by students of geology for decades. Although rich in fossils of marine invertebrates, fossils of land-living animals had been rare. The sauropod tooth was the first specimen of a sauropod dinosaur from the Jurassic of northern Germany.
After more fossil material was found, including bones, excavation of the bone-bearing layer commenced in April 1999, conducted by a local association of private fossil collectors. Although the quarry operator was cooperative, excavation was complicated by the near-vertical orientation of the layers that limited access, as well as by the ongoing quarrying. The sauropod material could not be excavated directly from the layer but had to be collected from lose blocks that resulting from blasting. The exact origin of the bone material was therefore unclear, but could later be traced to a single bed (bed 83).
An excavation conducted between July 20–28 of 2000 rescued ca. of bone-bearing blocks containing vertebrate remains.
Fossils were prepared and stored in the
Dinosaur Park Münchehagen (DFMMh), a private dinosaur open-air museum located close to
Hanover
Hanover ( ; ; ) is the capital and largest city of the States of Germany, German state of Lower Saxony. Its population of 535,932 (2021) makes it the List of cities in Germany by population, 13th-largest city in Germany as well as the fourth-l ...
. Due to the very good preservation of the bones, consolidating agents had to be applied only occasionally, and preparation could be conducted comparatively quickly as bone would separate easily from the surrounding rock. Bones of simple shape could sometimes be prepared in less than an hour, while the preparation of a
sacrum
The sacrum (: sacra or sacrums), in human anatomy, is a triangular bone at the base of the spine that forms by the fusing of the sacral vertebrae (S1S5) between ages 18 and 30.
The sacrum situates at the upper, back part of the pelvic cavity, ...
required a workload of three weeks. By January 2001, 200 single vertebrate bones had already been prepared. At this point, the highest bone density was found in a block measuring 70 x 70 cm, which contained ca. 20 bones.
By January 2002, preparation of an even larger block had revealed a partial sauropod skull – the first to be discovered in Europe. Before complete removal of the bones from the block, a silicon cast was made of the block to document the precise three-dimensional position of the individual bones.
Part of the ''Europasaurus'' fossil material got damaged or destroyed by arson fire in the night from the 4th to the 5th of October, 2003. The fire destroyed the laboratory and exhibition hall of the Dinosaur Park Münchehagen, resulting in the loss of 106 bones, which account for 15% of the bones prepared at the time. Furthermore, the fire affected most of the still unprepared blocks, with firefighting water hitting the hot stone causing additional crumbling.
Destroyed specimens include DFMMh/FV 100, which included the best preserved vertebral series and the only complete pelvis.

In
2006
2006 was designated as the International Year of Deserts and Desertification.
Events
January
* January 1– 4 – Russia temporarily cuts shipment of natural gas to Ukraine during a price dispute.
* January 12 – A stampede during t ...
, the new sauropod
taxon
In biology, a taxon (back-formation from ''taxonomy''; : taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular name and ...
was formally described as ''Europasaurus holgeri''. The given
etymology
Etymology ( ) is the study of the origin and evolution of words—including their constituent units of sound and meaning—across time. In the 21st century a subfield within linguistics, etymology has become a more rigorously scientific study. ...
for the
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 ...
name is "reptile from Europe", and the specific name honours Holger Lüdtke, the discoverer of the first fossils.
Given the comparatively small size of the bones, it was initially assumed that they stem from juvenile individuals. The 2006 publication, however, established that the majority of specimens were adult, and that ''Europasaurus'' was an island dwarf.
The number of individual sauropod bones had increased to 650 and include variously articulated individuals; the material was found within an area of squared.
From these specimens, 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 ...
was selected, a disarticulated but associated individual (DFMMh/FV 291).
The holotype includes multiple
cranial
Standard anatomical terms of location are used to describe unambiguously the anatomy of humans and other animals. The terms, typically derived from Latin or Greek language, Greek roots, describe something in its standard anatomical position. Thi ...
bones (
premaxilla
The premaxilla (or praemaxilla) is one of a pair of small cranial bones at the very tip of the upper jaw of many animals, usually, but not always, bearing teeth. In humans, they are fused with the maxilla. The "premaxilla" of therian mammals h ...
,
maxilla
In vertebrates, the maxilla (: maxillae ) is the upper fixed (not fixed in Neopterygii) bone of the jaw formed from the fusion of two maxillary bones. In humans, the upper jaw includes the hard palate in the front of the mouth. The two maxil ...
and
quadratojugal The quadratojugal is a skull bone present in many vertebrates, including some living reptiles and amphibians.
Anatomy and function
In animals with a quadratojugal bone, it is typically found connected to the jugal (cheek) bone from the front and ...
), a partial
braincase
In human anatomy, the neurocranium, also known as the braincase, brainpan, brain-pan, or brainbox, is the upper and back part of the skull, which forms a protective case around the brain. In the human skull, the neurocranium includes the calv ...
, multiple
mandible
In jawed vertebrates, the mandible (from the Latin ''mandibula'', 'for chewing'), lower jaw, or jawbone is a bone that makes up the lowerand typically more mobilecomponent of the mouth (the upper jaw being known as the maxilla).
The jawbone i ...
bones (
dentary
In jawed vertebrates, the mandible (from the Latin ''mandibula'', 'for chewing'), lower jaw, or jawbone is a bone that makes up the lowerand typically more mobilecomponent of the mouth (the upper jaw being known as the maxilla).
The jawbone ...
,
surangular
The surangular or suprangular is a jaw bone found in most land vertebrates, except mammals. Usually in the back of the jaw, on the upper edge, it is connected to all other jaw bones: dentary, angular bone, angular, splenial and articular. It is o ...
and
angular), large amounts of teeth,
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 ...
,
sacral vertebrae
The sacrum (: sacra or sacrums), in human anatomy, is a triangular bone at the base of the spine that forms by the fusing of the sacral vertebrae (S1S5) between ages 18 and 30.
The sacrum situates at the upper, back part of the pelvic cavity, ...
and
ribs
The rib cage or thoracic cage is an endoskeletal enclosure in the thorax of most vertebrates that comprises the ribs, vertebral column and sternum, which protect the vital organs of the thoracic cavity, such as the heart, lungs and great vessels ...
from the neck and torso. At least 10 other individuals were referred to the same taxon based on overlap in material.
A large-scale excavation campaign commenced in the summer of 2012, with the goal to excavate ''Europasaurus'' bones not only from lose blocks but directly from the rock layer. Access to the bone-bearing layer required the removal of some 600 tons of rock using excavators and wheel loaders, and the constant pumping out of water from the base of the quarry. Excavations continued in spring and summer 2013. The campaign resulted in the discovery of new fish, turtle, and crocodile remains, as well as valuable information of the bone-bearing layer; additional ''Europasaurus'' bones, however, could not be located. By 2014, around 1300 vertebrate bones had been prepared from bed 83, the majority of which stemming from ''Europasaurus''; an estimated 3000 additional bones await preparation. A minimum number of 20 individuals was identified based on jaw bones.
Description
''Europasaurus'' is a very small sauropod, measuring only long and weighed as an adult.
This length was estimated based on a partial femur, scaled to the size of a nearly complete ''
Camarasaurus
''Camarasaurus'' ( ) is a genus of sauropod dinosaur that lived in North America during the Late Jurassic period. Its fossil remains have been found in the Morrison Formation, dating to the Kimmeridgian and Tithonian ages of the Jurassic, betwe ...
'' specimen. Younger individuals are known, from sizes of to the youngest juvenile at .
Distinguishing characteristics

Aside from being a very small
neosauropod, ''Europasaurus'' was thought to have multiple unique morphological features to distinguish it from close relatives by its original describers, Sander ''et al.'' (2006
[). The nasal process of the premaxilla was thought to curve anteriorly while projecting upwards (now known to be preservational]), there is a notch on the upper surface of the centra
Centra is a convenience shop chain that operates throughout Ireland. The chain operates as a symbol group owned by Musgrave Group, the food wholesaler, meaning the individual shops are all owned by individual franchisees.
The chain has three ...
of cervical vertebrae, 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 ...
has a prominent process on the posterior surface of its body, and the astragalus
Astragalus may refer to:
* ''Astragalus'' (plant), a large genus of herbs and small shrubs
*Astragalus (bone)
The talus (; Latin for ankle or ankle bone; : tali), talus bone, astragalus (), or ankle bone is one of the group of foot bones known ...
(an ankle bone) is twice as wide as tall.
When compared to ''Camarasaurus'', ''Europasaurus'' has a different morphology of the postorbital
The ''postorbital'' is one of the bones in vertebrate skulls which forms a portion of the dermal skull roof and, sometimes, a ring about the orbit. Generally, it is located behind the postfrontal and posteriorly to the orbital fenestra. In some ve ...
where the posterior flange is not as short, a short contact between the nasal
Nasal is an adjective referring to the nose, part of human or animal anatomy. It may also be shorthand for the following uses in combination:
* With reference to the human nose:
** Nasal administration, a method of pharmaceutical drug delivery
* ...
and frontal bone
In the human skull, the frontal bone or sincipital bone is an unpaired bone which consists of two portions.'' Gray's Anatomy'' (1918) These are the vertically oriented squamous part, and the horizontally oriented orbital part, making up the bo ...
s of the skull, the shape of its parietal (rectangular in ''Europasaurus''), and the neural spines of its vertebrae in front of the pelvis
The pelvis (: pelves or pelvises) is the lower part of an Anatomy, anatomical Trunk (anatomy), trunk, between the human abdomen, abdomen and the thighs (sometimes also called pelvic region), together with its embedded skeleton (sometimes also c ...
are unsplit. Comparisons with ''Brachiosaurus
''Brachiosaurus'' () is a genus of sauropod dinosaur that lived in North America during the Late Jurassic, about . It was first Species description, described by American paleontologist Elmer S. Riggs in 1903 in paleontology, 1903 from fossi ...
'' (now named ''Giraffatitan
''Giraffatitan'' (name meaning "titanic giraffe") is a genus of sauropod dinosaur that lived during the late Jurassic Period (geology), Period (Kimmeridgian–Tithonian stages) in what is now Lindi Region, Tanzania. Only one species is known, ...
'') were also mentioned, and it was identified that ''Europasaurus'' has a shorter snout, a contact between the quadratojugal and squamosal
The squamosal is a skull bone found in most reptiles, amphibians, and birds. In fishes, it is also called the pterotic bone.
In most tetrapods, the squamosal and quadratojugal bones form the cheek series of the skull. The bone forms an ancestra ...
, and a humerus
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 (bone), radius and ulna, and consists of three sections. The humeral upper extrem ...
(upper forelimb bone) that has flattened and aligned proximal
Standard anatomical terms of location are used to describe unambiguously the anatomy of humans and other animals. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position prov ...
and distal
Standard anatomical terms of location are used to describe unambiguously the anatomy of humans and other animals. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position provi ...
surfaces. There were finally quick comparisons to the potential brachiosaurid '' Lusotitan'', which has a different ilium and astragalus shape, and '' Cetiosaurus humerocristatus'' (named ''Duriatitan
''Duriatitan'' is a genus of titanosauriform sauropod dinosaur that lived in the Late Jurassic in what is now England. The holotype specimen of ''Duriatitan'', BMNH 44635, is a partial left upper arm bone which was found by R.I. Smith nea ...
''), which has a deltopectoral crest that is less prominent and extends across less of the humerus.
Skull
Nearly all external skull bones have been preserved among ''Europasaurus'' specimens, except the prefrontals. Some additional bones are only represented by very fragmented and uninformative fossils, such as the lacrimals. Eight premaxillae are known, with a generally rectangular snout shape as found in ''Camarasaurus''. The anterior projection of the premaxilla identified in Sander ''et al.'' (2006[) was re-identified as a preservational artifact in Marpmann ''et al.'' (2014][), similar to the anatomy found in ''Camarasaurus'' and '' Euhelopus'' to a lesser degree. The ]dorsal
Dorsal (from Latin ''dorsum'' ‘back’) may refer to:
* Dorsal (anatomy), an anatomical term of location referring to the back or upper side of an organism or parts of an organism
* Dorsal, positioned on top of an aircraft's fuselage
The fus ...
projection of the premaxilla, the one which contacts the nasal bone, begins as a postero-dorsal projection, before becoming straight vertical at the point of the subnarial foramen
In anatomy and osteology, a foramen (; : foramina, or foramens ; ) is an opening or enclosed gap within the dense connective tissue (bones and deep fasciae) of extant and extinct amniote animals, typically to allow passage of nerves, artery, ...
, until it reaches the nasal. This weak "step" is seen in ''Camarasaurus'' and ''Euhelopus'', and is present more strongly in '' Abydosaurus'', ''Giraffatitan'' and a possible skull of ''Brachiosaurus''. These latter taxa also have a longer snout, with more distance from the first tooth until the nasal process of the premaxilla. As well, ''Europasaurus'' shares with the basal camarasauromorphs (brachiosaurids, ''Camarasaurus'', ''Euhelopus'' and '' Malawisaurus'') a similarly sized orbit
In celestial mechanics, an orbit (also known as orbital revolution) is the curved trajectory of an object such as the trajectory of a planet around a star, or of a natural satellite around a planet, or of an artificial satellite around an ...
and nasal fenestra
The skull, or cranium, is typically a bony enclosure around the brain of a vertebrate. In some fish, and amphibians, the skull is of cartilage. The skull is at the head end of the vertebrate.
In the human, the skull comprises two prominent ...
, whereas the nasal opening is significantly reduced in derived titanosaurs ('' Rapetosaurus'', '' Tapuiasaurus'' and '' Nemegtosaurus'').
A single maxilla is present in the well-preserved material of ''Europasaurus'', DFMMh/FV 291.17. This maxilla has a long body, with two elongate processes, a nasal and a posterior process. There is only a weak lacrimal process, like in most sauropods except ''Rapetosaurus''. The nasal process is elongate and covers the anterior and ventral rim of the antorbital fenestra
An antorbital fenestra (plural: fenestrae) is an opening in the skull that is in front of the eye sockets. This skull character is largely associated with Archosauriformes, archosauriforms, first appearing during the Triassic Period. Among Extant ...
. This process extends about 120º from the horizontal tooth row. The base of the nasal process also forms the body of the lacrimal process, and at their divergence is the antorbital fenestra, similar in shape to those of ''Camarasaurus'', ''Euhelopus'', ''Abydosaurus'' and ''Giraffatitan'', but about 1/2 taller proportionally. The pre-antorbital fenestra, a small opening in front of or beneath the antorbital opening, is well developed in taxa like ''Diplodocus
''Diplodocus'' (, , or ) is an extinct genus of diplodocid sauropod dinosaurs known from the Late Jurassic of North America. The first fossils of ''Diplodocus'' were discovered in 1877 by S. W. Williston. The generic name, coined by Othnie ...
'' and ''Tapuiasaurus'', is nearly absent, like in ''Camarasaurus'' and ''Euhelopus''. There were about 12–13 total teeth in the maxilla of ''Europasaurus'', fewer than in more basal taxa (16 teeth in ''Jobaria
''Jobaria'' is a genus of sauropod dinosaur that lived in what is now Niger during the middle Jurassic Period, between 164 and 161 million years ago. ''Jobaria'' is currently the only known valid sauropod from the Tiouraren, where it was disco ...
'' and 14–25 in '' Atlasaurus''), but falling within the range of variation in Brachiosauridae (15 in ''Brachiosaurus'' to 10 in ''Abydosaurus''). All of the unworn teeth preserved display up to four small denticles on their mesial
This is a list of definitions of commonly used terms of location and direction in dentistry. This set of terms provides orientation within the oral cavity, much as anatomical terms of location provide orientation throughout the body.
Terms
...
edges. A small amount of the posterior tooth crowns are slightly twisted (~15º), but much less than in brachiosaurids (30–45º).
Among the nasal bones of ''Europasaurus'', several are known, but few are complete or undistorted. The nasals are overlapped posteriorly by the frontal bones, and towards the side, they articulate bluntly with the prefrontals. Unlike the nasals of ''Giraffatitan'', those in ''Europasaurus'' project horizontally forwards, forming a small portion of the skull roof over the antorbital fenestrae. Four frontals are known from ''Europasaurus'', three being from the left and one being from the right. Because of their disarticulation, it is likely that the frontals never fused during growth, unlike in ''Camarasaurus''. The frontals form a portion of the skull roof, articulating with other bones such as the nasals, parietals, prefrontals and postorbitals, and they are longer antero-posteriorly than they are wide, a unique character among a eusauropoda
Eusauropoda (meaning "True Lizard Foot") is a derived clade of sauropod dinosaurs. Eusauropods represent the node-based group that includes all descendant sauropods starting with the basal eusauropods of '' Shunosaurus'', and possibly '' Barapas ...
n. Like in diplodocoids (''Amargasaurus
''Amargasaurus'' (; "La Amarga lizard") is a genus of sauropod dinosaur from the Early Cretaceous epoch (geology), epoch (129.4–122.46 Mya (unit), mya) of what is now Argentina. The only known skeleton was discovered in 1984 and is virtually ...
'', '' Dicraeosaurus'' and ''Diplodocus''), as well as ''Camarasaurus'', the frontals are excluded from the frontoparietal fenestra (or parietal fenestra when frontals are excluded). The frontals are also excluded from the supratemporal fenestra
Temporal fenestrae are openings in the temporal region of the skull of some amniotes, behind the orbit (eye socket). These openings have historically been used to track the evolution and affinities of reptiles. Temporal fenestrae are commonly (al ...
margin (a widespread character in sauropods more derived than ''Shunosaurus
''Shunosaurus'', meaning "Lizard from Sichuan", is a genus of sauropod dinosaur from Late Jurassic ( Oxfordian) beds in Sichuan Province in China, from 161 to 157 Million years ago. The name derives from "Shu", an ancient name for the Sichuan pr ...
''), and they only have a small, unornamented participation in the orbit
In celestial mechanics, an orbit (also known as orbital revolution) is the curved trajectory of an object such as the trajectory of a planet around a star, or of a natural satellite around a planet, or of an artificial satellite around an ...
. Several parietal bones are known in ''Europasaurus'', which show a rectangular shape much wider than long. the parietals are also wide when viewed from the back of the skull, being slightly taller than the foramen magnum
The foramen magnum () is a large, oval-shaped opening in the occipital bone of the skull. It is one of the several oval or circular openings (foramina) in the base of the skull. The spinal cord, an extension of the medulla oblongata, passes thro ...
(spinal cord opening). The parietals contribute to about half the post-temporal fenestra (opening above the very back of the skull) border, with the other region enclosed by the squamosal bone
The squamosal is a skull bone found in most reptiles, amphibians, and birds. In fishes, it is also called the pterotic bone.
In most tetrapods, the squamosal and quadratojugal bones form the cheek series of the skull. The bone forms an ancestral ...
s and some braincase bones. Parietals also form part of the edge of the supratemporal fenestra, which is wider than long in ''Europasaurus'', like in ''Giraffatitan'', ''Camarasaurus'' and ''Spinophorosaurus
''Spinophorosaurus'' is a genus of sauropod dinosaur that lived in what is now Niger during the Middle Jurassic period. The first two specimens were excavated in the 2000s by German and Spanish teams under difficult conditions. The skeletons were ...
''. Besides the before mentioned fenestra, the parietals also have a "postparietal fenestra", something rarely seen outside of Dicraeosauridae
Dicraeosauridae is a Family (biology), family of Diplodocoidea, diplodocoid sauropods who are the sister group to Diplodocidae. Dicraeosaurids are a part of the Flagellicaudata, along with Diplodocidae. Dicraeosauridae includes genera such as ''A ...
. A triradiate postorbital bone is present in ''Europasaurus'', which evolved as the fusion of the postfrontal and postosbital bone of more basal taxa. Between the anterior and ventrally projecting processes the postorbital forms the margin of the orbit, and between the posterior and ventral processes it borders the infratemporal fenestra.
Multiple jugals are known from ''Europasaurus'', which are more similar in morphology to basal sauropodomorph
Sauropodomorpha ( ; from Greek, meaning "lizard-footed forms") is an extinct clade of long-necked, herbivorous, saurischian dinosaurs that includes the sauropods and their ancestral relatives. Sauropods generally grew to very large sizes, had lo ...
s than other macronarians. It forms part of the border of the orbit, infratemporal fenestra and the bottom edge of the skull, but does not reach the antorbital fenestra. The posterior process of the jugal are very fragile and narrow, showing a bone scar from the articulation with the quadratojugal. There are two prominences projecting from the back of the jugal body, which diverge at 75º and form the bottom and front edges of the infratemporal fenestra. Like in ''Riojasaurus
''Riojasaurus'' (meaning "lizard from La Rioja") was a herbivorous sauropodomorph dinosaur named after La Rioja Province in Argentina where it was found in the Los Colorados Formation in the Ischigualasto-Villa Unión Basin by José Bonaparte. ...
'' and ''Massospondylus
''Massospondylus'' ( ) is a genus of sauropodomorph dinosaur from the Early Jurassic of southern Africa. It was described by Sir Richard Owen in 1854 from remains discovered in South Africa, and is thus one of the first dinosaurs to have been ...
'', two non-sauropod sauropodomorphs, the jugal forms a large part of the orbit edge, from the back to the front bottom corner. This feature has been seen in embryo
An embryo ( ) is the initial stage of development for a multicellular organism. In organisms that reproduce sexually, embryonic development is the part of the life cycle that begins just after fertilization of the female egg cell by the male sp ...
s of titanosaurs, but no adult individuals. The quadratojugal bone The quadratojugal is a skull bone present in many vertebrates, including some living reptiles and amphibians.
Anatomy and function
In animals with a quadratojugal bone, it is typically found connected to the jugal (cheek) bone from the front and ...
is an elongate element that has two projecting arms, one anterior and one dorsal. Like in other sauropods, the anterior process is longer than the dorsal, but in ''Europasaurus'' the arms are more similar lengths. The horizontal process is parallel to the tooth row of ''Europasaurus'', similar to in ''Camarasaurus'' but unlike in ''Giraffatitan'' and ''Abydosaurus''. There is a prominent ventral flange on the anterior arm of the bone, which is a possible synapomorphy
In phylogenetics, an apomorphy (or derived trait) is a novel Phenotypic trait, character or character state that has evolution, evolved from its ancestral form (or Plesiomorphy and symplesiomorphy, plesiomorphy). A synapomorphy is an apomorphy sh ...
of Brachiosauridae, although it is also found in some ''Camarasaurus'' individuals. The two quadratojugal processes diverge at a nearly right angle (90º), although the dorsal process curves as it follows the shape of the quadrate. Squamosals found from ''Europasaurus'' show the same approximate shape in lateral view as ''Camarasaurus'', that of a question mark
The question mark (also known as interrogation point, query, or eroteme in journalism) is a punctuation, punctuation mark that indicates a question or interrogative clause or phrase in many languages.
History
The history of the question mark is ...
. The squamosals articulate with many skull bones, including those of the skull roof, those of the ventral skull, and those of the braincase. Like the postorbitals, the squamosals are triradiate, with a ventral, anterior and medial
Medial may refer to:
Mathematics
* Medial magma, a mathematical identity in algebra Geometry
* Medial axis, in geometry the set of all points having more than one closest point on an object's boundary
* Medial graph, another graph that repr ...
process.
There are thirteen preserved elements of the palate
The palate () is the roof of the mouth in humans and other mammals. It separates the oral cavity from the nasal cavity.
A similar structure is found in crocodilians, but in most other tetrapods, the oral and nasal cavities are not truly sep ...
of ''Europasaurus'', including the quadrate, pterygoid and ectopterygoid. The quadrates articulate with the palate and braincase bones, as well as the external skull bones. They are similar in shape to those of ''Giraffatitan'' and ''Camarasaurus'', and have well-developed articular surfaces. A single shaft is present for a majority of the quadrates length, with a pterygoid wing along the medial side. Pterygoids are the largest of the sauropod palate bones, and it has a triradiate shape, like the postorbitals. An anterior projection contacts the opposite pterygoid, while a lateral wing contacts the ectopterygoid, and a posterior wing supports the quadrate and basipterygoid (a bone that provides connection between the palate and the braincase). The ectopterygoid is a small palate bone, which articulates the central palate bones (pterygoid and palatine
A palatine or palatinus (Latin; : ''palatini''; cf. derivative spellings below) is a high-level official attached to imperial or royal courts in Europe since Roman Empire, Roman times. ) with the maxilla. Ectopterygoids are L-shaped, with an anterior process attaching to the maxilla, and a dorsal process that meets the pterygoid.
Vertebrae
The cervical vertebrae of ''Europasaurus'' are the best preserved and most represented of the vertebral column. However, not the entire neck is known, so the cervical number could be between ''Camarasaurus'' (12 vertebrae) and ''Rapetosaurus'' (17 vertebrae). Additionally, the multiple cervical vertebrae come from different-aged individuals, and the centrum length and internal structure are known to change throughout development. The adult cervical centra are elongated and (anterior end is ball-shaped), with a notch in the top of the rear end of the centrum. This feature was described as characteristic of ''Europasaurus'' but is also known in ''Euhelopus'' and ''Giraffatitan''. In the side of the centra of ''Europasaurus'' there is an excavation which opens into the internal of the vertebrae. Unlike in ''Giraffatitan'' and brachiosaurids, ''Europasaurus'' does not have thin ridges () dividing this opening. ''Europasaurus'' shares laminae features on the upper vertebrae with basal macronarians and brachiosaurids. Differing from the anterior and middle cervicals, the posterior cervical vertebrae are less elongate, and taller proportionally, like in other macronarians, with significant changes in the positions of articular surfaces.
Front dorsal vertebrae are strongly opisthocoelous like the cervicals, and can be placed in the series based on the absence of the and low placement. The internal structures are open and like ''Camarasaurus'', ''Giraffatitan'' and '' Galvesaurus'', but unlike these taxa this pneumaticity does not extend into the middle and posterior dorsal vertebrae. The arrangement and presence of anterior laminae in ''Europasaurus'' is similar to other basal macronarians, but unlike more basal taxa (e.g. '' Mamenchisaurus'', '' Haplocanthosaurus'') and more derived taxa (e.g. ''Giraffatitan''). The middle dorsals possess a pneumatic cavity that extends upwards into the , like in ''Barapasaurus
''Barapasaurus'' ( ) is a genus of Basal (phylogenetics), basal sauropod dinosaur from Jurassic rocks of India. The only species is ''B. tagorei''. ''Barapasaurus'' comes from the lower part of the Kota Formation, which is of Early to Middle J ...
'', ''Cetiosaurus
''Cetiosaurus'' ( meaning 'whale lizard', from the Greek '/ meaning 'sea monster' (later, 'whale') and '/ meaning 'lizard'), is a genus of herbivorous sauropod dinosaur from the Middle Jurassic Period, living about 171 to 165 million years ago ...
'', '' Tehuelchesaurus'', and ''Camarasaurus''. The ventral edge of this opening is rhomboidal and well-defined. In the posterior vertebrae, the lateral pneumatic cavity has shifted higher on the centrum, a change seen in other basal macronarians. These are wide anteriorly, and narrow to become acutely angled posteriorly. The of ''Europasaurus'' stands vertically, a basal feature not seen in ''Brachiosaurus'' or more derived sauropods.
A series of all complete is only known from a single specimen, DFMMh/FV 100, which was destroyed in a fire in 2003. All five vertebrae, the characteristic number of more basal neosauropods, are incorporated into the . The third and fourth sacrals represent the primordial sacrals, present in all dinosaur groups. The second, S2, is the ancestral sauropodomorph sacral that was added in basal sauropodomorphs, who all share three sacrals to the exception of ''Plateosaurus
''Plateosaurus'' (probably meaning "broad lizard", often mistranslated as "flat lizard") is a genus of plateosaurid dinosaur that lived during the Late Triassic period, around 214 to 204 million years ago, in what is now Central and Northern Eu ...
''. The fifth sacral, fused behind the primordial pair, is a caudosacral, migrated from the tail into the pelvis in taxa around '' Leonerasaurus''. The first sacral, articulated with the ilium but not fused to the other vertebrae, represents the dorsosacral, bringing the count to five vertebrae found in all neosauropods. The level of fusion of the dorsosacral confirms the evolutionary history of the sauropod sacral count: the primordial pair incorporating first a dorsal (total of three), then a caudal (total of four), then another dorsal to make a total of five vertebrae.
Skin
Among macronarians, fossilized skin impressions are only known from ''Haestasaurus
''Haestasaurus'' is a genus of herbivorous sauropod dinosaur, belonging to the Macronaria, that during the Early Cretaceous lived in the area of present-day England. The only species is ''Haestasaurus becklesii''.Upchurch P., Mannion P.D., Taylor ...
'', '' Tehuelchesaurus'' and ''Saltasaurus
''Saltasaurus'' (which means "lizard from Salta") is a genus of saltasaurid dinosaur of the Late Cretaceous Period (geology), period of Argentina. Small among sauropods, though still heavy by the standards of modern creatures, ''Saltasaurus'' was ...
''. Both ''Tehuelchesaurus'' and ''Haestasaurus'' may be closely related to ''Europasaurus'', and the characteristics of all sauropod skin impressions are similar. ''Haestasaurus'', the first dinosaur known from skin impressions, preserved integument
In biology, an integument is the tissue surrounding an organism's body or an organ within, such as skin, a husk, Exoskeleton, shell, germ or Peel (fruit), rind.
Etymology
The term is derived from ''integumentum'', which is Latin for "a coverin ...
over a portion of the arm around the elbow joint. Dermal impressions are more widespread in the material of ''Tehuelchesaurus'', where they are known from the areas of the forelimb, scapula and torso. There are no bony plates or nodules, to indicate armour, but there are several types of scales. The skin types of ''Tehuelchesaurus'' are overall more similar to those found in diplodocids and ''Haestasaurus'' than in the titanosaur embryos of Auca Mahuevo. As the shape and articulation of the preserved tubercles in these basal macronarians are similar in other taxa where skin is preserved, including specimens of '' Brontosaurus excelsus'' and intermediate diplodocoids, such dermal structures are probably widespread throughout Neosauropoda.
Classification
When it was first named, ''Europasaurus'' was considered to be a taxon within Macronaria that didn't fall within the 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 ...
Brachiosauridae or the clade
In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
Titanosauromorpha. This indicated that the dwarfism of the taxon was a result of evolution, instead of being a characteristic of a group. Three matrices were analysed with the inclusion of ''Europasaurus'', that of Wilson (2002) and Upchurch (1998) and Upchurch ''et al.'' (2004). All analyses resulted in similar phylogenetics, where ''Europasaurus'' placed more derived than ''Camarasaurus'' but outside a clade of Brachiosauridae and Titanosauromorpha (now named Titanosauriformes). The results of the favoured analysis of Sander ''et al.'' (2006) are shown below on the left:
During a description of the vertebrae of ''Europasaurus'' by Carballido & Sander (2013), another phylogenetic analysis was conducted (right column above). The cladistic matrix was expanded to include more sauropod taxa, such as '' Bellusaurus'', '' Cedarosaurus'' and ''Tapuiasaurus''. The taxon ''Brachiosaurus'' was also separated into true ''Brachiosaurus'' (''B. altithorax'') and ''Giraffatitan'' (''B. brancai''), based on Taylor
Taylor, Taylors or Taylor's may refer to:
People
* Taylor (surname)
** List of people with surname Taylor
* Taylor (given name), including Tayla and Taylah
* Taylor sept, a branch of Scottish clan Cameron
* Justice Taylor (disambiguation)
...
(2009). Based on this newer and more expansive analysis, ''Europasaurus'' was found to be in a similar placement, as a basal camarasauromorph closer to titanosaurs than ''Camarasaurus''. However, ''Euhelopus'', ''Tehuelchesaurus'', '' Tastavinsaurus'' and '' Galvesaurus'' were placed between ''Europasaurus'' and Brachiosauridae.
Placement as a brachiosaurid
In a 2012 analysis of the phylogeny of Titanosauriformes, D'Emic (2012) considered ''Europasaurus'' to belong to Brachiosauridae, instead of being basal to the earliest brachiosaurids. The phylogeny resolved the most true brachiosaurids to date, although several potential brachiosaurids were instead determined to belong to Somphospondyli ('' Paluxysaurus'', ''Sauroposeidon
''Sauroposeidon'' ( ; meaning "lizard earthquake deity, god", after the Greek god Poseidon) is a genus of sauropod dinosaur known from several incomplete specimens including a bone bed and fossilized trackways that have been found in the U.S. st ...
'' and ''Qiaowanlong
''Qiaowanlong'' (meaning "Qiaowan dragon") is an extinct genus of sauropod dinosaur. Fossils belonging to the genus were found in the Yujinzi Basin of what is today Gansu Province, China. The remains come from a geological formation called the X ...
''). However, D'Emic was tentative in considering ''Europasaurus'' to be a confirmed brachiosaurid. While there was strong support in the phylogeny for its placement, ''Europasaurus'', one of few basal macronarians with a skull, lacks multiple bones that display characteristic features of the group, such as caudal vertebrae. The cladogram below on the left illustrates the phylogenetic results of D'Emic (2012), with Euhelopodidae
Euhelopodidae is a family of sauropod dinosaurs of disputed membership and affinities, which contains '' Euhelopus'' and its close relatives. Most proposed euhelopodids are from East Asia.
Euhelopodidae was first recognized by Carl Wiman in 1929 ...
and Titanosauria collapsed.
A later analysis on titanosauriformes agreed with D'Emic (2012) in the placement of ''Europasaurus''. It formed a polytomy
An internal node of a phylogenetic tree is described as a polytomy or multifurcation if (i) it is in a rooted tree and is linked to three or more child subtrees or (ii) it is in an unrooted tree and is attached to four or more branches. A tree ...
with ''Brachiosaurus'' and the "French ''Bothriospondylus
''Bothriospondylus'', from Ancient Greek βοθρίον (''bothríon''), meaning "trench", and σπόνδυλος (''spóndulos''), meaning "vertebra", is a nomen dubium, dubious genus of neosauropod sauropod dinosaur. It lived during the Late Ju ...
''" (named '' Vouivria'') as the basalmost brachiosaurids. Next most derived in the clade was '' Lusotitan'', with ''Giraffatitan'', ''Abydosaurus'', ''Cedarosaurus'' and '' Venenosaurus'' forming a more derived clade of brachiosaurids. The "twisted" teeth of ''Europasaurus'' were found to be one of the unique features of Brachiosauridae, which could mean a confident referral of isolated sauropod teeth to the clade.[ ]
A further phylogenetic analysis was performed on Brachiosauridae, based on that of D'Emic (2012). This phylogeny, conducted by D'Emic ''et al.'' (2016), resolved a very similar placement of ''Europasaurus'' within Brachiosauridae, although '' Sonorasaurus'' was placed in a clade with ''Giraffatitan'', and ''Lusotitan'' was placed in a polytomy with ''Abydosaurus'' and ''Cedarosaurus''. The remaining tree was the same as in D'Emic (2012), although ''Brachiosaurus'' was collapsed into a polytomy with more derived brachiosaurids. Another phylogeny, Mannion ''et al.'' (2017) found similar results to D'Emic (2012) and D'Emic ''et al.'' (2016). ''Europasaurus'' was the basalmost brachiosaurid, with the "French Bothriospondylus", or ''Vouivria'', as the next most basal brachiosaurid. ''Brachiosaurus'' was placed outside of a poltomy of all other brachiosaurids, ''Giraffatitan'', ''Abydosaurus'', ''Sonorasaurus'', ''Cedarosaurus'' and ''Venenosaurus''. A 2017 phylogeny, that of Royo-Torres ''et al.'' (2017), resolved more complex relations within Brachiosauridae. Besides ''Europasaurus'' as the basalmost brachiosaurid, there were two subgroups within the clade, one containing ''Giraffatitan'', ''Sonorasaurus'' and ''Lusotitan'', and another including almost all other brachiosaurids, as well as ''Tastavinsaurus''. This second clade would be termed Laurasiformes under the group's definition. ''Brachiosaurus'' was in a polytomy with the two subclades of Brachiosauridae. The phylogeny of Royo-Torres ''et al.'' (2017) can be seen above, in the right column.
Paleobiology
Growth
It was identified that ''Europasaurus'' was a unique dwarf species, and not a juvenile of an existing taxon like ''Camarasaurus'', by a histology
Histology,
also known as microscopic anatomy or microanatomy, is the branch of biology that studies the microscopic anatomy of biological tissue (biology), tissues. Histology is the microscopic counterpart to gross anatomy, which looks at large ...
analysis of multiple specimens of ''Europasaurus''. The youngest specimen (DFMMh/FV 009) was shown by this analysis to lack signs of aging such as growth marks or laminar bone tissue, and is also the smallest specimen at in length. Such bone tissue is an indicator of rapid growth, so the specimen is probably a young juvenile. A larger specimen (DFMMh/FV 291.9) at shows large amounts of laminar tissue, with no growth marks present, so is likely a juvenile as well. The next smallest specimen (DFMMh/FV 001) has shows the presence of growth marks (specifically annuli), and at the length of is possibly a subadult. Further larger, DFMMh/FV 495 displays mature osteons
In osteology, the osteon or haversian system (; named for Clopton Havers) is the fundamental functional unit of much compact bone. Osteons are roughly cylindrical structures that are typically between 0.25 mm and 0.35 mm in diameter. Their ...
as well as annuli, and is . The second largest analysed specimen (DFMMh/FV 153) also shows growth marks, but they are more frequent. This specimen is . A single partial femur represents the largest known ''Europasaurus'' individual, at a body length of . Unlike all other specimens, this one (DFMMh/FV 415) shows the presence of lines of arrested growth
Growth arrest lines, also known as Harris lines, are lines of increased bone density that represent the position of the growth plate at the time of insult to the organism and formed on long bones due to growth arrest. They are only visible by ra ...
, indicating it died after reaching full body size. The internal bone is also partially lamellar
A lamella (: lamellae) is a small plate or flake, from the Latin, and may also refer to collections of fine sheets of material held adjacent to one another in a gill-shaped structure, often with fluid in between though sometimes simply a set of ...
, which shows it had stopped growing recently.[
These combinations of growth factors show that ''Europasaurus'' developed its small size because of a largely reduced growth rate, gaining size slower than larger taxa such as ''Camarasaurus''. This slowing growth rate is the opposite of the general trend of sauropods and ]theropod
Theropoda (; from ancient Greek , (''therion'') "wild beast"; , (''pous, podos'') "foot"">wiktionary:ποδός"> (''pous, podos'') "foot" is one of the three major groups (clades) of dinosaurs, alongside Ornithischia and Sauropodom ...
s, who reached greater sizes with increased growth rates. Some of the close relatives of ''Europasaurus'' represent the largest dinosaurs known, including ''Brachiosaurus'' and ''Sauroposeidon''. Marpmann ''et al.'' (2014) proposed that the small size and reduced growth rate of ''Europasaurus'' was an effect of pedomorphism, where the adults of taxa retain juvenile characteristics, such as size.
Examinations of the inner ears of infant ''Europasaurus'' suggest they were precocial, and it is suggested that they would have been reliant on the protections of adults in a herd to some degree, something not seen in larger sauropods due to the massive size difference in parent and offspring. The structure and long length of the inner ear in this genus also suggests that they had good senses of hearing, with ''Europasaurus''. Intraspecific communication was also apparently important to this sauropod, based on these studies, suggesting this sauropod displayed clear, gregarious behavior.
Dwarfism
It has been suggested that an ancestor of ''Europasaurus'' would have quickly decreased in body size after emigrating to an island that existed at the time, as the largest of the islands in the region around northern Germany was smaller than squared, which may not have been able to support a community of large sauropods. Alternately, a macronarian may have shrunken concurrently with a larger landmass, until achieving the size of ''Europasaurus''. Previous studies on insular (island) dwarfism are largely restricted to 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 ...
of Haţeg Island in Romania
Romania is a country located at the crossroads of Central Europe, Central, Eastern Europe, Eastern and Southeast Europe. It borders Ukraine to the north and east, Hungary to the west, Serbia to the southwest, Bulgaria to the south, Moldova to ...
, home to the dwarf titanosaur ''Magyarosaurus'' and the dwarf hadrosaur
Hadrosaurids (), also hadrosaurs or duck-billed dinosaurs, are members of the ornithischian family Hadrosauridae. This group is known as the duck-billed dinosaurs for the flat duck-bill appearance of the bones in their snouts. The ornithopod fami ...
''Telmatosaurus
''Telmatosaurus'' (meaning "marsh lizard") is a genus of basal (phylogeny), basal hadrosauromorph dinosaur from the Late Cretaceous of Romania. It was a relatively small hadrosaur, measuring approximately in length and in body mass, which has b ...
''. ''Telmatosaurus'' is known to be from a small adult, and although it is very small, ''Magyarosaurus'' specimens of small sizes are known to be from adult to old individuals. ''Magyarosaurus dacus'' adults were a similar body size to ''Europasaurus'', but the largest of the latter had longer femora than the largest of the former, while ''Magyarosaurus hungaricus'' was significantly larger than either taxon. The dwarfism in ''Europasaurus'' represents the only significant rapid body mass change in derived Sauropodomorpha
Sauropodomorpha ( ; from Greek, meaning "lizard-footed forms") is an extinct clade of long-necked, herbivorous, saurischian dinosaurs that includes the sauropods and their ancestral relatives. Sauropods generally grew to very large sizes, had lo ...
, with the general trend of taxa being a growth in overall size in other groups.
Palaeoecology
The Langenberg locality in Germany, from the early Oxfordian to late Kimmeridgian, displays the variety of plant and animal life from an island ecosystem from the late Jurassic.[ During the Kimmeridgian the locality would have been marine, being located between the Rhenisch, Bohemian, and London-Brabant ]Massif
A massif () is a principal mountain mass, such as a compact portion of a mountain range, containing one or more summits (e.g. France's Massif Central). In mountaineering literature, ''massif'' is frequently used to denote the main mass of an ...
s. This does not indicate that the taxa present were marine, as the animals and plants may have been deposited allochthon
upright=1.6, Schematic overview of a thrust system. The hanging wall block is (when it has reasonable proportions) called a window. A klippe is a solitary outcrop of the nappe in the middle of autochthonous material.
An allochthon, or an alloc ...
ously (albeit only by a short distance) from the surrounding islands. The sediments to show that there was an occasional influx of fresh or brackish
Brackish water, sometimes termed brack water, is water occurring in a natural environment that has more salinity than freshwater, but not as much as seawater. It may result from mixing seawater (salt water) and fresh water together, as in estuari ...
water, and the fossils preserved display that. There are large numbers of marine bivalve
Bivalvia () or bivalves, in previous centuries referred to as the Lamellibranchiata and Pelecypoda, is a class (biology), class of aquatic animal, aquatic molluscs (marine and freshwater) that have laterally compressed soft bodies enclosed b ...
fossils, as well as echinoderm
An echinoderm () is any animal of the phylum Echinodermata (), which includes starfish, brittle stars, sea urchins, sand dollars and sea cucumbers, as well as the sessile sea lilies or "stone lilies". While bilaterally symmetrical as ...
s and microfossils present in the limestone of the quarry, although many of the animals
Animals are multicellular, eukaryotic organisms in the biological kingdom Animalia (). With few exceptions, animals consume organic material, breathe oxygen, have myocytes and are able to move, can reproduce sexually, and grow from a ...
and plants
Plants are the eukaryotes that form the kingdom Plantae; they are predominantly photosynthetic. This means that they obtain their energy from sunlight, using chloroplasts derived from endosymbiosis with cyanobacteria to produce sugars f ...
were terrestrial.
Many marine taxa are preserved at Langenberg, although they would not have co-existed often with ''Europasaurus''. There are at least three turtle
Turtles are reptiles of the order (biology), order Testudines, characterized by a special turtle shell, shell developed mainly from their ribs. Modern turtles are divided into two major groups, the Pleurodira (side necked turtles) and Crypt ...
genera, ''Plesiochelys
''Plesiochelys'' ("near turtle") is a genus of late Jurassic European and Asian turtle. The type species is ''Plesiochelys etalloni''.
Fossil records have discovered ''Plesiochelys bigleri'' and ''Plesiochelys etalloni'' from the Kimmeridge Clay ...
'', '' Thalassemys'' and up to two unnamed taxa. Actinopterygian
Actinopterygii (; ), members of which are known as ray-finned fish or actinopterygians, is a class of bony fish that comprise over 50% of living vertebrate species. They are so called because of their lightly built fins made of webbings of skin ...
fish are abundant, being represented by '' Lepidotes'', '' Macromesodon'', '' Proscinetes'', '' Coelodus'', '' Macrosemius'', '' Notagogus'', '' Histionotus'', '' Ionoscopus'', '' Callopterus'', ''Caturus
''Caturus'' (from , 'down' and 'tail') is an extinct genus of predatory marine fishes in the family Caturidae in the order Amiiformes, related to modern bowfin. It has been suggested that the genus is non-monophyletic with respect to other c ...
'', '' Sauropsis'', '' Belonostomus'', and '' Thrissops''. Also present are at least five distinct morphologies of hybodont sharks, the neoselachians '' Palaeoscyllium'', '' Synechodus'' and '' Asterodermus''. Two marine crocodyliform
Crocodyliformes is a clade of Crurotarsi, crurotarsan archosaurs, the group often traditionally referred to as "crocodilians". They are the first members of Crocodylomorpha to possess many of the features that define later relatives. They are the ...
s are known from Langenberg, '' Machimosaurus'' and '' Steneosaurus'', which likely fed off turtles and fish, and the amphibious crocodyliform '' Goniopholis'' has also been found.
Conifer
Conifers () are a group of conifer cone, cone-bearing Spermatophyte, seed plants, a subset of gymnosperms. Scientifically, they make up the phylum, division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a sin ...
cones and twigs can be identified as the araucarian ''Brachyphyllum
''Brachyphyllum'' (meaning "short leaf") is a form genus of fossil coniferous plant foliage. Plants of the genus have been variously assigned to several different conifer groups including Araucariaceae and Cheirolepidiaceae. They are known from ...
'', from the terrestrial fossils of the site. Deposited in the locality are many taxa, including a large accumulation of ''Europasaurus'' bones and individuals. At least 450 bones from ''Europasaurus'' were recovered from the Langenberg Quarry, with about 1/3 bearing tooth marks.[ Of these tooth marks, the sizes and shapes match well with the teeth of ]fish
A fish (: fish or fishes) is an aquatic animal, aquatic, Anamniotes, anamniotic, gill-bearing vertebrate animal with swimming fish fin, fins and craniate, a hard skull, but lacking limb (anatomy), limbs with digit (anatomy), digits. Fish can ...
, crocodyliforms or other scavengers, but no confirmed theropod marks. The high number of individuals present suggests that a herd of ''Europasaurus'' was crossing a tidal zone and drowned.[ While the dominant large-bodied animal present is ''Europasaurus'', there is also material from a diplodocid sauropod, a ]stegosauria
Stegosauria is a group of Herbivore, herbivorous ornithischian dinosaurs that lived during the Jurassic and early Cretaceous Period (geology), periods. Stegosaurian fossils have been found mostly in the Northern Hemisphere (North America, Europe a ...
n, and multiple theropods. Three cervicals of the diplodocid are preserved, and from their size it is possible that the taxon was also a dwarf. The stegosaurian and variety of theropods only preserve teeth, with the exception of a few bones possibly from a taxon in Ceratosauridae
Ceratosaurs are members of the clade Ceratosauria, a group of dinosaurs defined as all theropods sharing a more recent common ancestor with ''Ceratosaurus'' than with birds. The oldest known ceratosaur, ''Saltriovenator'', dates to the earliest ...
.[ Isolated teeth show that there were at least four different types of ]theropods
Theropoda (; from ancient Greek , (''therion'') "wild beast"; , (''pous, podos'') "foot"">wiktionary:ποδός"> (''pous, podos'') "foot" is one of the three major groups (clades) of dinosaurs, alongside Ornithischia and Sauropodom ...
present at the locality, including the megalosaurid '' Torvosaurus sp.'' as well as an additional megalosaurid and indeterminate members of the Allosauridae
Allosauridae is an extinct family of medium to large bipedal, carnivorous allosauroid theropod dinosaurs from the Late Jurassic. Allosauridae is a fairly old taxonomic group, having been first named by the American paleontologist Othniel Cha ...
and Ceratosauria
Ceratosaurs are members of the clade Ceratosauria, a group of dinosaurs defined as all theropods sharing a more recent common ancestor with '' Ceratosaurus'' than with birds. The oldest known ceratosaur, '' Saltriovenator'', dates to the earlies ...
; and there are also the oldest teeth known from Velociraptorinae.
Besides the dinosaurs, many small-bodied terrestrial vertebrates are also preserved in the Langenberg quarry. Such animals include a well-preserved three-dimensional pterosaur
Pterosaurs are 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 million to 66 million years ago). Pterosaurs are the earli ...
skeleton from Dsungaripteridae, and isolated remains from Ornithocheiroidea and Ctenochasmatidae
Ctenochasmatidae is a group of pterosaurs within the suborder Pterodactyloidea. They are characterized by their distinctive teeth, which are thought to have been used for filter-feeding. Ctenochasmatids lived from the Late Jurassic to the Earl ...
; a paramacellodid lizard
Lizard is the common name used for all Squamata, squamate reptiles other than snakes (and to a lesser extent amphisbaenians), encompassing over 7,000 species, ranging across all continents except Antarctica, as well as most Island#Oceanic isla ...
; and partial skeletons and skulls from a relative of '' Theriosuchus'' now named as the genus '' Knoetschkesuchus''. Teeth from dryolestid mammals are also preserved, as well as a docodont
Docodonta is an Order (biology), order of extinct Mesozoic Mammaliaformes, mammaliaforms (advanced cynodonts closely related to true Crown group, crown-group mammals). They were among the most common mammaliaforms of their time, persisting from t ...
, a taxon in Eobaataridae
Eobaataridae is a family of fossil mammals within the order Multituberculata. Remains are known from the Lower Cretaceous of Europe and Asia. They are among the most derived representatives of the informal suborder "Plagiaulacida", and closely re ...
, and a multituberculate
Multituberculata (commonly known as multituberculates, named for the multiple tubercles of their teeth) is an extinct order of rodent-like mammals with a fossil record spanning over 130 million years. They first appeared in the Middle Jurassic, a ...
with similarities to '' Proalbionbataar'' (now named '' Teutonodon'').
Extinction
Dinosaur footprints preserved at the Langenberg Quarry display a possible reason for the extinction of ''Europasaurus'', and potentially other insular dwarfs present on the islands of the region. The footprints are located above the deposit of ''Europasaurus'' individuals, which shows that at least 35,000 years after that deposit there was a drop in sea level which allowed for a faunal overturn. The inhabiting theropods of the island, that coexisted with ''Europasaurus'', would have been about , but the theropods that arrived over the land bridge preserve footprints
up to , which indicates a body size between if reconstructed as an allosaurian. It was suggested by the describers of these tracks (Jens Lallensack and colleagues), that these theropod taxa likely made the specialized dwarf fauna extinct, and the bed from which the footprints originated (Langenberg bed 92) is probably the youngest in which ''Europasaurus'' is present.
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{{Taxonbar, from=Q131712
Brachiosauridae
Dinosaur genera
Kimmeridgian dinosaurs
Fossil taxa described in 2006
Taxa named by Octávio Mateus
Dinosaurs of Germany