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Kentrosaurus

Kentrosaurus (/ˌkɛntrˈsɔːrəs/ KEN-troh-SOR-əs; lit.'prickle lizard') is a genus of stegosaurid dinosaur from the Late Jurassic in Lindi Region of Tanzania. The type species is K. aethiopicus, named and described by German palaeontologist Edwin Hennig in 1915. Often thought to be a "primitive" member of the Stegosauria, several recent cladistic analyses find it as more derived than many other stegosaurs, and a close relative of Stegosaurus from the North American Morrison Formation within the Stegosauridae.

Kentrosaurus
Temporal range: Late Jurassic (Tithonian), 152 Ma
Mounted skeleton, Museum für Naturkunde, Berlin
Scientific classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Clade: Dinosauria
Clade: Ornithischia
Clade: Thyreophora
Clade: Stegosauria
Family: Stegosauridae
Genus: Kentrosaurus
Hennig, 1915
Type species
Kentrosaurus aethiopicus
Hennig, 1915
Synonyms
  • Kentrurosaurus
    Hennig, 1916
  • Doryphorosaurus
    Nopcsa, 1916

Fossils of K. aethiopicus have been found only in the Tendaguru Formation, dated to the late Kimmeridgian and early Tithonian ages, about 152 million years ago. Hundreds of bones were unearthed by German expeditions to German East Africa between 1909 and 1912. Although no complete skeletons are known, the remains provided a nearly complete picture of the build of the animal. In the Tendaguru Formation, it coexisted with a variety of dinosaurs such as the carnivorous theropods Elaphrosaurus and Veterupristisaurus, giant herbivorous sauropods Giraffatitan and Tornieria, and the dryosaurid Dysalotosaurus.

Kentrosaurus generally measured around 4–4.5 metres (13–15 ft) in length as an adult, and weighed about 700–1,600 kilograms (1,500–3,500 lb). It walked on all fours with straight hindlimbs. It had a small, elongated head with a beak used to bite off plant material that would be digested in a large gut. It had a, probably double, row of small plates running down its neck and back. These plates gradually merged into spikes on the hip and tail. The longest spikes were on the tail end and were used to actively defend the animal. There also was a long spike on each shoulder. The thigh bones come in two different types, suggesting that one sex was larger and more stout than the other.

Discovery and naming edit

 
Outdated skeletal mount (lectotype and paralectotypes), Museum für Naturkunde. This mount was erected in 1925 and was disassembled in 2006. In 2007 it was reassembled with a slightly altered posture

The first fossils of Kentrosaurus were discovered by the German Tendaguru Expedition in 1909, recognised as belonging to a stegosaur by expedition leader Werner Janensch on 24 July 1910, and described by German palaeontologist Edwin Hennig in 1915.[1] The name Kentrosaurus was coined by Hennig and comes from the Greek kentron/κέντρον, meaning "sharp point" or "prickle", and sauros/σαῦρος meaning "lizard",[2] Hennig added the specific name aethiopicus to denote the provenance from Africa.[1] Soon after its description, a controversy arose over the stegosaur's name, which is very similar to the ceratopsian Centrosaurus. Under the rules of biological nomenclature, forbidding homonymy, two animals may not be given the same name. Hennig renamed his stegosaur Kentrurosaurus, "pointed-tail lizard", in 1916,[3] while Hungarian paleontologist Franz Nopcsa renamed the genus Doryphorosaurus, "lance-bearing lizard", the same year.[4][5] If a renaming had been necessary, Hennig's would have had priority.[6] However, because the spelling is different, both Doryphorosaurus and Kentrurosaurus are unneeded replacement names; Kentrosaurus remains the valid name for the genus with Kentrurosaurus and Doryphorosaurus being its junior objective synonyms.[7]

 
Endocasts of the sacral ganglion and brain at Museum of Paleontology of Tübingen.

Although no complete individuals were found, some material was discovered in association, including a nearly complete tail, hip, several dorsal vertebrae and some limb elements of one individual. These form the core of a mount in the Museum für Naturkunde by Janensch.[8] The mount was dismantled during the museum renovation in 2006/2007, and re-mounted in an improved pose by Research Casting International.[9] Some other material, including a braincase and spine, was thought to have been misplaced or destroyed during World War II.[10] However, all the supposedly lost cranial material was later found in a drawer of a basement cupboard.[11]

From 1909 onwards, Kentrosaurus remains were uncovered in four quarries in the Mittlere Saurierschichten (Middle Saurian Beds) and one quarry in the Obere Saurierschichten (Upper Saurian Beds).[12] During four field seasons, the German Expedition found over 1200 bones of Kentrosaurus, belonging to about fifty individuals,[13] many of which were destroyed during the Second World War.[14] Today, almost all remaining material is housed in the Museum für Naturkunde Berlin (roughly 350 remaining specimens), while the museum of the Institute for Geosciences of the University of Tübingen houses a composite mount, roughly 50% of it being original bones.[12]

 
Lateral view of a skeleton on display at the Museum für Naturkunde, Berlin

In the original description, Hennig did not designate a holotype specimen. However, in a detailed monography on the osteology, systematic position and palaeobiology of Kentrosaurus in 1925, Hennig picked the most complete partial skeleton, today inventorised as MB.R.4800.1 through MB.R.4800.37, as a lectotype (see syntype).[15][16] This material includes a nearly complete series of tail vertebrae, several vertebrae of the back, a sacrum with five sacral vertebrae and both ilia, both femora and an ulna, and is included in the mounted skeleton at the Museum für Naturkunde in Berlin, Germany. The type locality is Kindope, Tanzania, north of Tendaguru hill.[16]

Unaware that Hennig had already defined a lectotype, Peter Galton[17] selected two dorsal vertebrae, specimens MB.R.1930 and MB.R.1931, from the material figured in Hennig's 1915 description, as 'holotypes'. This definition of a holotype is not valid, because Hennig's selection has priority. In 2011, Heinrich Mallison clarified that all the material known to Hennig in 1915, i.e. all the bones discovered before 1912, when Hermann Heck concluded the last German excavations, are paralectotypes, and that MB.R.4800 is the correct lectotype.[18]

Description edit

 
Life restoration

Kentrosaurus was a small stegosaur. It had the typical dinosaurian body bauplan, characterised by a small head, a long neck, short forelimbs and long hindlimbs, and a long, horizontal and muscular tail. Typical stegosaurid traits included the elongation and flatness of the head, the powerful build of the forelimbs, erect and pillar-like hindlimbs and an array of plates and spikes running along both sides of the top mid-line of the animal.[15]

Size and posture edit

 
Size compared to a human

Kentrosaurus aethiopicus was a relatively small stegosaur, reaching 4–4.5 m (13–15 ft) in length and 700–1,600 kg (1,500–3,500 lb) in body mass.[8][A][20][21] Some specimens suggest that relatively larger individuals could have existed.[1][13] These specimens are comparable to some Stegosaurus specimens in terms of the olecranon process in development.[22]

The long tail of Kentrosaurus results in a position of the center of mass that is unusually far back for a quadrupedal animal. It rests just in front of the hip, a position usually seen in bipedal dinosaurs. However, the femora are straight in Kentrosaurus, as opposed to typical bipeds, indicating a straight and vertical limb position. Thus, the hindlimbs, though powered by massive thigh muscles attached to a long ilium, did not support the animal alone, and the very robust forelimbs took up 10 to 15% of the bodyweight.[9] The center of mass was not heavily modified by the osteoderms (bony structures in skin) in Kentrosaurus or Stegosaurus, which allowed the animals to stay mobile despite their armament. The hindlimbs’ thigh muscles were very powerful, allowing Kentrosaurus to reach a tripod stance on its hindlegs and tail.[23]

Skull and dentition edit

Eight specimens from the skull, mandible, and teeth have been collected and described from the Tendaguru Formation, most of them being isolated elements.[24] Two quadrates (bones from the jaw joint) were referred to Kentrosaurus, but they instead belong to a juvenile brachiosaurid.[25]

 
Braincase of Kentrosaurus in lateral view.

The long and narrow skull was small in proportion to the body. It had a small antorbital fenestra, the hole between the nose and eye common to most archosaurs, including modern birds, though lost in extant crocodylians. The skull's low position suggests that Kentrosaurus may have been a browser of low-growing vegetation. This interpretation is supported by the absence of premaxillary teeth and their likely replacement by a horny beak or rhamphotheca. The presence of a beak extended along much of the jaws may have precluded the presence of cheeks in stegosaurs.[26] Due to its phylogenetic position, it is unlikely that Kentrosaurus had an extensive beak like Stegosaurus and it instead probably had a beak restricted to the jaw tips.[27][28] Other researchers have interpreted these ridges as modified versions of similar structures in other ornithischians which might have supported fleshy cheeks, rather than beaks.[7]

There are two nearly complete braincases known from Kentrosaurus though they exhibit some taphonomic distortion.[24] The frontals and parietals are flat and broad, with the latter bearing two transversely concave ventral sides with a ridge running down the middle that divides them. The lateral surface of the frontals form part of the orbit (eye socket) and the medial side creates the anterior part of the endocranial cavity (braincase). Basioccipitals (where the skull articulated with the cervical vertebrae) form the posterior floor of the brain and the occipital condyle, which is large and spherical in Kentrosaurus. The rest of the braincase is formed by the presphenoid composing the anterior end. The overall braincase morphology is very similar to those of Tuojiangosaurus, Huayangosaurus, and Stegosaurus. However, the occipital condyle is a closer distance to the basisphenoid tubera (bone at the front of the braincase) in Kentrosaurus and Huayangosaurus than in Tuojiangosaurus and some specimens of Stegosaurus. Due to dinosaurs having more molding in their braincases, endocasts of Kentrosaurus can be reconstructed using the preserved fossils. The brain is relatively short, deep, and small, with a strong cerebral and pontine flexures and a steeply inclined posterodorsal edge when compared to those of other ornithischians. There is a small dorsal projection in the endocast where an unossified (lacking bone) region between the top of the supraoccipital (bone at the top-back of the braincase) and overlying parietal that was likely covered in cartilage. This characteristic is seen in other ornithischians. Because of the prominent flexures, many of the aspects of the brain can only be interpreted by the present structures.[24]

 
Skull and neck of the Berlin mount

In the mandible (lower jaw), only an incomplete right dentary is known from Kentrosaurus.[29] The deep dentary is almost identical in shape to that of Stegosaurus, albeit much smaller. Similarly, the tooth is a typical stegosaurian tooth, small with a widened base and vertical grooves creating five ridges. The dentary has 13 preserved alvelovi on the dorsomedial side and it they are slightly convex in lateral and dorsal views. On the surface adjacent to the alveoli, there is a shallow groove bearing small foramina (small openings in bone) that is similar to grooves on the dentary of the Cretaceous neornithischian Hypsilophodon, with one foramina per tooth position. Stegosaurian teeth were small, triangular, and flat; wear facets show that they did grind their food.[30] A single complete cheek tooth is preserved, with a large crown and long root. The crown notably has fewer marginal denticles and a prominent cingulum compared to Stegosaurus, Tuojiangosaurus, and Huayangosaurus.[24]

Postcrania edit

 
A dorsal (back) vertebra of Kentrosaurus in lateral and anterior views.

The neck was composed of 13 cervical (neck) vertebrae, the first being the atlas which was strongly fused to the occipital region of the skull and followed by the axis. The other 11 cervicals had hourglass-shaped centra (the base of a vertebra) and rounded ventral keels. The diapophyses are large and strongly angled posteriorily and parallel to each other. The spinous processes got larger towards the posterior end, while the postzygapophyses became smaller and less horizontal, giving the anterior part of the neck lots of mobility laterally. The dorsal column consists of 13 dorsal (back) vertebrae which are tall and have short centra. They have a neural arch more than twice as high as the centrum, the vertebral body, and almost completely occupied by the extremely spacious neural canal, a trait unique to Kentrosaurus. The diapophyses too were laterally elongated, creating a Y-shape in anterior view. The sacrum (part of pelvis with vertebrae) consists of 6 fused centra, the first being a loose sacrodorsal, while the rest of the centra's transverse processes (extensions of bone) are fused to the dorsal parts of the sacral ribs into a solid sacral plate. The ribs also fuse to the ilium (the upper part of the pelvis) creating a fully ankylosed and solid sacrum. The ilium is notable in that the preacetabular process, front blade, of the ilium widens laterally, to the front outer side, and does not taper unlike in all other stegosaurs. Another characteristic is that the length of the ilium equals, or is greater than, that of the thigh bone.[12] The caudal (tail) vertebrae are 29 in number, though 27-29 are coossified for attachment to the thagomizers (tail spikes). The caudal vertebrae are very unique, as they have a combination of transverse processes up to the 28th vertebra and rod-shaped processes on the posterior caudals. These posterior caudal processes have narrow bases that do not tough the plate formed by the fusion of the processes of the sacral vertebrae. Kentrosaurus can be distinguished from other members of the Stegosauria by a number of processes of the vertebrae, which in the tail do not run sub-parallel, as in most dinosaurs. In the front third of the tail, they point backwards, the usual direction. In the middle tail, however, they are almost vertical, and further back they are hook-shaped and point obliquely forward. The chevrons, bones pointing to below from the bottom side of the tail vertebrae, have the shape of an inverted T.[12]

 
Sacrum of Kentrosaurus in ventral view.

The scapula (shoulder blade) is sub-rectangular, with a robust blade. Though it is not always perfectly preserved, the acromion ridge is slightly smaller than in Stegosaurus. The blade is relatively straight, although it curves towards the back. There is a small bump on the back of the blade, that would have served as the base of the triceps muscle. The coracoid is sub-circular.[31] The fore limbs were much shorter than the stocky hind limbs, which resulted in an unusual posture. The humerus (upper arm bone), like other stegosaurs, has greatly expanded proximal and distal ends that were attachment points between the coracoid and ulna-radius (forearm bones) respectively. The radius was larger than the ulna and had a wedge-shaped proximal end. The manus (hand) was small and had five toes with 2 toes bearing only a single phalange. The hindlimbs were much larger and too are similar to those of other stegosaurs. The femur (thigh bone) is the longest element in the body, with the largest known femur measuring 665 mm from the proximal to distal end. The tibia (shin bone) was wide and robust, while the fibula was skinny and thin without a greatly expanded distal end. The pes (foot) terminated in 3 toes, all of which had hoof-like unguals (claws).[15][32][24]

Armour edit

 
Lectotype, partial individual from excavation 'St' at Kindope, Tendaguru, Tanzania

Typically for a stegosaur, Kentrosaurus had extensive osteoderm (bony structures in the skin) covering, including small plates (probably located on the neck and anterior trunk), and spikes of various shapes. The spikes of Kentrosaurus are very elongated, with one specimen having a bone core length of 731 millimetres.[20] The plates have a thickened section in the middle, as if they were modified spines.[33] The spikes and plates were likely covered by horn. Aside from a few exceptions they were not found in close association with other skeletal remains. Thus, the exact position of most osteoderms is uncertain. A pair of closely spaced spikes was found articulated with a tail tip, and a number of spikes were found apparently regularly spaced in pairs along the path of an articulated tail.[13]

 
Thagomizer at the Museum of Palaeontology of Tübingen

Hennig[13] and Janensch,[8] while grouping the dermal armour elements into four distinct types, recognised an apparently continuous change of shape among them, shorter and flatter plates at the front gradually merging into longer and more pointed spikes towards the rear, suggesting an uninterrupted distribution along the entire body, in fifteen pairs.[33] Because each type of osteoderm was found in mirrored left and right versions, it seems probable that all types of osteoderms were distributed in two rows along the back of the animal, a marked contrast to the better-known North American Stegosaurus, which had one row of plates on the neck, trunk and tail, and two rows of spikes on the tail tip. There is one type of spike that differs from all others in being strongly, and not only slightly, asymmetrical, and having a very broad base. Because of bone morphology classic reconstructions placed it on the hips, at the iliac blade, while many recent reconstructions place it on the shoulder, because a similarly shaped spike is known to have existed on the shoulder in the Chinese stegosaurs Gigantspinosaurus and Huayangosaurus.[33]

Classification and species edit

Like the spikes and shields of ankylosaurs, the bony plates and spines of stegosaurians evolved from the low-keeled osteoderms characteristic of basal thyreophorans.[34] Galton (2019) interpreted plates of an armored dinosaur from the Lower Jurassic (Sinemurian-Pliensbachian) Lower Kota Formation of India as fossils of a member of Ankylosauria; the author argued that this finding indicates a probable early Early Jurassic origin for both Ankylosauria and its sister group Stegosauria.[35]

The vast majority of stegosaurian dinosaurs thus far recovered belong to the Stegosauridae, which lived in the later part of the Jurassic and early Cretaceous, and which were defined by Paul Sereno as all stegosaurians more closely related to Stegosaurus than to Huayangosaurus.[36] This group is widespread, with members across the Northern Hemisphere, Africa and possibly South America.[37] The South American remains come from Chubut, Argentina and consist only of a partial humerus, but the anatomy of the humerus is very similar to that of Kentrosaurus and both date to the Late Jurassic. In a phylogenetic analysis, the Chubut stegosaurid was recovered in polytomy with Kentrosaurus as basal stegosaurids, further suggesting that they are closely related.[37]

In Hennig's 1915 description, Kentrosaurus was assigned to the family Stegosauridae due to the preservation of dermal armor and features like posterodorsally angled neural spines on the caudal vertebrae.[1] This is confirmed by modern cladistic analyses, although in 1915 Stegosauridae was a far more inclusive concept that included some taxa now classified as ankylosaurs. A consecutive narrowing down of this concept caused Kentrosaurus, until the 1980s to be seen as a typical "primitive" stegosaurian,[38] to be placed in a more derived, higher, position in the stegosaur evolutionary tree. However, recent analyses have consistently found Kentrosaurus to be in Stegosauridae, though typically as one of the most basal genera in the family.[39][37][40] Kentrosaurus has many traits not seen in other stegosaurids but seen in basal stegosaurians, such as the presence of a parascapular spine and maxillary teeth with only seven denticles at the margin.[17][41] Cladogram of Stegosauria below that includes nearly every known stegosaur genus, recovering Kentrosaurus as a basal stegosaurid:[40]

The type and sole accepted species of Kentrosaurus is Kentrosaurus aethiopicus, named by Hennig in 1915. Fragmentary fossil material from Wyoming, named Stegosaurus longispinus by Charles Gilmore in 1914,[42] was in 1993 classified as a North American species of Kentrosaurus, as K. longispinus.[43] However, this action was not accepted by the paleontological community, and S. longispinus has been assigned to its own genus, Alcovasaurus, differing from Kentrosaurus in having more elongated tail spikes and the structure of the pelvis and vertebrae.[44][45] Cladogram of the phylogenetic analysis of Stegosauridae conducted by Maidment et al (2019), which recovers a distinct Alcovasaurus:[46]

Paleobiology edit

Feeding edit

 
Depiction of Kentrosaurus reaching for leaves on a tree

Like all ornithischians, Kentrosaurus was a herbivore. The fodder was barely chewed and swallowed in large chunks. One hypothesis on stegosaurid diet holds that they were low-level browsers, eating foliage and low-growing fruit from various non-flowering plants.[47] Kentrosaurus was capable of eating at heights of up to 1.7 m (5 ft 7 in) when on all fours. It may also have been possible for it to rear up on its hindlegs to reach vegetation higher in trees.[9]

With its centre of mass close to the hind-limbs, the animal could potentially support itself as it stood up. The hips were likely capable of allowing a vertical trunk rotation of about 60 degrees and the tail probably would either have been fully lifted, not blocking this movement or have enough curvature to rest on the ground; thus it could have provided additional support, though precisely because of this flexibility it is not certain whether much support was actually provided: it was not stiff enough to function as a "third leg" as had been suggested by Robert Thomas Bakker. In this pose, Kentrosaurus could have fed at heights of 3.3 m (11 ft).[9]

Sexual dimorphism edit

Differences in the proportions, not the size, of the femurs (thighbones) led Holly Barden and Susannah Maidment to realize that Kentrosaurus probably showed sexual dimorphism. This dimorphism of the femurs consisted in them being either more or less robust than the other. The occurrence ratio of the robust morph to the gracile one was 2:1, and it is likely that the higher percentage of animals were females. Because of this ratio, it was considered reasonable to assume that in their society, Kentrosaurus males mated with more than one female, a behaviour also found in other vertebrates.[48]

The problem posed by the ratio is that the multiple specimens studied, died in the same place, but probably not in a sudden mass-death and so do not represent a single herd or contemporary population. The results may have been distorted by a greater chance for robust animals of getting fossilised or discovered. In an earlier study by Galton in 1982, it was suggested that individual difference in the sacral rib count of both Kentrosaurus and Dacentrurus might be an indication of dimorphism: females would have had an extra pair of sacral ribs, having also the first sacral vertebra connected to the ilium, in addition to the subsequent four sacrals.[48]

Reproduction and growth edit

As the plates and spikes would have been obstacles during copulation, it is possible that pairs mated back-to-back with the female staying still in a lordosis posture as the male maneuvers his penis into her cloaca. The shoulder spikes would have made the female unable to lie on her side during mating as is proposed for Stegosaurus.[49]

In 2013, a study by Ragna Redelstorff e.a. concluded that the bone histology of Kentrosaurus indicated that it had a higher growth rate than reported for Stegosaurus and Scutellosaurus, in view of the relatively rapid deposition of highly vascularised fibrolamellar bone. As Stegosaurus was larger than Kentrosaurus, this contradicts the general rule that larger dinosaurs grew quicker than smaller ones.[50]

Defense edit

 
Thagomizer on the Museum für Naturkunde Berlin mount

Because the tail had at least forty caudal vertebrae,[13] it was highly mobile.[9] It could possibly swing at an arc of 180 degrees, covering the entire half circle behind it.[20][9] Swing speeds at the tail end may have been as high as 50 km/h. Continuous rapid swings would have allowed the spikes to slash open the skin of its attacker or to stab the soft tissues and break the ribs or facial bones. More directed blows would have resulted in the sides of the spikes fracturing even sturdy longbones of the legs by blunt trauma. These attacks would have crippled small and medium-sized theropods and may even have done some damage to large ones.[20] Earlier interpretations of the defensive behaviour of Kentrosaurus included the suggestion that the animal might have charged to the rear, to run through attackers with its spines, in the way of modern porcupines.[38]

Though Kentrosaurus likely stood with forelimbs erect like in other dinosaurs, it is hypothesised that the animal adopted a sprawling posture when defending itself. Its neck was flexible enough to allow it to keep sight of predators, as it could reach the sides of its body with its snout and look over the back. In addition, the posterior position of the center of mass may not have been advantageous for rapid locomotion, but meant that the animal could quickly rotate around the hips by pushing sideways with the arms, keeping the tail pointed at the attacker.[9] Kentrosaurus was nevertheless not invulnerable. A quick predator could have made it to the tail base (where the impact speed would be much lower) when the tail passed and the neck and upper-part of the body would have been unprotected by the tail swings. A successful predation of Kentrosaurus may have required group hunting. Compared to the more robust spikes of Stegosaurus, the thinner spikes of Kentrosaurus were at greater risk of bending.[20]

Paleoecology edit

Kentrosaurus lived in what is now Tanzania in the Late Jurassic Tendaguru Formation. The main Kentrosaurus quarries were located in the Middle Saurian Beds dating from the upper Kimmeridgian. Some remains were found in the Upper Saurian Beds dating from the Tithonian.[51] Since 2012, the boundary between the Kimmeridgian and Tithonian is dated at 152.1 million year ago.[52]

The Tendaguru ecosystem primarily consisted of three types of environment: shallow, lagoon-like marine environments, tidal flats and low coastal environments; and vegetated inland environments. The marine environment existed above the fair weather wave base and behind siliciclastic and ooid barriers. It appeared to have had little change in salinity levels and experienced tides and storms. The coastal environments consisted of brackish coastal lakes, ponds and pools. These environments had little vegetation and were probably visited by herbivorous dinosaurs mostly during droughts. The well vegetated inlands were dominated by conifers. Overall, the Late Jurassic Tendaguru climate was subtropical to tropical with seasonal rains and pronounced dry periods. During the Early Cretaceous, the Tendaguru became more humid.[53] The Tendaguru Beds are similar to the Morrison Formation of North America except in its marine interbeds.[54]

Kentrosaurus would have coexisted with fellow ornithischians like Dysalotosaurus lettowvorbecki; the sauropods Giraffatitan brancai, Dicraeosaurus hansemanni and D. sattleri, Janenschia africana, Tendaguria tanzaniensis and Tornieria africanus; theropods "Allosaurus" tendagurensis, "Ceratosaurus" roechlingi, "Torvosaurus" ingens, Elaphrosaurus bambergi, Veterupristisaurus milneri and Ostafrikasaurus crassiserratus; and the pterosaur Tendaguripterus recki.[55][56][57][58] Other organisms that inhabited the Tendaguru included corals, echinoderms, cephalopods, bivalves, gastropods, decapods, sharks, neopterygian fish, crocodilians and small mammals like Brancatherulum tendagurensis.[59]

See also edit

Notes edit

  1. ^ p. 223 in Paul (2010)[19]

References edit

  1. ^ a b c d Hennig, E. (1915). "Kentrosaurus aethiopicus, der Stegosauride des Tendaguru [Kentrosaurus aethiopicus, the stegosaur of the Tendaguru]" (PDF). Sitzungsberichte der Gesellschaft Naturforschender Freunde zu Berlin. 1915: 219–247.
  2. ^ Liddell, Henry George and Robert Scott (1980). A Greek-English Lexicon (Abridged ed.). United Kingdom: Oxford University Press. ISBN 978-0-19-910207-5.
  3. ^ Hennig, E. (1916). "Zweite Mitteilung über den Stegosauriden vom Tendaguru [Second report on the stegosaurid of the Tendaguru]". Sitzungsberichte der Gesellschaft Naturforschender Freunde zu Berlin (in German). 1916 (6): 175–182.
  4. ^ Nopcsa, Baron. F. (1915). "Die Dinosaurier der Siebenbürgischen Landesteile Ungarns [The dinosaurs of the Siebenbürgen part of the Hungarian Empire]". Mitteilungen aus dem Jahrbuche der Königlich Ungarischen Geologischen Reichsanstalt (in German). 23: 1–26.
  5. ^ Nopcsa, F. (1916). "Doryphorosaurus nov. nom. für Kentrosaurus HENNIG 1915". Centralblatt für Mineralogie, Geologie und Paläontologie. 1916: 511–512.
  6. ^ Hennig, E. (1916). "Kentrurosaurus, non Doryphorosaurus". Centralblatt für Mineralogie, Geologie und Paläontologie (in German). 1916: 578.
  7. ^ a b Galton PM, Upchurch P (2004). "Stegosauria". In Weishampel DB, Dodson P, Osmólska H (eds.). The Dinosauria (2nd ed.). University of California Press. ISBN 978-0-520-24209-8.
  8. ^ a b c Janensch, W. (1925). "Ein aufgestelltes Skelett des Stegosauriers Kentrurosaurus aethiopicus HENNIG 1915 aus den Tendaguru-Schichten Deutsch-Ostafrikas [A mounted skeleton of the Stegosaur Kentrurosaurus aethiopicus HENNIG 1915 from the Tendaguru layers of German East Africa]". Palaeontographica. Supplement 7 (in German): 257–276.
  9. ^ a b c d e f g Mallison, H. (2010). "CAD assessment of the posture and range of motion of Kentrosaurus aethiopicus HENNIG 1915". Swiss Journal of Geosciences. 103 (2): 211–233. doi:10.1007/s00015-010-0024-2. S2CID 132746786.
  10. ^ Glut, Donald F. (1997). "Kentrosaurus". Dinosaurs: The Encyclopedia. Jefferson, North Carolina: McFarland & Co. pp. 516–519. ISBN 978-0-89950-917-4.
  11. ^ Galton, P.M. (1988). "Skull bones and endocranial casts of stegosaurian dinosaur Kentrosaurus HENNIG, 1915 from Upper Jurassic of Tanzania, East Africa". Geologica et Palaeontologica. 22: 123–143.
  12. ^ a b c d Mallison, H. (2011). "The real lectotype of Kentrosaurus aethiopicus HENNIG 1915" (PDF). Neues Jahrbuch für Geologie und Paläontologie. 259 (2): 197–206. doi:10.1127/0077-7749/2011/0114.
  13. ^ a b c d e Hennig, E. (1925). "Kentrurosaurus aethiopicus. Die Stegosaurier-Funde vom Tendaguru, Deutsch-Ostafrika [Kentrurosaurus aethiopicus. The Stegosaur find from Tendaguru, German East-Africa]". Palaeontographica. Supplement 7 (in German): 101–254.
  14. ^ Maier, G (2003). African Dinosaurs Unearthed. The Tendaguru Expeditions. Bloomington: Indiana University Press. p. 432. ISBN 978-0-253-34214-0.
  15. ^ a b c Hennig, E. (1925). Kentrurosaurus aethiopicus; die Stegosaurierfunde vom Tendaguru, Deutsch-Ostafrika. Palaeontographica-Supplementbände, 101-254.
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External links edit

  • from Thescelosaurus.com (Includes details on Kentrosaurus, its junior synonyms, and other material)

kentrosaurus, confused, with, centrosaurus, ɔːr, troh, prickle, lizard, genus, stegosaurid, dinosaur, from, late, jurassic, lindi, region, tanzania, type, species, aethiopicus, named, described, german, palaeontologist, edwin, hennig, 1915, often, thought, pri. Not to be confused with Centrosaurus Kentrosaurus ˌ k ɛ n t r oʊ ˈ s ɔːr e s KEN troh SOR es lit prickle lizard is a genus of stegosaurid dinosaur from the Late Jurassic in Lindi Region of Tanzania The type species is K aethiopicus named and described by German palaeontologist Edwin Hennig in 1915 Often thought to be a primitive member of the Stegosauria several recent cladistic analyses find it as more derived than many other stegosaurs and a close relative of Stegosaurus from the North American Morrison Formation within the Stegosauridae KentrosaurusTemporal range Late Jurassic Tithonian 152 Ma PreꞒ Ꞓ O S D C P T J K Pg N Mounted skeleton Museum fur Naturkunde Berlin Scientific classification Domain Eukaryota Kingdom Animalia Phylum Chordata Clade Dinosauria Clade Ornithischia Clade Thyreophora Clade Stegosauria Family Stegosauridae Genus KentrosaurusHennig 1915 Type species Kentrosaurus aethiopicusHennig 1915 Synonyms Kentrurosaurus Hennig 1916 Doryphorosaurus Nopcsa 1916 Fossils of K aethiopicus have been found only in the Tendaguru Formation dated to the late Kimmeridgian and early Tithonian ages about 152 million years ago Hundreds of bones were unearthed by German expeditions to German East Africa between 1909 and 1912 Although no complete skeletons are known the remains provided a nearly complete picture of the build of the animal In the Tendaguru Formation it coexisted with a variety of dinosaurs such as the carnivorous theropods Elaphrosaurus and Veterupristisaurus giant herbivorous sauropods Giraffatitan and Tornieria and the dryosaurid Dysalotosaurus Kentrosaurus generally measured around 4 4 5 metres 13 15 ft in length as an adult and weighed about 700 1 600 kilograms 1 500 3 500 lb It walked on all fours with straight hindlimbs It had a small elongated head with a beak used to bite off plant material that would be digested in a large gut It had a probably double row of small plates running down its neck and back These plates gradually merged into spikes on the hip and tail The longest spikes were on the tail end and were used to actively defend the animal There also was a long spike on each shoulder The thigh bones come in two different types suggesting that one sex was larger and more stout than the other Contents 1 Discovery and naming 2 Description 2 1 Size and posture 2 2 Skull and dentition 2 3 Postcrania 2 4 Armour 3 Classification and species 4 Paleobiology 4 1 Feeding 4 2 Sexual dimorphism 4 3 Reproduction and growth 4 4 Defense 5 Paleoecology 6 See also 7 Notes 8 References 9 External linksDiscovery and naming edit nbsp Outdated skeletal mount lectotype and paralectotypes Museum fur Naturkunde This mount was erected in 1925 and was disassembled in 2006 In 2007 it was reassembled with a slightly altered postureThe first fossils of Kentrosaurus were discovered by the German Tendaguru Expedition in 1909 recognised as belonging to a stegosaur by expedition leader Werner Janensch on 24 July 1910 and described by German palaeontologist Edwin Hennig in 1915 1 The name Kentrosaurus was coined by Hennig and comes from the Greek kentron kentron meaning sharp point or prickle and sauros saῦros meaning lizard 2 Hennig added the specific name aethiopicus to denote the provenance from Africa 1 Soon after its description a controversy arose over the stegosaur s name which is very similar to the ceratopsian Centrosaurus Under the rules of biological nomenclature forbidding homonymy two animals may not be given the same name Hennig renamed his stegosaur Kentrurosaurus pointed tail lizard in 1916 3 while Hungarian paleontologist Franz Nopcsa renamed the genus Doryphorosaurus lance bearing lizard the same year 4 5 If a renaming had been necessary Hennig s would have had priority 6 However because the spelling is different both Doryphorosaurus and Kentrurosaurus are unneeded replacement names Kentrosaurus remains the valid name for the genus with Kentrurosaurus and Doryphorosaurus being its junior objective synonyms 7 nbsp Endocasts of the sacral ganglion and brain at Museum of Paleontology of Tubingen Although no complete individuals were found some material was discovered in association including a nearly complete tail hip several dorsal vertebrae and some limb elements of one individual These form the core of a mount in the Museum fur Naturkunde by Janensch 8 The mount was dismantled during the museum renovation in 2006 2007 and re mounted in an improved pose by Research Casting International 9 Some other material including a braincase and spine was thought to have been misplaced or destroyed during World War II 10 However all the supposedly lost cranial material was later found in a drawer of a basement cupboard 11 From 1909 onwards Kentrosaurus remains were uncovered in four quarries in the Mittlere Saurierschichten Middle Saurian Beds and one quarry in the Obere Saurierschichten Upper Saurian Beds 12 During four field seasons the German Expedition found over 1200 bones of Kentrosaurus belonging to about fifty individuals 13 many of which were destroyed during the Second World War 14 Today almost all remaining material is housed in the Museum fur Naturkunde Berlin roughly 350 remaining specimens while the museum of the Institute for Geosciences of the University of Tubingen houses a composite mount roughly 50 of it being original bones 12 nbsp Lateral view of a skeleton on display at the Museum fur Naturkunde Berlin In the original description Hennig did not designate a holotype specimen However in a detailed monography on the osteology systematic position and palaeobiology of Kentrosaurus in 1925 Hennig picked the most complete partial skeleton today inventorised as MB R 4800 1 through MB R 4800 37 as a lectotype see syntype 15 16 This material includes a nearly complete series of tail vertebrae several vertebrae of the back a sacrum with five sacral vertebrae and both ilia both femora and an ulna and is included in the mounted skeleton at the Museum fur Naturkunde in Berlin Germany The type locality is Kindope Tanzania north of Tendaguru hill 16 Unaware that Hennig had already defined a lectotype Peter Galton 17 selected two dorsal vertebrae specimens MB R 1930 and MB R 1931 from the material figured in Hennig s 1915 description as holotypes This definition of a holotype is not valid because Hennig s selection has priority In 2011 Heinrich Mallison clarified that all the material known to Hennig in 1915 i e all the bones discovered before 1912 when Hermann Heck concluded the last German excavations are paralectotypes and that MB R 4800 is the correct lectotype 18 Description edit nbsp Life restoration Kentrosaurus was a small stegosaur It had the typical dinosaurian body bauplan characterised by a small head a long neck short forelimbs and long hindlimbs and a long horizontal and muscular tail Typical stegosaurid traits included the elongation and flatness of the head the powerful build of the forelimbs erect and pillar like hindlimbs and an array of plates and spikes running along both sides of the top mid line of the animal 15 Size and posture edit nbsp Size compared to a human Kentrosaurus aethiopicus was a relatively small stegosaur reaching 4 4 5 m 13 15 ft in length and 700 1 600 kg 1 500 3 500 lb in body mass 8 A 20 21 Some specimens suggest that relatively larger individuals could have existed 1 13 These specimens are comparable to some Stegosaurus specimens in terms of the olecranon process in development 22 The long tail of Kentrosaurus results in a position of the center of mass that is unusually far back for a quadrupedal animal It rests just in front of the hip a position usually seen in bipedal dinosaurs However the femora are straight in Kentrosaurus as opposed to typical bipeds indicating a straight and vertical limb position Thus the hindlimbs though powered by massive thigh muscles attached to a long ilium did not support the animal alone and the very robust forelimbs took up 10 to 15 of the bodyweight 9 The center of mass was not heavily modified by the osteoderms bony structures in skin in Kentrosaurus or Stegosaurus which allowed the animals to stay mobile despite their armament The hindlimbs thigh muscles were very powerful allowing Kentrosaurus to reach a tripod stance on its hindlegs and tail 23 Skull and dentition edit Eight specimens from the skull mandible and teeth have been collected and described from the Tendaguru Formation most of them being isolated elements 24 Two quadrates bones from the jaw joint were referred to Kentrosaurus but they instead belong to a juvenile brachiosaurid 25 nbsp Braincase of Kentrosaurus in lateral view The long and narrow skull was small in proportion to the body It had a small antorbital fenestra the hole between the nose and eye common to most archosaurs including modern birds though lost in extant crocodylians The skull s low position suggests that Kentrosaurus may have been a browser of low growing vegetation This interpretation is supported by the absence of premaxillary teeth and their likely replacement by a horny beak or rhamphotheca The presence of a beak extended along much of the jaws may have precluded the presence of cheeks in stegosaurs 26 Due to its phylogenetic position it is unlikely that Kentrosaurus had an extensive beak like Stegosaurus and it instead probably had a beak restricted to the jaw tips 27 28 Other researchers have interpreted these ridges as modified versions of similar structures in other ornithischians which might have supported fleshy cheeks rather than beaks 7 There are two nearly complete braincases known from Kentrosaurus though they exhibit some taphonomic distortion 24 The frontals and parietals are flat and broad with the latter bearing two transversely concave ventral sides with a ridge running down the middle that divides them The lateral surface of the frontals form part of the orbit eye socket and the medial side creates the anterior part of the endocranial cavity braincase Basioccipitals where the skull articulated with the cervical vertebrae form the posterior floor of the brain and the occipital condyle which is large and spherical in Kentrosaurus The rest of the braincase is formed by the presphenoid composing the anterior end The overall braincase morphology is very similar to those of Tuojiangosaurus Huayangosaurus and Stegosaurus However the occipital condyle is a closer distance to the basisphenoid tubera bone at the front of the braincase in Kentrosaurus and Huayangosaurus than in Tuojiangosaurus and some specimens of Stegosaurus Due to dinosaurs having more molding in their braincases endocasts of Kentrosaurus can be reconstructed using the preserved fossils The brain is relatively short deep and small with a strong cerebral and pontine flexures and a steeply inclined posterodorsal edge when compared to those of other ornithischians There is a small dorsal projection in the endocast where an unossified lacking bone region between the top of the supraoccipital bone at the top back of the braincase and overlying parietal that was likely covered in cartilage This characteristic is seen in other ornithischians Because of the prominent flexures many of the aspects of the brain can only be interpreted by the present structures 24 nbsp Skull and neck of the Berlin mount In the mandible lower jaw only an incomplete right dentary is known from Kentrosaurus 29 The deep dentary is almost identical in shape to that of Stegosaurus albeit much smaller Similarly the tooth is a typical stegosaurian tooth small with a widened base and vertical grooves creating five ridges The dentary has 13 preserved alvelovi on the dorsomedial side and it they are slightly convex in lateral and dorsal views On the surface adjacent to the alveoli there is a shallow groove bearing small foramina small openings in bone that is similar to grooves on the dentary of the Cretaceous neornithischian Hypsilophodon with one foramina per tooth position Stegosaurian teeth were small triangular and flat wear facets show that they did grind their food 30 A single complete cheek tooth is preserved with a large crown and long root The crown notably has fewer marginal denticles and a prominent cingulum compared to Stegosaurus Tuojiangosaurus and Huayangosaurus 24 Postcrania edit nbsp A dorsal back vertebra of Kentrosaurus in lateral and anterior views The neck was composed of 13 cervical neck vertebrae the first being the atlas which was strongly fused to the occipital region of the skull and followed by the axis The other 11 cervicals had hourglass shaped centra the base of a vertebra and rounded ventral keels The diapophyses are large and strongly angled posteriorily and parallel to each other The spinous processes got larger towards the posterior end while the postzygapophyses became smaller and less horizontal giving the anterior part of the neck lots of mobility laterally The dorsal column consists of 13 dorsal back vertebrae which are tall and have short centra They have a neural arch more than twice as high as the centrum the vertebral body and almost completely occupied by the extremely spacious neural canal a trait unique to Kentrosaurus The diapophyses too were laterally elongated creating a Y shape in anterior view The sacrum part of pelvis with vertebrae consists of 6 fused centra the first being a loose sacrodorsal while the rest of the centra s transverse processes extensions of bone are fused to the dorsal parts of the sacral ribs into a solid sacral plate The ribs also fuse to the ilium the upper part of the pelvis creating a fully ankylosed and solid sacrum The ilium is notable in that the preacetabular process front blade of the ilium widens laterally to the front outer side and does not taper unlike in all other stegosaurs Another characteristic is that the length of the ilium equals or is greater than that of the thigh bone 12 The caudal tail vertebrae are 29 in number though 27 29 are coossified for attachment to the thagomizers tail spikes The caudal vertebrae are very unique as they have a combination of transverse processes up to the 28th vertebra and rod shaped processes on the posterior caudals These posterior caudal processes have narrow bases that do not tough the plate formed by the fusion of the processes of the sacral vertebrae Kentrosaurus can be distinguished from other members of the Stegosauria by a number of processes of the vertebrae which in the tail do not run sub parallel as in most dinosaurs In the front third of the tail they point backwards the usual direction In the middle tail however they are almost vertical and further back they are hook shaped and point obliquely forward The chevrons bones pointing to below from the bottom side of the tail vertebrae have the shape of an inverted T 12 nbsp Sacrum of Kentrosaurus in ventral view The scapula shoulder blade is sub rectangular with a robust blade Though it is not always perfectly preserved the acromion ridge is slightly smaller than in Stegosaurus The blade is relatively straight although it curves towards the back There is a small bump on the back of the blade that would have served as the base of the triceps muscle The coracoid is sub circular 31 The fore limbs were much shorter than the stocky hind limbs which resulted in an unusual posture The humerus upper arm bone like other stegosaurs has greatly expanded proximal and distal ends that were attachment points between the coracoid and ulna radius forearm bones respectively The radius was larger than the ulna and had a wedge shaped proximal end The manus hand was small and had five toes with 2 toes bearing only a single phalange The hindlimbs were much larger and too are similar to those of other stegosaurs The femur thigh bone is the longest element in the body with the largest known femur measuring 665 mm from the proximal to distal end The tibia shin bone was wide and robust while the fibula was skinny and thin without a greatly expanded distal end The pes foot terminated in 3 toes all of which had hoof like unguals claws 15 32 24 Armour edit nbsp Lectotype partial individual from excavation St at Kindope Tendaguru Tanzania Typically for a stegosaur Kentrosaurus had extensive osteoderm bony structures in the skin covering including small plates probably located on the neck and anterior trunk and spikes of various shapes The spikes of Kentrosaurus are very elongated with one specimen having a bone core length of 731 millimetres 20 The plates have a thickened section in the middle as if they were modified spines 33 The spikes and plates were likely covered by horn Aside from a few exceptions they were not found in close association with other skeletal remains Thus the exact position of most osteoderms is uncertain A pair of closely spaced spikes was found articulated with a tail tip and a number of spikes were found apparently regularly spaced in pairs along the path of an articulated tail 13 nbsp Thagomizer at the Museum of Palaeontology of Tubingen Hennig 13 and Janensch 8 while grouping the dermal armour elements into four distinct types recognised an apparently continuous change of shape among them shorter and flatter plates at the front gradually merging into longer and more pointed spikes towards the rear suggesting an uninterrupted distribution along the entire body in fifteen pairs 33 Because each type of osteoderm was found in mirrored left and right versions it seems probable that all types of osteoderms were distributed in two rows along the back of the animal a marked contrast to the better known North American Stegosaurus which had one row of plates on the neck trunk and tail and two rows of spikes on the tail tip There is one type of spike that differs from all others in being strongly and not only slightly asymmetrical and having a very broad base Because of bone morphology classic reconstructions placed it on the hips at the iliac blade while many recent reconstructions place it on the shoulder because a similarly shaped spike is known to have existed on the shoulder in the Chinese stegosaurs Gigantspinosaurus and Huayangosaurus 33 Classification and species editLike the spikes and shields of ankylosaurs the bony plates and spines of stegosaurians evolved from the low keeled osteoderms characteristic of basal thyreophorans 34 Galton 2019 interpreted plates of an armored dinosaur from the Lower Jurassic Sinemurian Pliensbachian Lower Kota Formation of India as fossils of a member of Ankylosauria the author argued that this finding indicates a probable early Early Jurassic origin for both Ankylosauria and its sister group Stegosauria 35 The vast majority of stegosaurian dinosaurs thus far recovered belong to the Stegosauridae which lived in the later part of the Jurassic and early Cretaceous and which were defined by Paul Sereno as all stegosaurians more closely related to Stegosaurus than to Huayangosaurus 36 This group is widespread with members across the Northern Hemisphere Africa and possibly South America 37 The South American remains come from Chubut Argentina and consist only of a partial humerus but the anatomy of the humerus is very similar to that of Kentrosaurus and both date to the Late Jurassic In a phylogenetic analysis the Chubut stegosaurid was recovered in polytomy with Kentrosaurus as basal stegosaurids further suggesting that they are closely related 37 In Hennig s 1915 description Kentrosaurus was assigned to the family Stegosauridae due to the preservation of dermal armor and features like posterodorsally angled neural spines on the caudal vertebrae 1 This is confirmed by modern cladistic analyses although in 1915 Stegosauridae was a far more inclusive concept that included some taxa now classified as ankylosaurs A consecutive narrowing down of this concept caused Kentrosaurus until the 1980s to be seen as a typical primitive stegosaurian 38 to be placed in a more derived higher position in the stegosaur evolutionary tree However recent analyses have consistently found Kentrosaurus to be in Stegosauridae though typically as one of the most basal genera in the family 39 37 40 Kentrosaurus has many traits not seen in other stegosaurids but seen in basal stegosaurians such as the presence of a parascapular spine and maxillary teeth with only seven denticles at the margin 17 41 Cladogram of Stegosauria below that includes nearly every known stegosaur genus recovering Kentrosaurus as a basal stegosaurid 40 Eurypoda Ankylosauria Stegosauria Huayangosauridae Huayangosaurus taibaii Chungkingosaurus jiangbeiensis Tuojiangosaurus multispinus Paranthodon africanus Stegosauridae Jiangjunosaurus junggarensis Gigantspinosaurus sichuanensis Kentrosaurus aethiopicus Dacentrurus armatus Loricatosaurus priscus Hesperosaurus mjosi Miragaia longicollum Stegosaurus stenops Wuerhosaurus homheni The type and sole accepted species of Kentrosaurus is Kentrosaurus aethiopicus named by Hennig in 1915 Fragmentary fossil material from Wyoming named Stegosaurus longispinus by Charles Gilmore in 1914 42 was in 1993 classified as a North American species of Kentrosaurus as K longispinus 43 However this action was not accepted by the paleontological community and S longispinus has been assigned to its own genus Alcovasaurus differing from Kentrosaurus in having more elongated tail spikes and the structure of the pelvis and vertebrae 44 45 Cladogram of the phylogenetic analysis of Stegosauridae conducted by Maidment et al 2019 which recovers a distinct Alcovasaurus 46 Stegosauridae Alcovasaurus Paranthodon Tuojiangosaurus Kentrosaurus Adratiklit Dacentrurus Miragaia Jiangjunosaurus Hesperosaurus Loricatosaurus StegosaurusPaleobiology editFeeding edit nbsp Depiction of Kentrosaurus reaching for leaves on a tree Like all ornithischians Kentrosaurus was a herbivore The fodder was barely chewed and swallowed in large chunks One hypothesis on stegosaurid diet holds that they were low level browsers eating foliage and low growing fruit from various non flowering plants 47 Kentrosaurus was capable of eating at heights of up to 1 7 m 5 ft 7 in when on all fours It may also have been possible for it to rear up on its hindlegs to reach vegetation higher in trees 9 With its centre of mass close to the hind limbs the animal could potentially support itself as it stood up The hips were likely capable of allowing a vertical trunk rotation of about 60 degrees and the tail probably would either have been fully lifted not blocking this movement or have enough curvature to rest on the ground thus it could have provided additional support though precisely because of this flexibility it is not certain whether much support was actually provided it was not stiff enough to function as a third leg as had been suggested by Robert Thomas Bakker In this pose Kentrosaurus could have fed at heights of 3 3 m 11 ft 9 Sexual dimorphism edit Differences in the proportions not the size of the femurs thighbones led Holly Barden and Susannah Maidment to realize that Kentrosaurus probably showed sexual dimorphism This dimorphism of the femurs consisted in them being either more or less robust than the other The occurrence ratio of the robust morph to the gracile one was 2 1 and it is likely that the higher percentage of animals were females Because of this ratio it was considered reasonable to assume that in their society Kentrosaurus males mated with more than one female a behaviour also found in other vertebrates 48 The problem posed by the ratio is that the multiple specimens studied died in the same place but probably not in a sudden mass death and so do not represent a single herd or contemporary population The results may have been distorted by a greater chance for robust animals of getting fossilised or discovered In an earlier study by Galton in 1982 it was suggested that individual difference in the sacral rib count of both Kentrosaurus and Dacentrurus might be an indication of dimorphism females would have had an extra pair of sacral ribs having also the first sacral vertebra connected to the ilium in addition to the subsequent four sacrals 48 Reproduction and growth edit As the plates and spikes would have been obstacles during copulation it is possible that pairs mated back to back with the female staying still in a lordosis posture as the male maneuvers his penis into her cloaca The shoulder spikes would have made the female unable to lie on her side during mating as is proposed for Stegosaurus 49 In 2013 a study by Ragna Redelstorff e a concluded that the bone histology of Kentrosaurus indicated that it had a higher growth rate than reported for Stegosaurus and Scutellosaurus in view of the relatively rapid deposition of highly vascularised fibrolamellar bone As Stegosaurus was larger than Kentrosaurus this contradicts the general rule that larger dinosaurs grew quicker than smaller ones 50 Defense edit nbsp Thagomizer on the Museum fur Naturkunde Berlin mount Because the tail had at least forty caudal vertebrae 13 it was highly mobile 9 It could possibly swing at an arc of 180 degrees covering the entire half circle behind it 20 9 Swing speeds at the tail end may have been as high as 50 km h Continuous rapid swings would have allowed the spikes to slash open the skin of its attacker or to stab the soft tissues and break the ribs or facial bones More directed blows would have resulted in the sides of the spikes fracturing even sturdy longbones of the legs by blunt trauma These attacks would have crippled small and medium sized theropods and may even have done some damage to large ones 20 Earlier interpretations of the defensive behaviour of Kentrosaurus included the suggestion that the animal might have charged to the rear to run through attackers with its spines in the way of modern porcupines 38 Though Kentrosaurus likely stood with forelimbs erect like in other dinosaurs it is hypothesised that the animal adopted a sprawling posture when defending itself Its neck was flexible enough to allow it to keep sight of predators as it could reach the sides of its body with its snout and look over the back In addition the posterior position of the center of mass may not have been advantageous for rapid locomotion but meant that the animal could quickly rotate around the hips by pushing sideways with the arms keeping the tail pointed at the attacker 9 Kentrosaurus was nevertheless not invulnerable A quick predator could have made it to the tail base where the impact speed would be much lower when the tail passed and the neck and upper part of the body would have been unprotected by the tail swings A successful predation of Kentrosaurus may have required group hunting Compared to the more robust spikes of Stegosaurus the thinner spikes of Kentrosaurus were at greater risk of bending 20 Paleoecology editKentrosaurus lived in what is now Tanzania in the Late Jurassic Tendaguru Formation The main Kentrosaurus quarries were located in the Middle Saurian Beds dating from the upper Kimmeridgian Some remains were found in the Upper Saurian Beds dating from the Tithonian 51 Since 2012 the boundary between the Kimmeridgian and Tithonian is dated at 152 1 million year ago 52 The Tendaguru ecosystem primarily consisted of three types of environment shallow lagoon like marine environments tidal flats and low coastal environments and vegetated inland environments The marine environment existed above the fair weather wave base and behind siliciclastic and ooid barriers It appeared to have had little change in salinity levels and experienced tides and storms The coastal environments consisted of brackish coastal lakes ponds and pools These environments had little vegetation and were probably visited by herbivorous dinosaurs mostly during droughts The well vegetated inlands were dominated by conifers Overall the Late Jurassic Tendaguru climate was subtropical to tropical with seasonal rains and pronounced dry periods During the Early Cretaceous the Tendaguru became more humid 53 The Tendaguru Beds are similar to the Morrison Formation of North America except in its marine interbeds 54 Kentrosaurus would have coexisted with fellow ornithischians like Dysalotosaurus lettowvorbecki the sauropods Giraffatitan brancai Dicraeosaurus hansemanni and D sattleri Janenschia africana Tendaguria tanzaniensis and Tornieria africanus theropods Allosaurus tendagurensis Ceratosaurus roechlingi Torvosaurus ingens Elaphrosaurus bambergi Veterupristisaurus milneri and Ostafrikasaurus crassiserratus and the pterosaur Tendaguripterus recki 55 56 57 58 Other organisms that inhabited the Tendaguru included corals echinoderms cephalopods bivalves gastropods decapods sharks neopterygian fish crocodilians and small mammals like Brancatherulum tendagurensis 59 See also edit nbsp Dinosaurs portal Timeline of stegosaur researchNotes edit p 223 in Paul 2010 19 References edit a b c d Hennig E 1915 Kentrosaurus aethiopicus der Stegosauride des Tendaguru Kentrosaurus aethiopicus the stegosaur of the Tendaguru PDF Sitzungsberichte der Gesellschaft Naturforschender Freunde zu Berlin 1915 219 247 Liddell Henry George and Robert Scott 1980 A Greek English Lexicon Abridged ed United Kingdom Oxford University Press ISBN 978 0 19 910207 5 Hennig E 1916 Zweite Mitteilung uber den Stegosauriden vom Tendaguru Second report on the stegosaurid of the Tendaguru Sitzungsberichte der Gesellschaft Naturforschender Freunde zu Berlin in German 1916 6 175 182 Nopcsa Baron F 1915 Die Dinosaurier der Siebenburgischen Landesteile Ungarns The dinosaurs of the Siebenburgen part of the Hungarian Empire Mitteilungen aus dem Jahrbuche der Koniglich Ungarischen Geologischen Reichsanstalt in German 23 1 26 Nopcsa F 1916 Doryphorosaurus nov nom fur Kentrosaurus HENNIG 1915 Centralblatt fur Mineralogie Geologie und Palaontologie 1916 511 512 Hennig E 1916 Kentrurosaurus non Doryphorosaurus Centralblatt fur Mineralogie Geologie und Palaontologie in German 1916 578 a b Galton PM Upchurch P 2004 Stegosauria In Weishampel DB Dodson P Osmolska H eds The Dinosauria 2nd ed University of California Press ISBN 978 0 520 24209 8 a b c Janensch W 1925 Ein aufgestelltes Skelett des Stegosauriers Kentrurosaurus aethiopicus HENNIG 1915 aus den Tendaguru Schichten Deutsch Ostafrikas A mounted skeleton of the Stegosaur Kentrurosaurus aethiopicus HENNIG 1915 from the Tendaguru layers of German East Africa Palaeontographica Supplement 7 in German 257 276 a b c d e f g Mallison H 2010 CAD assessment of the posture and range of motion of Kentrosaurus aethiopicus HENNIG 1915 Swiss Journal of Geosciences 103 2 211 233 doi 10 1007 s00015 010 0024 2 S2CID 132746786 Glut Donald F 1997 Kentrosaurus Dinosaurs The Encyclopedia Jefferson North Carolina McFarland amp Co pp 516 519 ISBN 978 0 89950 917 4 Galton P M 1988 Skull bones and endocranial casts of stegosaurian dinosaur Kentrosaurus HENNIG 1915 from Upper Jurassic of Tanzania East Africa Geologica et Palaeontologica 22 123 143 a b c d Mallison H 2011 The real lectotype of Kentrosaurus aethiopicus HENNIG 1915 PDF Neues Jahrbuch fur Geologie und Palaontologie 259 2 197 206 doi 10 1127 0077 7749 2011 0114 a b c d e Hennig E 1925 Kentrurosaurus aethiopicus Die Stegosaurier Funde vom Tendaguru Deutsch Ostafrika Kentrurosaurus aethiopicus The Stegosaur find from Tendaguru German East Africa Palaeontographica Supplement 7 in German 101 254 Maier G 2003 African Dinosaurs Unearthed The Tendaguru Expeditions Bloomington Indiana University Press p 432 ISBN 978 0 253 34214 0 a b c Hennig E 1925 Kentrurosaurus aethiopicus die Stegosaurierfunde vom Tendaguru Deutsch Ostafrika Palaeontographica Supplementbande 101 254 a b Janensch W 1925 Ein aufgestelltes skelett des stegosauriers Kentrurosaurus aethiopicus E Hennig aus den Tendaguru schichten Deutsch Ostafrikas Palaeontographica Supplementbande 255 276 a b Galton P M 1982 The postcranial anatomy of stegosaurian dinosaur Kentrosaurus from the Upper Jurassic of Tanzania East Africa Geologica et Palaeontologica 15 139 165 Mallison H 2011 The real lectotype of Kentrosaurus aethiopicus HENNIG 1915 Neues Jahrbuch fur Geologie und Palaontologie Abhandlungen 197 206 Paul G S 2010 The Princeton Field Guide to Dinosaurs Princeton University Press a b c d e Mallison H 2011 Defense capabilities of Kentrosaurus aethiopicus HENNIG 1915 Palaeontologia Electronica 14 2 10 Benson Roger B J Campione Nicolas E Carrano Matthew T Mannion Philip D Sullivan Corwin Upchurch Paul Evans David C 2014 05 06 Rates of Dinosaur Body Mass Evolution Indicate 170 Million Years of Sustained Ecological Innovation on the Avian Stem Lineage PLOS Biology 12 5 e1001853 doi 10 1371 journal pbio 1001853 ISSN 1545 7885 PMC 4011683 PMID 24802911 Woodruff D C Trexler D Maidment S C R 2019 Two New Stegosaur Specimens from the Upper Jurassic Morrison Formation of Montana USA Acta Palaeontologica Polonica 64 3 461 480 doi 10 4202 app 00585 2018 S2CID 201310639 Mallison H 2014 03 07 Osteoderm distribution has low impact on the centre of mass of stegosaurs Fossil Record 17 1 33 39 doi 10 5194 fr 17 33 2014 ISSN 2193 0066 a b c d e Galton P M 1988 Skull bones and endocranial casts of stegosaurian dinosaur Kentrosaurus Hennig 1915 from Upper Jurassic of Tanzania East Africa Geologica et Palaeontologica 22 123 143 Maidment S C Norman D B Barrett P M amp Upchurch P 2008 Systematics and phylogeny of Stegosauria Dinosauria Ornithischia Journal of Systematic Palaeontology 6 4 367 407 Czerkas S 1999 The beaked jaws of stegosaurs and their implications for other ornithischians Miscellaneous Publication of the Utah Geological Survey 99 1 143 150 Knoll F 2008 Buccal soft anatomy in Lesothosaurus Dinosauria Ornithischia PDF Neues Jahrbuch fur Geologie und Palaontologie Abhandlungen 248 3 355 364 doi 10 1127 0077 7749 2008 0248 0355 Barrett P M 2001 Tooth wear and possible jaw action of Scelidosaurus harrisonii Owen and a review of feeding mechanisms in other thyreophoran dinosaurs Pp 25 52 in Carpenter K ed The Armored Dinosaurs Bloomington Indiana University Press Hennig E 1936 Ein dentale von Kentrurosaurus aethiopicus Hennig Palaeontographica Supplementbande 309 312 Fastovsky DE Weishampel DB 2005 Stegosauria Hot Plates In Fastovsky DE Weishampel DB eds The Evolution and Extinction of the Dinosaurs 2nd ed Cambridge University Press pp 107 30 ISBN 978 0 521 81172 9 Maidment Susannah Catherine Rose Brassey Charlotte Barrett Paul Michael 2015 10 14 The Postcranial Skeleton of an Exceptionally Complete Individual of the Plated Dinosaur Stegosaurus stenops Dinosauria Thyreophora from the Upper Jurassic Morrison Formation of Wyoming U S A PLOS ONE 10 10 e0138352 Bibcode 2015PLoSO 1038352M doi 10 1371 journal pone 0138352 ISSN 1932 6203 PMC 4605687 PMID 26466098 Galton P M 1982 The postcranial anatomy of stegosaurian dinosaur Kentrosaurus from the Upper Jurassic of Tanzania East Africa a b c Galton P M and P Upchurch 2004 Stegosauria pp 343 362 in D B Weishampel P Dodson and H Osmolska eds The Dinosauria 2nd Edition University of California Press Berkeley CA Norman David 2001 Scelidosaurus the earliest complete dinosaur in The Armored Dinosaurs pp 3 24 Bloomington Indiana University Press ISBN 0 253 33964 2 Peter M Galton 2019 Earliest record of an ankylosaurian dinosaur Ornithischia Thyreophora Dermal armor from Lower Kota Formation Lower Jurassic of India Neues Jahrbuch fur Geologie und Palaontologie Abhandlungen 291 2 205 219 doi 10 1127 njgpa 2019 0800 S2CID 134302379 Sereno P C 1998 A rationale for phylogenetic definitions with application to the higher level taxonomy of Dinosauria Neues Jahrbuch fur Geologie und Palaontologie Abhandlungen 210 41 83 a b c Rauhut Oliver W M Carballido Jose Luis Pol Diego 2020 12 10 First osteological record of a stegosaur Dinosauria Ornithischia from the Upper Jurassic of South America Journal of Vertebrate Paleontology 40 6 e1862133 doi 10 1080 02724634 2020 1862133 ISSN 0272 4634 S2CID 234161169 a b Norman D B 1985 The Illustrated Encyclopedia of Dinosaurs Salamander Books Ltd London Hui Dai Ning Li Maidment Susannah C R Guangbiao Wei Yuxuan Zhou Xufeng Hu Qingyu Ma Xunqian Wang Haiqian Hu Guangzhao Peng 2022 03 30 New stegosaurs from the Middle Jurassic Lower Member of the Shaximiao Formation of Chongqing China Journal of Vertebrate Paleontology 41 5 e1995737 doi 10 1080 02724634 2021 1995737 ISSN 0272 4634 S2CID 247267743 a b Raven Thomas J Maidment Susannah C R 2017 A new phylogeny of Stegosauria Dinosauria Ornithischia Palaeontology 60 3 401 408 doi 10 1111 pala 12291 hdl 10044 1 45349 S2CID 55613546 Galton P M 1990 Stegosauria in D B Weishampel P Dodson amp H Osmolska eds The Dinosauria University of California Press pp 435 455 Gilmore C W 1914 Osteology of the armored Dinosauria in the United States National Museum with special reference to the genus Stegosaurus United States National Museum Bulletin 81 1 136 doi 10 5962 bhl title 63658 Olshevsky G Ford T L 1993 The Origin and Evolution of the Stegosaurs Kyoryugaku Saizensen 4 64 103 Ulansky R E 2014 Evolution of the stegosaurs Dinosauria Ornithischia Dinologia 35 pp in Russian DOWNLOAD PDF http dinoweb narod ru Ulansky 2014 Stegosaurs evolution pdf Ulansky RE 2014 Natronasaurus longispinus 100 years with another name Dinologia 10 pp In Russian Maidment Susannah C R Raven Thomas J Ouarhache Driss Barrett Paul M 2020 01 01 North Africa s first stegosaur Implications for Gondwanan thyreophoran dinosaur diversity Gondwana Research 77 82 97 Bibcode 2020GondR 77 82M doi 10 1016 j gr 2019 07 007 hdl 10141 622706 ISSN 1342 937X S2CID 202188261 Weishampel DB 1984 Interactions between Mesozoic Plants and Vertebrates Fructifications and seed predation N Jb Geol Palaontol Abhandl 167 224 250 a b Barden H E Maidment S C R 2011 Evidence for sexual dimorphism in the stegosaurian dinosaur Kentrosaurus aethiopicus from the Upper Jurassic of Tanzania Journal of Vertebrate Paleontology 31 3 641 651 doi 10 1080 02724634 2011 557112 S2CID 83521877 Isles T 2009 The socio sexual behaviour of extant archosaurs Implications for understanding dinosaur behaviour PDF Historical Biology 21 3 4 139 214 doi 10 1080 08912960903450505 S2CID 85417112 Ragna Redelstorff Tom R Hubner Anusuya Chinsamy P Martin Sander 2013 Bone histology of the stegosaur Kentrosaurus aethiopicus Ornithischia Thyreophora from the Upper Jurassic of Tanzania The Anatomical Record 296 6 933 952 doi 10 1002 ar 22701 PMID 23613282 S2CID 23433029 Robert Bussert Wolf Dieter Heinrich and Martin Aberhan 2009 The Tendaguru Formation Late Jurassic to Early Cretaceous southern Tanzania definition palaeoenvironments and sequence stratigraphy Fossil Record 12 2 2009 141 174 Gradstein F M Ogg J G Schmitz M D amp Ogg G M 2012 A Geologic Time Scale 2012 Elsevier Aberhan Martin Bussert Robert Heinrich Wolf Dieter Schrank Eckhart Schultka Stephan Sames Benjamin Kriwet Jurgen Kapilima Saidi 2002 Palaeoecology and depositional environments of the Tendaguru Beds Late Jurassic to Early Cretaceous Tanzania Fossil Record 5 1 19 44 doi 10 1002 mmng 20020050103 Mateus Octavio 2006 Late Jurassic dinosaurs from the Morrison Formation USA the Lourinha and Alcobaca formations Portugal and the Tendaguru Beds Tanzania a comparison In Foster J R Lucas S G eds Paleontology and Geology of the Upper Morrison Formation New Mexico Museum of Natural History and Science bulletin 36 pp 223 232 ISSN 1524 4156 a href Template Cite book html title Template Cite book cite book a journal ignored help Weishampel David B et al 2004 Dinosaur distribution Late Jurassic Africa In Weishampel David B Dodson Peter and Osmolska Halszka eds The Dinosauria 2nd Berkeley University of California Press p 552 ISBN 0 520 24209 2 Barrett P M Butler R J Edwards N P amp Milner A R Pterosaur distribution in time and space an atlas p61 107 in Flugsaurier Pterosaur papers in honour of Peter Wellnhofer 2008 Hone D W E and Buffetaut E eds Zitteliana B 28 264pp 1 Rauhut Oliver W M 2011 Theropod dinosaurs from the Late Jurassic of Tendaguru Tanzania Special Papers in Palaeontology 86 195 239 Buffetaut Eric 2012 An early spinosaurid dinosaur from the Late Jurassic of Tendaguru Tanzania and the evolution of the spinosaurid dentition Oryctos 10 1 8 Heinrich Wolf Dieter et al 2001 The German Tanzanian Tendaguru Expedition 2000 Fossil Record 4 1 223 237 doi 10 1002 mmng 20010040113 External links edit nbsp Wikimedia Commons has media related to Kentrosaurus nbsp Wikispecies has information related to Kentrosaurus Stegosauria from Thescelosaurus com Includes details on Kentrosaurus its junior synonyms and other material Retrieved from https en wikipedia org w index php title Kentrosaurus amp oldid 1219458679, wikipedia, wiki, book, books, library,

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