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Dimetrodon
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It has been suggested that this article be merged with List of Dimetrodon species and Dimetrodon borealis. (Discuss) Proposed since May 2017.
Dimetrodon
Temporal range: Cisuralian, 295–272 Ma
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Skeleton of D. incisivum, Staatliches Museum für Naturkunde Karlsruhe
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Clade: Tetrapodomorpha
Family: †Sphenacodontidae
Subfamily: †Sphenacodontinae
Genus: †Dimetrodon
Cope, 1878
Type species
†Dimetrodon limbatus
Cope, 1877
(originally Clepsydrops limbatus)
Species
See below
Synonyms
Genus synonymy[show]
Embolophorus
Cope, 1878
Theropleura
Cope, 1880
Bathyglyptus
Case, 1911
Eosyodon
Olson, 1962
Species synonymy[show]
Clepsydrops limbatus
Cope, 1877
Clepsydrops gigas
Cope, 1878
Clepsydrops natalis
Cope, 1878
Dimetrodon gigas
Cope, 1878
Dimetrodon incisivus
Cope, 1878
Dimetrodon rectiformis
Cope, 1878
Embolophorus dollovianus
Cope, 1878
Dimetrodon semiradicatus
Cope, 1881
Clepsydrops macrospondylus
Cope 1884
Dimetrodon platycentrus
Case, 1907
Theropleura grandis
Case, 1907
Bathyglyptus theodori
Case, 1911
Dimetrodon maximus
Romer, 1936
Eosyodon hudsoni
Olson, 1962
Dimetrodon (/daɪˈmiːtrədɒn/ ( listen)[1] or /daɪˈmɛtrədɒn/,[2] meaning "two measures of teeth") is an extinct genus of synapsids that lived during the Cisuralian (Early Permian), around 295–272 million years ago (Ma).[3][4][5] It is a member of the family Sphenacodontidae. The most prominent feature of Dimetrodon is the large neural spine sail on its back formed by elongated spines extending from the vertebrae. It walked on four legs and had a tall, curved skull with large teeth of different sizes set along the jaws. Most fossils have been found in the southwestern United States, the majority coming from a geological deposit called the Red Beds of Texas and Oklahoma. More recently, fossils have been found in Germany. Over a dozen species have been named since the genus was first described in 1878.
Dimetrodon is often mistaken for a dinosaur or as a contemporary of dinosaurs in popular culture, but it became extinct some 40 million years before the first appearance of dinosaurs. Reptile-like in appearance and physiology, Dimetrodon is nevertheless more closely related to mammals than to modern reptiles, though it is not a direct ancestor of mammals.[4] Dimetrodon is assigned to the "non-mammalian synapsids", a group traditionally called "mammal-like reptiles".[4] This groups Dimetrodon together with mammals in a clade (evolutionary group) called Synapsida, while placing dinosaurs, reptiles and birds in a separate clade, Sauropsida. Single openings in the skull behind each eye, known as temporal fenestrae, and other skull features distinguish Dimetrodon and mammals from most of the earliest sauropsids.
Dimetrodon was probably one of the apex predators of the Cisuralian ecosystems, feeding on fish and tetrapods, including reptiles and amphibians. Smaller Dimetrodon species may have had different ecological roles. The sail of Dimetrodon may have been used to stabilize its spine or to heat and cool its body as a form of thermoregulation. Some recent studies argue that the sail would have been ineffective at removing heat from the body, and was most likely used in courtship display.
Contents
1 Description
1.1 Skull
1.1.1 Teeth
1.1.2 Nasal cavity
1.1.3 Jaw joint and ear
1.2 Tail
1.3 Sail
1.4 Skin
2 Classification history
2.1 First descriptions by Cope
2.2 Early 20th century descriptions
2.3 New specimens
2.4 Species
3 Phylogenetic classification
3.1 Phylogenetic taxonomy of Synapsida
4 Paleobiology
4.1 Function of neural spines
4.1.1 Thermoregulation
4.1.2 Sexual selection
4.2 Sexual dimorphism
5 Paleoecology
5.1 Food web
6 References
7 External links
Description[edit]
Restoration of D. giganhomogenes with exposed neural spine tips
Dimetrodon was a quadrupedal, sail-backed synapsid. Most Dimetrodon species ranged in length from 1.7 to 4.6 metres (6 to 15 ft) and are estimated to have weighed between 28 and 250 kilograms (60 and 550 lb).[6] The largest known species of Dimetrodon is D. angelensis at 4.6 metres (15 ft) and the smallest is D. teutonis at 60 centimetres (24 in).[7][8] The larger species of Dimetrodon were among the largest predators of the Early Permian, although the closely related Tappenosaurus, known from skeletal fragments in slightly younger rocks, may have been even larger at an estimated 18 feet (5.5 m) in total body length.[9][10] Although some Dimetrodon species could grow very large, many juvenile specimens are known.[11]
Skull[edit]
A single large opening on either side of the back of the skull links Dimetrodon with mammals and distinguishes it from most of the earliest sauropsids, which either lack openings or have two openings. Features such as ridges on the inside of the nasal cavity and a ridge at the back of the lower jaw are thought to be part of an evolutionary progression from early tetrapods (four-limbed vertebrates) to mammals.
The skull of Dimetrodon is tall and compressed laterally, or side-to-side. The eye sockets are positioned high and far back in the skull. Behind each eye socket is a single hole called an infratemporal fenestra. An additional hole in the skull, the supratemporal fenestra, can be seen when viewed from above. The back of the skull (the occiput) is oriented at a slight upward angle, a feature that it shares with all other early synapsids.[12] The upper margin of the skull slopes downward in a convex arc to the tip of the snout. The tip of the upper jaw, formed by the premaxilla bone, is raised above the part of the jaw formed by the maxilla bone to form a maxillary "step." Within this step is a diastema, or gap in the tooth row. Its skull was more heavily built than a dinosaur's.
Bones of the skull of Dimetrodon
Teeth[edit]
A skull of D. grandis
The size of the teeth varies greatly along the length of the jaws, lending Dimetrodon its name, which means "two measures of tooth" in reference to sets of small and large teeth.[13] One or two pairs of caniniforms (large pointed canine-like teeth) extend from the maxilla. Large incisor teeth are also present at the tips of the upper and lower jaws, rooted in the premaxillae and dentary bones. Small teeth are present around the maxillary "step" and behind the caniniforms, becoming smaller further back in the jaw.[14]
Many teeth are widest at their midsections and narrow closer to the jaws, giving them the appearance of a teardrop. Teardrop-shaped teeth are unique to Dimetrodon and other closely related sphenacodontids, and help distinguish them from other early synapsids.[8] As in many other early synapsids, the teeth of most Dimetrodon species are serrated at their edges.[8] The serrations of Dimetrodon teeth were so fine that they resembled tiny cracks.[15] The dinosaur Albertosaurus had similarly crack-like serrations, but, at the base of each serration was a round void, which would have functioned to distribute force over a larger surface area and prevent the stresses of feeding from causing the crack to spread through the tooth. Unlike Albertosaurus, Dimetrodon teeth lacked adaptations that would stop cracks from forming at their serrations.[15] The teeth of D. teutonis lack serrations, but still have sharp edges.[8]
A study in 2014 shows that Dimetrodon was in an arms race against its prey. The smaller species, D.milleri, had no serrations since it ate small prey. As prey grew larger, severalDimetrodon species started developing serrations on their teeth and increasing in size. For instance, D.limbatus had enamel serrations that helped it cut through flesh (which were similar to the serrations that can be found on Secodontosaurus). The second largest dimetrodon species, D.grandis, has denticle serrations similar to those of sharks and theropod dinosaurs, making its teeth even more specialized for slicing through flesh. This study demonstrates that, as Dimetrodon's prey grew larger, it responded by growing larger itself and developing ever-sharper teeth.[16]
Nasal cavity[edit]
On the inner surface of the nasal section of skull are ridges called nasoturbinals, which may have supported cartilage that increased the area of the olfactory epithelium, the layer of tissue that detects odors. These ridges are much smaller than those of later synapsids from the Late Permian and Triassic, whose large nasoturbinals are taken as evidence for warm-bloodedness because they may have supported mucous membranes that warmed and moistened incoming air. Thus, the nasal cavity of Dimetrodon is transitional between those of early land vertebrates and mammals.[17]
Jaw joint and ear[edit]
Another transitional feature of Dimetrodon is a ridge in the back of the jaw called the reflected lamina. The reflected lamina is found on the articular bone, which connects to the quadrate bone of the skull to form the jaw joint. In later mammal ancestors, the articular and quadrate separated from the jaw joint while the articular developed into the malleus bone of the middle ear. The reflected lamina became part of a ring called the tympanic annulus that supports the ear drum in all living mammals.[18]
Tail[edit]
An outdated restoration of Dimetrodon from 1908 showing a short tail, made before the discovery of skeletons with complete tails.
The tail of Dimetrodon makes up a large portion of its total body length and includes around 50 caudal vertebrae. Tails were missing or incomplete in the first described skeletons of Dimetrodon; the only caudal vertebrae known were the eleven closest to the hip. Since these first few caudal vertebrae narrow rapidly as they progress farther from the hip, many paleontologists in the late nineteenth and early twentieth centuries thought that Dimetrodon had a very short tail. It was not until 1927 that a largely complete tail of Dimetrodon was described.[19]
Sail[edit]
Two D. grandis skeletons, Royal Tyrrell Museum
The sail of Dimetrodon is formed by elongated neural spines projecting from the vertebrae. Each spine varies in cross-sectional shape from its base to its tip in what is known as "dimetrodont" differentiation.[20] Near the vertebra body, the spine cross section is laterally compressed into a rectangular shape, and closer to the tip, it takes on a figure-eight shape as a groove runs along either side of the spine. The figure-eight shape is thought to reinforce the spine, preventing bending and fractures.[21] A cross section of the spine of one specimen of Dimetrodon giganhomogenes is rectangular in shape but preserves figure-eight shaped rings close to its center, indicating that the shape of spines may change as individuals age.[22] The microscopic anatomy of each spine varies from base to tip, indicating where it was embedded in the muscles of the back and where it was exposed as part of a sail. The lower or proximal portion of the spine has a rough surface that would have served as an anchoring point for the epaxial muscles of the back, and also has a network of connective tissues called Sharpey's fibers that indicate it was embedded within the body. Higher up on the distal (outer) portion of the spine, the bone surface is smoother. The periosteum, a layer of tissue surrounding the bone, is covered in small grooves that presumably supported the blood vessels that vascularized the sail.[23]
The large groove that runs the length of the spine was once thought to be a channel for blood vessels, but since the bone does not contain vascular canals, the sail is not thought to have been as highly vascularized as once thought. Some specimens of Dimetrodon preserve deformed areas of the neural spines that appear to be healed-over fractures. The cortical bone that grew over these breaks is highly vascularized, suggesting that soft tissue must have been present on the sail to supply the site with blood vessels.[21] Layered lamellar bone makes up most of the neural spine's cross-sectional area, and contains lines of arrested growth that can be used to determine the age of each individual at death.[24] In many specimens of D. gigashomogenes the distal portions of spines bend sharply, indicating that the sail would have had an irregular profile in life. Their crookedness suggests that soft tissue may not have extended all the way to the tips of the spines, meaning that the sail's webbing may not have been as extensive as it is commonly imagined.[20]
Skin[edit]
No fossil evidence of Dimetrodon's skin has yet been found. Impressions of the skin of a related animal, Estemmenosuchus, indicate that it would have been smooth and well-provided with glands.[25] Dimetrodon also may have had large scutes on the underside of its tail and belly, as other synapsids did.[26][27] Recent evidence from the varanopid Ascendonanus suggests that some early synapsids had squamate-like scales.[28]
Classification history[edit]
First descriptions by Cope[edit]
Fossils now attributed to Dimetrodon were first studied by American paleontologist Edward Drinker Cope in the 1870s. Cope had obtained the fossils along with those of many other Permian tetrapods (four-limbed vertebrates) from several collectors who had been exploring a group of rocks in Texas called the Red Beds. Among these collectors were Swiss naturalist Jacob Boll, Texas geologist W. F. Cummins, and amateur paleontologist Charles Hazelius Sternberg.[29] Most of Cope's specimens went to the American Museum of Natural History or to the University of Chicago's Walker Museum (most of the Walker fossil collection is now housed in the Field Museum of Natural History).
Sternberg sent some of his own specimens to German paleontologist Ferdinand Broili at Munich University, although Broili was not as prolific as Cope when it came to describing specimens. Cope's rival Othniel Charles Marsh also collected some bones of Dimetrodon, which he sent to the Walker Museum.[30] The first use of the name Dimetrodon came in 1878 when Cope named the species Dimetrodon incisivus, Dimetrodon rectiformis, and Dimetrodon gigas in the scientific journal Proceedings of the American Philosophical Society.[31]
However, the first description of a Dimetrodon fossil came a year earlier, when Cope named the species Clepsydrops limbatus from the Texas Red Beds.[32] (The name Clepsydrops was first coined by Cope in 1875 for sphenacodontid remains from Vermilion County, Illinois, and was later employed for many sphenacontid specimens from Texas; many new species of sphenacodontids from Texas were assigned to either Clepsydrops or Dimetrodon in the late nineteenth and early twentieth centuries.) C. limbatus was reclassified as a species of Dimetrodon in 1940, meaning that Cope's 1877 paper was the first record of Dimetrodon.
Cope was the first to describe a sail-backed synapsid with the naming of Clepsydrops natalis in his 1878 paper, although he called the sail a fin and compared it to the crests of the modern basilisk lizard (Basilicus). Sails were not preserved in the specimens of D. incisivus and D. gigas that Cope described in his 1878 paper, but elongated spines were present in the D. rectiformis specimen he described.[31] Cope commented on the purpose of the sail in 1886, writing, "The utility is difficult to imagine. Unless the animal had aquatic habits, and swam on its back, the crest or fin must have been in the way of active movements... The limbs are not long enough nor the claws acute enough to demonstrate arboreal habits, as in the existing genus Basilicus, where a similar crest exists."[14]
Early 20th century descriptions[edit]
Restoration of Dimetrodon and Edaphosaurus (background), Charles R. Knight, 1897
In the first few decades of the twentieth century, American paleontologist E. C. Case authored many studies on Dimetrodon and described several new species. He received funding from the Carnegie Institution for his study of many Dimetrodon specimens in the collections of the American Museum of Natural History and several other museums.[30] Many of these fossils had been collected by Cope but had not been thoroughly described, as Cope was known for erecting new species on the basis of only a few bone fragments.
Beginning in the late 1920s, paleontologist Alfred Romer restudied many Dimetrodon specimens and named several new species. In 1940, Romer coauthored a large study with Llewellyn Ivor Price called "Review of the Pelycosauria" in which the species of Dimetrodon named by Cope and Case were reassessed.[33] Most of the species names considered valid by Romer and Price are still used today.[23]
New specimens[edit]
In the decades following Romer and Price's monograph, many Dimetrodon specimens were described from localities outside Texas and Oklahoma. The first was described from the Four Corners region of Utah in 1966[34] and another was described from Arizona in 1969.[35] In 1975, Olson reported Dimetrodon material from Ohio.[36] A new species of Dimetrodon called D. occidentalis (meaning "western Dimetrodon") was named in 1977 from New Mexico.[37] The specimens found in Utah and Arizona probably also belong to D. occidentalis.[38]
Before these discoveries, a theory existed that a Midcontinental seaway separated what is now Texas and Oklahoma from more western lands during the Early Permian, isolating Dimetrodon to a small region of North America while a smaller sphenacodontid called Sphenacodon dominated the western area. While this seaway probably did exist, the discovery of fossils outside Texas and Oklahoma show that its extent was limited and that it was not an effective barrier to the distribution of Dimetrodon.[37][39]
In 2001, a new species of Dimetrodon called D. teutonis was described from the Lower Permian Bromacker locality at the Thuringian Forest of Germany, extending the geographic range of Dimetrodon outside North America for the first time.[6]
Species[edit]
Main article: List of Dimetrodon species
Species
Authority
Location
Status
Synonyms
Images
Dimetrodon angelensis Olson, 1962 Texas Valid
Dimetrodon borealis Leidy, 1854 Prince Edward Island Valid Previously known as Bathygnathus borealis
Dimetrodon booneorum Romer, 1937 Texas Valid
Dimetrodon dollovianus Case, 1907 Texas Valid Embolophorus dollovianus Cope, 1888
Dimetrodon gigahomogenes Case, 1907 Texas Valid
Dimetrodon grandis Romer and Price, 1940 Oklahoma
Texas Valid Clepsydrops gigas Cope, 1878
Dimetrodon gigas Cope, 1878
Theropleura grandis Case, 1907
Bathyglyptus theodori Case, 1911
Dimetrodon maximus Romer 1936
Dimetrodon kempae Romer, 1937 Texas Possible nomen dubium
Dimetrodon limbatus Romer and Price, 1940 Oklahoma
Texas Valid Clepsydrops limbatus Cope, 1877
Dimetrodon incisivus Cope, 1878
Dimetrodon rectiformis Cope, 1878
Dimetrodon semiradicatus Cope, 1881
Dimetrodon loomisi Romer, 1937 Texas
Oklahoma Valid
Dimetrodon macrospondylus Case, 1907 Texas Valid Clepsydrops macrospondylus Cope, 1884
Dimetrodon platycentrus Case, 1907
Dimetrodon milleri Romer, 1937 Texas Valid
Dimetrodon natalis Romer, 1936 Texas Valid Clepsydrops natalis Cope, 1878
Dimetrodon occidentalis Berman, 1977 Arizona
New Mexico
Utah Valid
Dimetrodon teutonis Berman et al., 2001 Germany Valid
Phylogenetic classification[edit]
See also: Synapsid § Linnaean and cladistic classifications
Dimetrodon is an early member of a group called synapsids, which include mammals and many of their extinct relatives, though it is not an ancestor of any mammal (which appeared millions of years later[40]). It is often mistaken for a dinosaur in popular culture, despite having become extinct some 40 million years (Ma) before the first appearance of dinosaurs in the Triassic period. As a synapsid, Dimetrodon is more closely related to mammals than to dinosaurs or any living reptile. By the early 1900s most paleontologists called Dimetrodon a reptile in accordance with Linnean taxonomy, which ranked Reptilia as a class and Dimetrodon as a genus within that class. Mammals were assigned to a separate class, and Dimetrodon was described as a "mammal-like reptile". Paleontologists theorized that mammals evolved from this group in (what they called) a reptile-to-mammal transition.
Phylogenetic taxonomy of Synapsida[edit]
D. grandis skeleton, North American Museum of Ancient Life
Under phylogenetic systematics, the descendants of the last common ancestor of Dimetrodon and all living reptiles would include all mammals because Dimetrodon is more closely related to mammals than to any living reptile. Thus, if it is desired to avoid the clade that contains both mammals and the living reptiles, then Dimetrodon must not be included in that clade—nor any other "mammal-like reptile". Descendants of the last common ancestor of mammals and reptiles (which appeared around 310 Ma in the Late Carboniferous) are therefore split into two clades: Synapsida, which includes Dimetrodon and mammals, and Sauropsida, which includes living reptiles and all extinct reptiles more closely related to them than to mammals.[4]
Within clade Synapsida, Dimetrodon is part of the clade Sphenacodontia, which was first proposed as an early synapsid group in 1940 by paleontologists Alfred Romer and Llewellyn Ivor Price, along with the groups Ophiacodontia and Edaphosauria.[33] All three groups are known from the Late Carboniferous and Early Permian. Romer and Price distinguished them primarily by postcranial features such as the shapes of limbs and vertebrae. Ophiacodontia was considered the most primitive group because its members appeared the most reptilian, and Sphenacodontia was the most advanced because its members appeared the most like a group called Therapsida, which included the closest relatives to mammals. Romer and Price placed another group of early synapsids called varanopids within Sphenacodontia, considering them to be more primitive than other sphenacodonts like Dimetrodon.[41] They thought varanopids and Dimetrodon-like sphenacodonts were closely related because both groups were carnivorous, although varanopids are much smaller and more lizard-like, lacking sails.
The modern view of synapsid relationships was proposed by paleontologist Robert R. Reisz in 1986, whose study included features mostly found in the skull rather than in the postcranial skeleton.[42] Dimetrodon is still considered a sphenacodont under this phylogeny, but varanodontids are now considered more basal synapsids, falling outside clade Sphenacodontia. Within Sphenacodontia is the group Sphenacodontoidea, which in turn contains Sphenacodontidae and Therapsida. Sphenacodontidae is the group containing Dimetrodon and several other sail-backed synapsids like Sphenacodon and Secodontosaurus, while Therapsida includes mammals and their mostly Permian and Triassic relatives.
Below is the cladogram Clade Synapsida, which follows this phylogeny of Synapsida as modified from the analysis of Benson (2012).[41]
Restoration of various Dimetrodon species to scale
Amniota
Sauropsida (including dinosaurs, living reptiles and birds)
Synapsida
†Ophiacodontidae
†Varanopidae
†Caseasauria
†Ianthodon schultzei
†Edaphosauridae
Sphenacodontia
†Haptodus garnettensis
†Pantelosaurus saxonicus
†Sphenacodontidae
†Cutleria wilmarthi
†Secodontosaurus obtusidens
†Cryptovenator hirschbergeri
†Dimetrodon spp.
†Sphenacodon spp.
Therapsida (including mammals)
The below cladogram shows the relationships of a few Dimetrodon species, from Brink et al., (2015).[43]
Sphenacodontidae
Secodontosaurus
Sphenacodon
Ctenospondylus
Dimetrodon milleri
Dimetrodon limbatus
Dimetrodon borealis
Dimetrodon grandis
Paleobiology[edit]
Function of neural spines[edit]
D. grandis warming up at sunrise
Paleontologists have proposed many ways in which the sail could have functioned in life. Some of the first to think about its purpose suggested that the sail may have served as camouflage among reeds while Dimetrodon waited for prey, or as an actual boat-like sail to catch the wind while the animal was in the water.[44] Another is that the long neural spines could have stabilized the trunk by restricting up-and-down movement, which would allow for a more efficient side-to-side movement while walking.[21]
Thermoregulation[edit]
In 1940, Alfred Romer and Llewellyn Ivor Price proposed that the sail served a thermoregulatory function, allowing individuals to warm their bodies with the sun's heat. In the following years, many models were created to estimate the effectiveness of thermoregulation in Dimetrodon. For example, in a 1973 article in the journal Nature, paleontologists C. D. Bramwell and P. B. Fellgett estimated that it took a 200 kilograms (440 lb) individual about one and a half hours for its body temperature to rise from 26 to 32 °C (79 to 90 °F).[45] In 1986, Steven C. Haack concluded that the warming was slower than previously thought and that the process probably took four hours. Using a model based on a variety of environmental factors and hypothesized physiological aspects of Dimetrodon, Haack found that the sail allowed Dimetrodon to warm faster in the morning and reach a slightly higher body temperature during the day, but that it was ineffective in releasing excess heat and did not allow Dimetrodon to retain a higher body temperature at night.[46] In 1999, a group of mechanical engineers created a computer model to analyze the ability of the sail to regulate body temperature during different seasons, and concluded that the sail was beneficial for capturing and releasing heat at all times in the year.[47]
The comparatively small D. milleri
Most of these studies give two thermoregulatory roles for the sail of Dimetrodon: one as a means of warming quickly in the morning, and another as a way to cool down when body temperature becomes high. Dimetrodon and all other Early Permian land vertebrates are assumed to have been cold-blooded or poikilothermic, relying on the sun to maintain a high body temperature. Because of its large size, Dimetrodon had high thermal inertia, meaning that changes in body temperature occurred more slowly in it than in smaller-bodied animals. As temperatures rose in the mornings, the small-bodied prey of Dimetrodon could warm their bodies much faster than could something the size of Dimetrodon. Many paleontologists including Haack have proposed that the sail of Dimetrodon may have allowed it to warm quickly in the morning in order to keep pace with its prey.[46] The sail's large surface area also meant heat could dissipate quickly into the surroundings, useful if the animal needed to release excess heat produced by metabolism or absorbed from the sun. Dimetrodon may have angled its sail away from the sun to cool off or restricted blood flow to the sail to maintain heat at night.[44]
In 1986, J. Scott Turner and C. Richard Tracy proposed that the evolution of a sail in Dimetrodon was related to the evolution of warm-bloodedness in mammal ancestors. They thought that the sail of Dimetrodon enabled it to be homeothermic, maintaining a constant, albeit low, body temperature. Mammals are also homeothermic, although they differ from Dimetrodon in being endothermic, controlling their body temperature internally through heightened metabolism. Turner and Tracy noted that early therapsids, a more advanced group of synapsids closely related to mammals, had long limbs which can release heat in a manner similar to that of the sail of Dimetrodon. The homeothermy that developed in animals like Dimetrodon may have carried over to therapsids through a modification of body shape, which would eventually develop into the warm-bloodedness of mammals.[48]
Sphenacodon had a low crest along its back
Recent studies on the sail of Dimetrodon and other sphenacodontids support Haack's 1986 contention that the sail was poorly adapted to releasing heat and maintaining a stable body temperature. The presence of sails in small-bodied species of Dimetrodon such as D. milleri and D. teutonis does not fit the idea that the sail's purpose was thermoregulation because smaller sails are less able to transfer heat and because small bodies can absorb and release heat easily on their own. Moreover, close relatives of Dimetrodon such as Sphenacodon have very low crests that would have been useless as thermoregulatory devices.[23] The large sail of Dimetrodon is thought to have developed gradually from these smaller crests, meaning that over most of the sail's evolutionary history, thermoregulation could not have served an important function.[49]
Larger bodied specimens of Dimetrodon have larger sails relative to their size, an example of positive allometry. Positive allometry may benefit thermoregulation because it means that, as individuals get larger, surface area increases faster than mass. Larger-bodied animals generate a great deal of heat through metabolism, and the amount of heat that must be dissipated from the body surface is significantly greater than what must be dissipated by smaller-bodied animals. Effective heat dissipation can be predicted across many different animals with a single relationship between mass and surface area. However, a 2010 study of allometry in Dimetrodon found a different relationship between its sail and body mass: the actual scaling exponent of the sail was much larger than the exponent expected in an animal adapted to heat dissipation. The researchers concluded that the sail of Dimetrodon grew at a much faster rate than was necessary for thermoregulation, and suggested that sexual selection was the primary reason for its evolution.[49]
Sexual selection[edit]
The allometric exponent for sail height is similar in magnitude to the scaling of interspecific antler length to shoulder height in cervids. Furthermore, as Bakker (1970) observed in the context of Dimetrodon, many lizard species raise a dorsal ridge of skin during threat and courtship displays, and positively allometric, sexually dimorphic frills and dewlaps are present in extant lizards (Echelle et al. 1978; Christian et al. 1995). There is also evidence of sexual dimorphism both in the robustness of the skeleton and in the relative height of the spines of D. limbatus (Romer and Price 1940).[49]
Sexual dimorphism[edit]
Dimetrodon may have been sexually dimorphic, meaning that males and females had slightly different body sizes. Some specimens of Dimetrodon have been hypothesized as males because they have thicker bones, larger sails, longer skulls, and more pronounced maxillary "steps" than others. Based on these differences, the mounted skeletons in the American Museum of Natural History (AMNH 4636) and the Field Museum of Natural History may be males and the skeletons in the Denver Museum of Nature and Science (MCZ 1347) and the University of Michigan Museum of Natural History may be females.[33]
The mounted D. limbatus skeleton AMNH 4636 may represent the male type.
The D. incisivus skeleton in the University of Michigan Museum of Natural History may represent the female type.
Paleoecology[edit]
Possible Dimetrodon footprint, Prehistoric Trackways National Monument
Fossils of Dimetrodon are known from the United States (Texas, Oklahoma, New Mexico, Arizona, Utah and Ohio) and Germany, areas that were part of the supercontinent Euramerica during the Early Permian. Within the United States, almost all material attributed to Dimetrodon has come from three geological groups in north-central Texas and south-central Oklahoma: the Clear Fork Group, the Wichita Group, and the Pease River Group.[50][51] Most fossil finds are part of lowland ecosystems which, during the Permian, would have been vast wetlands. In particular, the Red Beds of Texas is an area of great diversity of fossil tetrapods, or four-limbed vertebrates. In addition to Dimetrodon, the most common tetrapods in the Red Beds and throughout Early Permian deposits in the southwestern United States, are the amphibians Archeria, Diplocaulus, Eryops, and Trimerorhachis, the reptiliomorph Seymouria, the reptile Captorhinus, and the synapsids Ophiacodon and Edaphosaurus. These tetrapods made up a group of animals that paleontologist Everett C. Olson called the "Permo-Carboniferous chronofauna," a fauna that dominated the continental Euramerican ecosystem for several million years.[52] Based on the geology of deposits like the Red Beds, the fauna is thought to have inhabited a well-vegetated lowland deltaic ecosystem.[53]
Food web[edit]
Restoration of D. grandis and the temnospondyl Eryops, both found in the Red Beds of Texas
Olson made many inferences on the paleoecology of the Texas Red beds and the role of Dimetrodon within its ecosystem. He proposed several main types of ecosystems in which the earliest tetrapods lived. Dimetrodon belonged to the most primitive ecosystem, which developed from aquatic food webs. In it, aquatic plants were the primary producers and were largely fed upon by fish and aquatic invertebrates. Most land vertebrates fed on these aquatic primary consumers. Dimetrodon was probably the top predator of the Red Beds ecosystem, feeding on a variety of organisms such as the shark Xenacanthus[citation needed], the aquatic amphibians Trimerorhachis and Diplocaulus, and the terrestrial tetrapods Seymouria and Trematops. Insects are known from the Early Permian Red Beds and were probably involved to some degree in the same food web as Dimetrodon, feeding small reptiles like Captorhinus. The Red Beds assemblage also included some of the first large land-living herbivores like Edaphosaurus and Diadectes. Feeding primarily on terrestrial plants, these herbivores did not derive their energy from aquatic food webs. According to Olson, the best modern analogue for the ecosystem Dimetrodon inhabited is the Everglades.[53] The exact lifestyle of Dimetrodon (amphibious to terrestrial) has long been controversial, but bone microanatomy supports a terrestrial lifestyle,[54] which implies that it would have fed mostly on land, on the banks, or in very shallow water. Evidence also exists for Dimetrodon preying on aestivating Diplocaulus during times of drought, with three partially eaten juvenile Diplocaulus in a burrow of eight bearing teeth marks from a Dimetrodon that unearthed and killed them.[55]
The only species of Dimetrodon found outside the southwestern United States is D. teutonis from Germany. Its remains were found in the Tambach Formation in a fossil site called the Bromacker locality. The Bromacker's assemblage of Early Permian tetrapods is unusual in that there are few large-bodied synapsids serving the role of top predators. D. teutonis is estimated to have been only 1.7 metres (5.6 ft) in length, too small to prey on the large diadectid herbivores that are abundant in the Bromacker assemblage. It more likely ate small vertebrates and insects. Only three fossils can be attributed to large predators, and they are thought to have been either large varanopids or small sphenacodonts, both of which could potentially prey on D. teutonis. In contrast to the lowland deltaic Red Beds of Texas, the Bromacker deposits are thought to have represented an upland environment with no aquatic species. It is possible that large-bodied carnivores were not part of the Bromacker assemblage because they were dependent on large aquatic amphibians for food.[6]
References[edit]
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^ Jump up to: a b Case, E.C. (1907). Revision of the Pelycosauria of North America. Washington, D.C.: Carnegie Institution of Washington. pp. 1–176.
^ Jump up to: a b Cope, E.D. (1878). "Descriptions of extinct Batrachia and Reptilia from the Permian formation of Texas". Proceedings of the American Philosophical Society. 17 (101): 505–530. JSTOR 982652.
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Jump up ^ http://www.livescience.com/40842-finned-monster-chomped-heads-off-ancient-amphibians.html
External links[edit]
Wikimedia Commons has media related to Dimetrodon.
Wikispecies has information related to Dimetrodon
Paleontology portal
Dimetrodon Palaeos page on Dimetrodon
Introduction to the Pelycosaurs University of California Museum of Paleontology webpage on early synapsids, including Dimetrodon
"Dimetrodon: Our Most Unlikely Ancestor". PBS Eons. August 21, 2017 – via YouTube.
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Wikidata: Q130869
Wikispecies: Dimetrodon
EoL: 4530661
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Music in Buffy the Vampire Slayer and Angel
From Wikipedia, the free encyclopedia
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This article's tone or style may not reflect the encyclopedic tone used on Wikipedia. See Wikipedia's guide to writing better articles for suggestions. (December 2007) (Learn how and when to remove this template message)
The use of music was a key component in the fictional Buffyverse established by the TV shows Buffy the Vampire Slayer and Angel. Many actors of both series are gifted and professional singers/musicians (Anthony Head, James Marsters, Amber Benson, Andy Hallett and Christian Kane to just cite a few).
Contents
[hide]
1 Themes
1.1 Buffy theme
1.2 Angel theme
2 Buffyverse discography
3 Score
3.1 Walter Murphy
3.2 Christophe Beck
3.3 Thomas Wanker
3.4 Robert J. Kral
3.5 Robert Duncan
3.6 Douglas Romayne
4 The Bronze
5 Dingoes Ate My Baby
6 Lorne and Caritas
7 Shy
8 Soundtrack
8.1 Velvet Chain
9 Sarah McLachlan
10 Notes and references
11 External links
11.1 Themes
11.1.1 Darling Violetta
11.1.2 Nerf Herder
11.2 Composers
Themes[edit]
Buffy theme[edit]
The "Buffy theme" is the music played alongside the opening credits of the show. The theme itself has no lyrics; it begins with several notes played by an organ, a signifier for horror in movie culture from the 1930s onwards. This is quickly replaced by upbeat rock music; an electric guitar riff signifies youth culture and a post-modern twist on the horror genre.
The theme was played by the pop punk band Nerf Herder. In an interview they explain how they came to produce the theme:
"They [the Buffy production staff] had hired some famous theme song composer to come up with a theme and they didn’t like the one he came up with so they were kind of on a low budget, and asked local bands to come up with theme song ideas." [1]
In the DVD commentary for "Welcome to the Hellmouth," Whedon explained that part of his decision to go with Nerf Herder's theme was that Alyson Hannigan had made him listen to the band's music.
Nerf Herder later recorded a second version of the theme which was used for the opening titles from season 3 on.

Back in 1999, Nerf Herder lead singer Parry Grip gave the background on the writing of the melody. The original melody was not written specifically for the show; it had been in place before Whedon contacted the band asking them to submit a demo for a theme tune:
“ We did the theme song before the show was on the air. It was a song that had already been written, but we really didn't know where to take it. I had some ideas - it was originally going to have some kind of science fiction theme, which is weird, because we don't do songs like that. I remember having the title of 'Outer Space Rock' or something like that; that was a working thing.[2] ”
"Codo" has a science fiction theme and the lyrics reference an alien and flying objects. Their performances of the song featured outer space themes.
Angel theme[edit]
Darling Violetta is an alternative rock group that performed two songs, "Cure" and "Blue Sun" in the Buffy episode "Faith, Hope & Trick." They cowrote the song with songwriter Holly Knight (Love Is A Battlefield, Obsession. The Best).
The next year, Angel invited bands to submit demos for the theme music to the show. They asked bands to use "dark superhero ideas" and "Cello-rock."[3] Darling Violetta watched pivotal Angel-related episodes of Buffy like "Passion" and "Becoming, Part One" & "Becoming, Part Two" for inspiration along with cowriter and music producer, Holly Knight. Eventually Joss Whedon accepted Holly Knight/Darling Violetta's interpretation of an Angel theme as that most suitable for the show.
The theme has a slower tempo than the Buffy theme. It has heavier use of acoustic instruments such as cello and violin.
In 2005, Holly Knight and the band composed an extended version of the Angel theme called "The Sanctuary Extended Remix," which featured on the soundtrack of the series Live Fast - Die Never.
Joss Whedon and Alexis Denisof sing their own lyrics to the theme song in the DVD commentary to the episode "Spin the Bottle":
Buffyverse discography[edit]
A number of CD releases have been released linked to the Buffyverse:
Buffyverse discography
Release date
Buffy film soundtrack 1992
Buffy album 1999
Once More, with Feeling 2002
Radio Sunnydale 2003
Live Fast, Die Never 2005
Buffy score 2008
Score[edit]
Many people have worked on the scoring of Buffy and Angel. Some of the main composers were:
Walter Murphy[edit]
Main article: Walter Murphy
Murphy was the main composer on Buffy's first season.
Christophe Beck[edit]
Main article: Christophe Beck
Beck was the main composer on Buffy during seasons 2-4. He continued to compose key episodes after such as "The Gift" and "Once More, with Feeling." An album dedicated to his compositions for the show, entitled Buffy the Vampire Slayer: The Score, was released on September 9, 2008.[4]
Memorable tracks include:
"Close Your Eyes" plays during the award-winning "Becoming." This song incorporates the "Buffy/Angel Love Theme" that is used significantly throughout seasons 2 and 3. (This appears on Buffy album.)
"Remembering Jenny" plays after the death of Jenny Calendar ("Passion"). This song also features backing vocals from Anthony Stewart Head who played Rupert Giles on the show.
"Slayer's Elegy" plays during the final assault of "The Wish."
"Loneliness of Six," which plays at the end of "Lovers Walk" as various heartbroken characters walk off in different directions.
"Sacrifice" plays when Buffy sacrifices herself for her sister and for the world ("The Gift"). (This appears on Once More, With Feeling soundtrack along with Beck's suites from "Restless" and "Hush.")
"I'm Game" plays at the end of the penultimate episode of Angel ("Power Play") as the gang decides to go for a surely-suicidal final assault against the Circle of the Black Thorn. "I'm Game" was the theme used throughout the series whenever there was a need for "heroic music," and debuted in the first episode ("City Of"), when Angel decides to fight evil in Los Angeles.
Thomas Wanker[edit]
Main article: Thomas Wanker
The main composer for the majority of the 5th and 6th seasons of Buffy. His work tended to be subtle, and he thrived in "unassuming mysterious underscore."[5] Some of his best work appears in "Shadow" and "Into the Woods," more specifically, the tragic melodies which are used in musical montages shown along with Buffy's and Dawn's coping with their mother's illness, and Riley's feelings of alienation from Buffy.
Robert J. Kral[edit]
Main article: Robert J. Kral
Kral was the main composer for the series Angel, writing the music for 112 episodes for seasons 1 through 5. Initially hired by Christophe Beck for Season 1 with shared credit, Kral was employed by 20th Century Fox directly for Seasons 2 through 5. Kral worked for Beck on initial episodes, with Kral then writing most of the episodes for season one. Kral employed Zoran Boris and Douglas Romayne to write additional music for the final two seasons. The soundtrack Live Fast, Die Never mostly consists of his scores from throughout Angel. Fans were asked via internet and email for their favorite tracks and the resulting poll determined the bulk of the soundtrack released by Rounder, EMI and 20th Century Fox Television.
Memorable music scenes and themes by Kral for Angel include:
"Hero" for the character Doyle
"Darla's Theme" which recurs throughout the series
"Rebellion" for an episode with crossover plots and characters from Buffy
"Home" for Angel's son Connor, as he eventually lives out his life with no memory of his father
"Darla's Sacrifice" for an intense change in the story arc of the series
"Pylea," which features a score style change for several complete episodes in a different "dimension" of medieval style fantasy
"Vespa Chase" featuring a comedic motorcycle chase through Italy
"The End Of The World"
Robert Duncan[edit]
Main article: Robert Duncan (composer)
Duncan is credited as composer for Buffy episodes 7.02 and 7.04-7.22. Memorable tracks have included:
"Chosen," Buffy and Spike's love theme, Faith and Robin's theme, and the theme of Slayer Victory (during the fight in the Hellmouth).
"Every Girl, A Slayer," which plays whilst Buffy proposes her idea to overcome the First.
Douglas Romayne[edit]
Main article: Douglas Romayne
Douglas Romayne was hired by Joss Whedon to score episodes of Buffy season 7. He scored additional music for 33 episodes of Angel seasons four and five for Robert J. Kral. He was Associate Music Director on the Buffy musical "Once More, with Feeling," which was nominated for an Emmy for Outstanding Music Direction (C. Beck and J. Tobias nominated). He also arranged Joss Whedon's song "Mrs" in "Selfless." Memorable tracks have included:
"In Westbury Field" which plays when Giles is introduced in England ("Lessons")
The suite of "Istanbul" "Just In Time" and "Lesson One" is the score to the opening sequence of the season - from a slayer being chased and murdered in Turkey to Buffy training Dawn to fight vampires to Buffy's worries about the new school ("Lessons")
"On The Mend" which plays when Buffy helps Willow heal from the Gnarl ("Same Time, Same Place")
"Another Way Out" which plays when Angel, Wes, Lilah, Fred and Connor are trying to escape the Beast, set loose at Wolfram and Hart ("Habeas Corpses")
"Chasing Lilah" ("Calvary")
"Cordy's Journey" which plays over a montage of scenes showing that Cordy is possessed ("Inside Out")
"The Hellmouth" which plays when Spike magically appears in Angel's office straight from the Buffy series finale ("Unleashed")
"Puppet Fight" which plays when Angel, transformed into a puppet, has a showdown with evil puppets taking over the lives of children in LA ("Smile Time")
"One Day To Live" which plays while Angel and the gang try to save Fred from dying of a mystical disease ("A Hole in the World")
Music from Buffy, Angel and other television series and films can be heard at Romayne's official website.
The Bronze[edit]
The Bronze is a nightclub in Sunnydale. It hosts live music and serves as a key place which the Scooby Gang spend time for leisure. In the opening episodes, Sprung Monkey and Dashboard Prophets establish that much of the music featured at the club is of the alternative rock and roll, though numerous styles of music are also later heard, such as trip hop, ballad and new wave/electronic music.
The Bronze appears in 66 of the 144 episodes of Buffy. The Bronze acted as a way for the production team to showcase new bands from the Los Angeles area, as well as more well known artists and bands such as Aimee Mann, Cibo Matto and Michelle Branch.
Some key musical moments at the Bronze:
Buffy and Angel dancing to Sophie Zelmani's "I'll Remember You" ("Angel")
Buffy taunting Angel by intimately dancing with Xander to Cibo Matto's "Sugar Water" ("When She Was Bad")
Buffy and Faith flirtatiously dancing to Curve's "Chinese Burn" ("Bad Girls")
Jonathan, the superstar, performs various hits from Royal Crown Revue ("Superstar")
Oz along with the other Scoobies hears the band, "Shy" (a fictional front for the real-life band, T.H.C.), and their lead singer Veruca ("Beer Bad"/"Wild at Heart")
Giles performs 'Giles's Epiphany' along with Christophe Beck and Four Star Mary ("Restless")
Willow and Tara dance together during Melanie Doane's "I Can't Take My Eyes Off You" ("Family")
Michelle Branch plays "Goodbye to You" - A performance which became particularly well noted amongst fans due to the meaning of the episode ("Tabula Rasa" featured the break-up of Willow and Tara; the departure of Giles to England; and the beginning of Buffy and Spike's relationship). The version used on the show was slightly different from the radio version. ("Tabula Rasa")
The Breeders performed in the episode "Him " that aired November 5, 2002.
Angie Hart performs "Blue" with her band Splendid. The song was co-written by Hart and Joss Whedon ("Conversations with Dead People")
Aimee Mann performs "This Is How It Goes" and "Pavlov's Bell" ("Sleeper")
Dingoes Ate My Baby[edit]
Oz's band Dingoes Ate My Baby appeared on the show many times. The name alludes to the widespread news coverage of the death of nine-week-old Azaria Chamberlain in Australia in 1980. The real band that provided the music for Dingoes Ate My Baby was Four Star Mary; the song titles below are Four Star Mary tracks.
Episode
Location
Songs
"Inca Mummy Girl" The Bronze "Fate" & "Shadows"
"Bewitched, Bothered and Bewildered" The Bronze "Pain"
"Dead Man's Party" Buffy's House "Never mind," "Pain" & "Sway"
"Homecoming" The Bronze "She knows"
"Band Candy" The Bronze "Violent"
"Revelations" The Bronze "Run"
"Living Conditions" Buffy's dorm room "Pain" (on stereo)
"The Harsh Light of Day" The Bronze "Dilate"
"The Initiative" The Bronze "Fate" (on stereo)
Lorne and Caritas[edit]
Main articles: Lorne & Caritas
Lorne was born in a dimension called Pylea, a world without music. In 1996, he happened upon a dimensional portal and was sucked through it. He landed in Los Angeles and discovered music and culture. Soon after he opened up a karaoke bar, Caritas. He honed the ability to read people's auras when they sang and he could give those people direction.
Some key musical moments include:
The first song from Caritas seen on screen is Lorne singing Gloria Gaynor's "I Will Survive" ("Judgment"). Lorne was seen in many more Season 2 episodes singing at his bar.
In the same episode, Angel, a long-time fan of Barry Manilow, sang "Mandy." When he needed guidance once again he chose Wang Chung's "Everybody Have Fun Tonight" ("Dear Boy").
Cordy, Wes and Gunn drunkenly sing Queen's "We Are The Champions" after they have been sacked from Angel Investigations ("Redefinition").
Darla sings Ella Fitzgerald's "Ill Wind" when asking Lorne for advice. ("The Trial") .
Lindsey McDonald sings "L.A. Song." The music and lyrics written by David Greenwalt ("Dead End"). Eve also sang a little bit of this to Lorne to make sure she didn't have anything to do with what was happening to Fred ("A Hole In The World") .
The last performance seen at Caritas was Patsy Cline's "Crazy" sung by Fred ("That Old Gang of Mine").
After the destruction of Caritas, in the Angel finale, Lorne visits another bar and sings Regina Belle's "If I Ruled The World" ("Not Fade Away").
Shy[edit]
After Veruca makes a brief solo appearance ("Living Conditions"), her band Shy appears in two episodes.[6] Shy performs "Overfire" at The Bronze in "Beer Bad," and plays two other songs, "Dip" and "Need to Destroy" (rehearsal), in "Wild at Heart."[7] Shy's music was actually done by THC, a Los Angeles-based trip hop duo consisting of singer/lyricist Sarah Folkman and producer/composer/musician George Sarah. The three songs featured on Buffy appear on THC's 1999 album Adagio.[8]
Soundtrack[edit]
Velvet Chain[edit]
Main article: The Buffy EP
The Buffy EP was released by Velvet Chain in February 1999, following their appearance in "Never Kill a Boy on the First Date." The title track, "Buffy" is a retelling of the relationship between Buffy and Angel during seasons 1 and 2 of the show. The album also features the songs, "Strong" and "Treason," the two songs the band played during their appearance on the show.
Sarah McLachlan[edit]
Main article: Sarah McLachlan
Sarah McLachlan's music is used twice on Buffy at two key moments (finales of Season 2 and 6).
Full of Grace
In Becoming, Part Two "Full of Grace" plays when Buffy leaves Sunnydale. The song and Buffy's abandonment signify the character reaching an all-time low.
Prayer of St. Francis
At the end of the sixth season in Grave, "Prayer of St. Francis" plays as Buffy climbs out of the ground with Dawn. The song is all about overcoming that which is wrong, and renewal.
"The song, appears in the U.S. version of the Radio Sunnydale (12 songs), but not in the UK one (21 songs).
Notes and references[edit]
Jump up ^ Members.lycos.co.uk - Interview with Nerf Herder
Jump up ^ Springer, Matt, "Slayerpalooza," from Buffy the Vampire Slayer magazine #1 (UK, Oct 1999), page 18.
Jump up ^ Websites.cable.ntl.com - Interview with Darling Violetta lead guitarist
Jump up ^ "Buffy the Vampire Slayer: The Score [SOUNDTRACK]". Amazon.com. Retrieved 2008-08-18.
Jump up ^ bluntinstrument.org.uk Bluntinstrument.org.uk - Review of Thomas's work on Buffy
Jump up ^ Childs, T. Mike (2004). The Rocklopedia Fakebandica (1st ed.). New York: St. Martin's Press. p. 197. ISBN 978-0312329440.
Jump up ^ Tracy, Kathleen (2003). The Girl's Got Bite: The Original Unauthorized Guide to Buffy's World (Revised ed.). New York: St. Martin's Press. pp. 221–2. ISBN 978-0312312589.
Jump up ^ Dial-Driver, Emily; Emmons-Featherston, Sally; Ford, Jim; Taylor, Carolyn Anne (2008). The Truth of Buffy: Essays on Fiction Illuminating Reality (1st ed.). Jefferson, N.C.: McFarland & Company. p. 223. ISBN 978-0786437993.
External links[edit]
Bluntinstrument.org - Buffyverse music
Themes[edit]
Slayageonline.com - Music, Gender, and Identity in Buffy the Vampire Slayer and Angel
Darling Violetta[edit]
Websites.cable.ntl.com - Interview with Darling Violetta lead singer
Websites.cable.ntl.com - Interview with Darling Violetta lead guitarist
The11thhour.com - Interview with Darling Violetta
Nerf Herder[edit]
Members.lycos.co.uk - Interview with Nerf Herder
Composers[edit]
Bluntinstrument.org - Christophe Beck
Cityofangel.com - Interview with Robert Kral
Bbc.co.uk - Interview with Robert Kral
DouglasRomayne.com - Official site
Buffyfans.com Music Section
Duncanmusic.com - Official site
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Retrieved from "https://en.wikipedia.org/w/index.php?title=Music_in_Buffy_the_Vampire_Slayer_and_Angel&oldid=809577955"
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