Wikipedia

Spinosaurus

Spinosaurus(/ˌspaɪnəˈsɔːrəs/;lit. 'spine lizard') is agenusofspinosauriddinosaurthat lived in what now isNorth Africaduring theCenomanianstageof theLate Cretaceousperiod,about 100 to 94million years ago.The genus was known first from Egyptian remains discovered in 1912 and described by GermanpalaeontologistErnst Stromerin1915.The original remains were destroyed inWorld War II,but additional material came to light in the early 21st century. It is unclear whether one or two species are represented in the fossils reported in the scientific literature. Thetype speciesisS. aegyptiacusfromEgyptandMorocco.Although a potential second dubious species,S. maroccanus,has been recovered from Morocco, thisdubiousspecies is likely a junior synonym ofS. aegyptiacus.Other possible junior synonyms includeSigilmassasaurusfrom theKem Kem bedsin Morocco andOxalaiafrom theAlcântara FormationinBrazil,though other researchers propose both genera to be distincttaxa.

Spinosaurus
Temporal range:Late Cretaceous(Cenomanian),100–94Ma PossibleAlbianrecords[1][2]
Reconstructed skeleton based on theNeotypespecimen FSAC-KK-11888 in swimming posture,Field Museum of Natural History
Scientific classificationEdit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Clade: Dinosauria
Clade: Saurischia
Clade: Theropoda
Family: †Spinosauridae
Tribe: †Spinosaurini
Genus: †Spinosaurus
Stromer,1915
Type species
†Spinosaurus aegyptiacus
Stromer, 1915
Synonyms

Spinosaurusis the longest known terrestrialcarnivore;other large carnivores comparable toSpinosaurusincludetheropodssuch asTyrannosaurus,GiganotosaurusandCarcharodontosaurus.The most recent study suggests that previous body size estimates are overestimated, and thatS. aegyptiacusreached 14 m (46 ft) in length and 7.4 t (8.2 short tons) in body mass. The skull ofSpinosauruswas long, low, and narrow, similar to that of a moderncrocodilian,and bore straight conical teeth with noserrations.It would have had large, robust forelimbs bearing three-fingered hands, with an enlarged claw on the firstdigit.The distinctiveneural spinesofSpinosaurus,which were long extensions of thevertebrae(or backbones), grew to at least 1.65 m (5.4 ft) long and were likely to have had skin connecting them, forming asail-like structure,although some authors have suggested that the spines were covered in fat and formed a hump.[4]The hip bones ofSpinosauruswere reduced, and the legs were very short in proportion to the body. Its long and narrow tail was deepened by tall, thin neural spines and elongatedchevrons,forming a flexible fin or paddle-like structure.

Spinosaurusis known to have eaten fish and small to medium terrestrial prey as well.[5]Evidence suggests that it wassemiaquatic;how capable it was of swimming has been strongly contested.Spinosaurus's leg bones hadosteosclerosis(high bone density), allowing for betterbuoyancycontrol. Multiple functions have been put forward for the dorsal sail, includingthermoregulationanddisplay;either to intimidate rivals or attract mates. It lived in a humid environment oftidal flatsandmangroveforests alongside many other dinosaurs, as well as fish,crocodylomorphs,lizards, turtles,pterosaurs,andplesiosaurs.

Discovery and naming

Naming of species

Plate I inStromer(1915) showingS. aegyptiacusholotype elements

TwospeciesofSpinosaurushave been named:Spinosaurus aegyptiacus(meaning "Egyptian spine lizard" ) and the disputedSpinosaurus maroccanus(meaning "Moroccan spine lizard" ).[6][7]The first described remains ofSpinosauruswere found and described in the early 20th century. In 1912, Richard Markgraf discovered a partial skeleton of a giant theropod dinosaur in theBahariya Formationof western Egypt. In 1915, German paleontologist Ernst Stromer published an article assigning the specimen to a new genus and species,Spinosaurus aegyptiacus.[8][6]

Fragmentary additional remains from Bahariya, includingvertebraeand hindlimb bones, were designated by Stromer as "Spinosaurus B"in 1934.[9]Stromer considered them different enough to belong to another species, and this has been borne out. With the advantage of more expeditions and material, it appears that they pertain either toCarcharodontosaurus[10]or toSigilmassasaurus.[7]

S. maroccanuswas originally described byDale Russellin 1996 as a new species based on the length of its neck vertebrae. Specifically, Russell claimed that the ratio of the length of thecentrum(body of vertebra) to the height of the posterior articular facet was 1.1 inS. aegyptiacusand 1.5 inS. maroccanus.[7]Later authors have been split on this topic. Some authors note that the length of the vertebrae can vary from individual to individual, that the holotype specimen was destroyed and thus cannot be compared directly with theS. maroccanusspecimen, and that it is unknown which cervical vertebrae theS. maroccanusspecimens represent. Therefore, though some have retained the species as valid without much comment,[11][12][13]most researchers regardS. maroccanusas anomen dubium(dubious name)[14][1][15]or as a junior synonym ofS. aegyptiacus.[10][16]Some studies have referred the holotype and other referred specimens ofS. maroccanus(NMC 50791 and MNHN SAM 124-128) asS.cf.aegyptiacus.[17][18]The specimens previously ascribed as paratypes ofS. maroccanus(NMC 41768 and NMC 50790) are reidentified as indeterminate spinosaurid specimens that are currently not identifiable at the generic level.[19][20]

Specimens

Digital skeletal reconstruction ofSpinosaurus,showing known bones based on theholotype(red), proposedneotype(blue), and referred specimens (yellow), according toSerenoand colleagues

Six main partial specimens ofSpinosaurushave been described. BSP 1912 VIII 19, described by Stromer in 1915 from the Bahariya Formation, was theholotype.[8][6]The material consisted of the following items, most of which were incomplete: right and leftdentariesandsplenialsfrom the lower jaw measuring 75 centimeters (30 in) long; a straight piece of the leftmaxillathat was described but not drawn; 20 teeth; 2cervicalvertebrae; 7 dorsal (trunk) vertebrae; 3 sacral vertebrae; 1caudalvertebra; 4 thoracic ribs; andgastralia.Of the nine neural spines whose heights are given, the longest ( "i," associated with a dorsal vertebra) was 1.65 meters (5.4 ft) in length. Stromer claimed that the specimen was from the early Cenomanian, about 97 million years ago.[8][6]

It was destroyed inWorld War II,specifically "during the night of 24/25 April 1944 in a British bombing raid of Munich" that severely damaged the building housing thePaläontologisches Museum München(Bavarian State Collection of Paleontology). However, detailed drawings and descriptions of the specimen remain. Stromer's son donated Stromer's archives to the Paläontologische Staatssammlung München in 1995, and Smith and colleagues analyzed two photographs of theSpinosaurusholotype specimen BSP 1912 VIII 19 discovered in the archives in 2000. On the basis of a photograph of the lower jaw and a photograph of the entire specimen as mounted, Smith concluded that Stromer's original 1915 drawings were slightly inaccurate.[6]In 2003, Oliver Rauhut suggested that Stromer'sSpinosaurusholotypewas achimera,composed of vertebrae and neural spines from acarcharodontosauridsimilar toAcrocanthosaurusand adentaryfromBaryonyxorSuchomimus.[15]The analysis was rejected in at least one subsequent paper.[14]

NMC 50791, held by theCanadian Museum of Nature,is a mid-cervical vertebra which is 19.5 centimeters (7.7 in) long from theKem Kem BedsofMorocco.It is the holotype ofSpinosaurus maroccanus,as described by Russell in 1996. Other specimens referred toS. maroccanusin the same paper were two other mid-cervical vertebrae (NMC 41768 and NMC 50790), an anterior dentary fragment (NMC 50832), a mid-dentary fragment (NMC 50833), and an anterior dorsalneural arch(NMC 50813). Russell stated that "only general locality information could be provided" for the specimen, and therefore it could be dated only "possibly" to the Albian.[7]

Comparison between the snouts MNSM V4047 and MNHN SAM 124

MNHN SAM 124, housed at theMuséum National d'Histoire Naturelle,is a snout (consisting of partial premaxillae, partial maxillae,vomers,and a dentary fragment). Described by Taquet and Russell in 1998, the specimen is 13.4 to 13.6 centimeters (5.3–5.4 in) in width; no length was stated. The specimen was located in Algeria, and "is of Albian age." Taquet and Russell believed that the specimen, along with a premaxilla fragment (SAM 125), two cervical vertebrae (SAM 126–127), and a dorsal neural arch (SAM 128), belonged toS. maroccanus.[2]Although it was originally ascribed toS. maroccanus,based on their examination of this cranial material, the 2016 study considered the difference between the two species to be not taxonomically significant and either ontogenetic or intraspecific, and thus tentatively assigned the specimen toS. aegyptiacus.[16]The 2017 study considered MNHN SAM 124 to belong to same taxon as MSNM V4047.[21]

BM231 (in the collection of the Office National des Mines, Tunis) was described by Buffetaut and Ouaja in 2002. It consists of a partial anterior dentary 11.5 centimeters (4.5 in) in length from an early Albianstratumof theChenini FormationofTunisia.The dentary fragment, which included fouralveoliand two partial teeth, was "extremely similar" to existing material ofS. aegyptiacus.[1]

UCPC-2 in theUniversity of ChicagoPaleontological Collection consists mainly of two narrow connectednasalswith a fluted (ridged) crest from the region between the eyes. The specimen, which is 18.0 centimeters (7.1 in) long, was located in an early Cenomanian part of the Moroccan Kem Kem Beds in 1996 and described in the scientific literature in 2005 byCristiano Dal Sassoof theCivic Natural History MuseuminMilanand colleagues.[14]

MSNM V4047 (in theMuseo di Storia Naturale di Milano), described by Dal Sasso and colleagues in 2005 asSpinosauruscf.S. aegyptiacus,consists of a snout (premaxillae, partial maxillae, and partial nasals) 98.8 centimeters (38.9 in) long from the Kem Kem Beds.[14]An isolated fish vertebra, tentatively referred toOnchopristis,has been associated with the tooth alveolus of this specimen.[14]Similarly, the dentary fragment ofSpinosaurus aegyptiacus,MPDM 31, is associated with the rostral tooth ofOnchopristis.[22]Like UCPC-2, it is thought to have come from the early Cenomanian. Arden and colleagues in 2018 tentatively assigned this specimen toSigilmassasaurus brevicollisgiven its size.[23]However, this assignment was later rejected by other researchers who considered the uniqueness of this specimen to be based on misinterpretations and poor preservation of another specimen, NHMUK R16665, another snout stored that is stored in theNatural History Museum, London.[3]

FSAC-KK 11888 is a partial subadult skeleton recovered from the Kem Kem beds of North Africa. It was described byIbrahimand colleagues in 2014 and designated as the neotype specimen,[24]though Evers and colleagues rejected the neotype designation for FSAC-KK-11888 in 2015.[19]It includes cervical vertebrae, dorsal vertebrae, neural spines, a complete sacrum, femora, tibiae, pedal phalanges, caudal vertebra, several dorsal ribs, and fragments of the skull.[24]The body proportions of the specimen have been debated, as the hind limbs are disproportionately shorter in the specimen than in previous reconstructions. However, it has been demonstrated by multiple paleontologists that the specimen is not a chimera, and is indeed a specimen ofSpinosaurusthat suggests that the animal had much smaller hind limbs than previously thought.[25][26][27]

Other known specimens consist mainly of very fragmentary remains and scattered teeth. These include:

  • A 1986 paper described prismatic structures intooth enamelfrom twoSpinosaurusteeth from Tunisia.[28]
  • Buffetaut (1989, 1992) referred three specimens from the Institut und Museum für Geologie und Paläontologie of theUniversity of Göttingenin Germany toSpinosaurus:a right maxilla fragment IMGP 969–1, a jaw fragment IMGP 969–2, and a tooth IMGP 969–3.[29][30]These had been found in a Lower Cenomanian or Upper Albian deposit in southeastern Morocco in 1971.[29]
  • Kellner and Mader (1997) described two unserrated spinosaurid teeth from Morocco (LINHM 001 and 002) that were "highly similar" to the teeth of theS. aegyptiacusholotype.[31]
  • Teeth from the Chenini Formation in Tunisia which are "narrow, somewhat rounded in cross-section, and lack the anterior and posterior serrated edges characteristic of theropods and basalarchosaurs"were assigned toSpinosaurusin 2000.[32]
  • Material possibly belonging toSpinosaurusfrom theTurkana GritsofKenyahas been noted in 2004.[33]
  • Teeth from theEchkar FormationofNigerwere tentatively referred toSpinosaurusin 2007.[34]
  • A partial tooth 8 centimeters (3.1 in) long purchased at a fossil trade show, reportedly from the Kem Kem Bed of Morocco and attributed toSpinosaurus maroccanus,showed 1 to 5 millimeters (0.039 to 0.197 in) wide longitudinal striations and micro-structures (irregular ridges) among the striations in a 2010 paper.[13]
  • Isolated teeth attributed toS. aegyptiacusare reported from Algeria in 2015.[35]
  • Pedal ungual (MSNM V6894), cervical vertebra (FSAC-KK-7280) and dorsal vertebra (FSAC-KK-18118) from the Kem Kem beds are referred to juvenile cf.Spinosaurus aegyptiacus.[36][37]

MHNM.KK374, MHNM.KK375, MHNM.KK376, MHNM.KK377, MHNM.KK378 and MSNM V6896 are six isolated quadrates (skull bones) of different sizes that were collected by locals and acquired commercially in the Kem Kem region of southeastern Morocco, provided by François Escuillié and are deposited in the collections of the Muséum d’Histoire Naturelle of Marrakech. Only MHNM.KK376 is assigned toSigilmassasaurus brevicollis,while the other five specimens are assigned toS. aegyptiacus,since the quadrates show two different morphologies, suggesting the existence of two spinosaurines in Morocco.[16]However, a 2020 study on variation withinSpinosaurusconsiders these differences in morphology to be indicative of variation in skull morphology within a single species, as is the case inAllosaurus.[3]

Possible synonyms

Sigilmassasaurus

Some scientists have considered the genusSigilmassasaurusa junior synonym ofSpinosaurus.In Ibrahim and colleagues (2014), the specimens ofSigilmassasauruswas referred toSpinosaurus aegyptiacustogether with "Spinosaurus B" as theneotypeandSpinosaurus maroccanuswas considered as anomen dubiumfollowing the conclusions of the other papers.[14][24][10]A 2015 re-description ofSigilmassasaurusdisputed these conclusions, and considered the genus valid, with inclusion ofS. maroccanusas a synonym ofSigilmassasaurusinstead.[19]This conclusion was further supported in 2018 by Arden and colleagues, who considerSigilmassasaurusto be a distinct genus, though a very close relative ofSpinosaurus,the two unified in the tribe Spinosaurini, coined in the study.[23]

The 2020 study indicates synonymy betweenSpinosaurusandSigilmassasaurus,and considered specimens previously referred toSigilmassasaurusas those ofSpinosaurus.[3]For instance, the referral of an isolated quadrate (specimen MHNM.KK376) toSigilmassasaurus brevicollis,based on its difference from other specimens assigned toSpinosaurus aegyptiacus,[16]was rejected by the 2020 study which noted that these differences in morphology are indicative of variation in skull morphology within a single species.[3]The 2019 study assigned a juvenile specimen FSAC-KK-18122 toSigilmassasaurus brevicollisbased on its identical proportion to BSPG 2011 I 115 which was assigned to the taxon in a 2015 study,[37][19]but this referral was also rejected in a 2020 study based on the fact that the median tubercle and median suture is present in BSPG 2011 I 115 but absent in FSAC-KK-18122, so the presence or absence of such feature should not be used to taxonomically separate isolated spinosaurid remains.[3]

Regardless of the synonymy ofSigilmassasauruswithSpinosaurus,some authors consider the possibility that there could be a second distinct spinosaurid in North Africa during the Cenomanian age.[20]Additionally, in 2024, a complete posterior cervical vertebra (specimen NHMUK PV R 38358) was assigned toSigilmassasaurus brevicollis.[38]

Oxalaia

Since theNational Museum of Brazil firein 2018 engulfed the palace housing the museum,[39]with specimens ofOxalaiapossibly being destroyed,[40]any classification should remain tentative. In a 2020 paper written by Symthet al.in assessing spinosaurine specimens from theKem Kem Groupsuggested theBrazilianspinosaurineOxalaiato be a potential junior synonym ofSpinosaurus aegyptiacus.This was based on looking at the specimens assigned toOxalaia,and the supposed autapomorphies of this taxon to be insignificant and fall within the hypodigm ofSpinosaurus aegyptiacus.If supported by future studies, this would implySpinosaurus aegyptiacushad a wider distribution and supports the faunal exchange between South America and Africa during this time.[3]

However, subsequent studies have rejected the synonymy ofOxalaiawithSpinosaurus aegyptiacusbased on diagnostic features of the holotype (MN 6117-V) and the referred specimen (MN 6119-V). In 2021, Lacerda, Grillo and Romano noted that the anteromedial processes of the holotype maxillae (MN 6117-V) contact medially, a condition not observed in MSNM V4047 which has been referred to as a specimen ofSpinosaurus,and thus adding a new possible diagnostic feature ofOxalaia.They also suggested that the premaxilla ofOxalaiais wider in the posterior portion than that of MSNM V4047, and that the lateral morphology of its rostrum was distinguished from other spinosaurines based on their morphometric analysis.[41]In 2023, Isasmendi and colleagues consideredOxalaiaas a valid taxon based on the examination of its referred maxilla (MN 6119-V) which suggests that the position of its external naris would have been more anteriorly located, a condition similar to that ofIrritatorandbaryonychines,differing from African spinosaurines includingSpinosaurus aegyptiacus.[42]

Description

Size

Size comparison of selected giant theropod dinosaurs,S. aegyptiacusin red

Since its discovery,Spinosaurushas been a contender for the largest theropod dinosaur.[43]BothFriedrich von Huenein 1926[44]andDonald F. Glutin 1982 listed it as among the most massive theropods in their surveys, at 15 m (49 ft) in length and upwards of 6 t (6.6 short tons) in weight.[45]In 1988,Gregory S. Paulalso listed it as the longest theropod at 15 m (49 ft), but gave a lower mass estimate of 4 t (4.4 short tons).[46]

In 2005, Dal Sasso and colleagues assumed thatSpinosaurusand the relatedSuchomimushad the same body proportions in relation to their skull lengths, and thereby calculated thatSpinosauruswas 16 to 18 m (52 to 59 ft) in length and 7 to 9 t (7.7 to 9.9 short tons) in weight.[14]The estimates were criticized because the skull length estimate was uncertain, and (assuming that body mass increases as the cube of body length) scalingSuchomimus,which was 11 m (36 ft) long and 3.8 t (4.2 short tons) in mass, to the range of estimated lengths ofSpinosauruswould produce an estimated body mass of 11.7 to 16.7 t (12.9 to 18.4 short tons).[47]

Estimated size of the largest known, holotype, neotype, and smallest known specimen with a human

François Therrien and Donald Henderson, in a 2007 paper using scaling based on skull length, challenged previous estimates of the size ofSpinosaurus,finding the length too great and the weight too small. Based on estimated skull lengths of 1.5 to 1.75 m (4 ft 11 in to 5 ft 9 in), their estimates include a body length of 12.6 to 14.3 m (41 to 47 ft) and a body mass of 12 to 20.9 t (13.2 to 23.0 short tons). The lower estimates forSpinosauruswould imply that the animal was shorter and lighter thanCarcharodontosaurusandGiganotosaurus.[47]The Therrien and Henderson study has been criticized for the choice of theropods used for comparison (e.g., most of the theropods used to set the initial equations weretyrannosauridsandcarnosaurs,which have a different build than spinosaurids), and for the assumption that theSpinosaurusskull could be as little as 1.5 m (4 ft 11 in) in length.[48][49]

In 2014, Ibrahim and his colleagues suggested thatSpinosaurus aegyptiacuscould reach over 15 m (49 ft) in length.[24]In 2022, however, Paul Sereno and his colleagues suggested thatSpinosaurus aegyptiacusreached a maximum body length of 14 m (46 ft) and a maximum body mass of 7.4 t (8.2 short tons) by constructing a CT-based 3D skeletal model "with the axial column in neutral pose."[50]They argued that the 2D graphical reconstruction of the aquatic hypothesis by Ibrahim and his colleagues in 2020[22]overestimated the presacral column length by 10%, ribcage depth by 25%, and forelimb length by 30% over dimensions based on CT-scanned fossils; these proportional overestimates shift the center of mass anteriorly when translated to a flesh model, and thus the estimate from Ibrahim and his colleagues cannot be considered a reliable body size estimate.[50]

Skull

Annotated diagram of the reconstructed skull
Annotated diagram of the reconstructed skull muscles of Spinosaurus

Its skull had a narrow snout filled with straight conical teeth that lacked serrations. There were six or seven teeth on each side of the very front of the upper jaw, in thepremaxillae,and another twelve in bothmaxillaebehind them. The second and third teeth on each side were noticeably larger than the rest of the teeth in the premaxilla, creating a space between them and the large teeth in the front of the maxilla; large teeth in the lower jaw faced this space. The very tip of the snout holding those few large front teeth was expanded, and a small crest was present in front of the eyes. Using the dimensions of three specimens known as MSNM V4047, UCPC-2, and BSP 1912 VIII 19, and assuming that the postorbital part of the skull of MSNM V4047 had a shape similar to the postorbital part of the skull ofIrritator,Dal Sasso and colleagues (2005) estimated that the skull ofSpinosauruswas 1.75 meters (5.7 ft) long,[14]but more recent estimates suggest a length of 1.6–1.68 meters (5.2–5.5 ft).[16][51]The Dal Sasso and colleagues skull length estimate is questioned because skull shapes can vary across spinosaurid species and because MSNM V4047 may not belong toSpinosaurusitself,[47][23]though recent studies have reconfirmed it as a specimen ofSpinosaurus.[50][3]

Postcranial skeleton

Reconstructed replicas of the holotypevertebrae,National Geographic Museum,Washington, D. C.

As a spinosaurid,Spinosauruswould have had a long, muscular neck, curved in asigmoid,or S-shape. Its shoulders were prominent, and the forelimbs large and stocky, bearing three claweddigitson each hand. The first finger (or "thumb" ) would have been the largest.Spinosaurushad longphalanges(finger bones), and only somewhat recurvedclaws,suggesting that its hands were longer compared to those of other spinosaurids.[24][23][52]

Very tallneural spinesgrowing on the back vertebrae ofSpinosaurusformed the basis of what is usually called the animal's "sail".The lengths of the neural spines reached over 10 times the diameters of thecentra(vertebral bodies) from which they extended.[53][54]The neural spines were slightly longer front to back at the base than higher up, and were unlike the thin rods seen in thepelycosaurfinbacksEdaphosaurusandDimetrodon,contrasting also with the thicker spines in theiguanodontianOuranosaurus.[53]

Life restoration

Spinosaurussails were unusual, although other dinosaurs, namelyOuranosaurus,which lived a few million years earlier in the same general region asSpinosaurus,and theEarly CretaceousSouth American sauropodAmargasaurus,might have developed similar structural adaptations of their vertebrae. The sail may be ananalogof the sail of thePermiansynapsidDimetrodon,which lived before the dinosaurs even appeared, produced byconvergent evolution.[53]

The structure may also have been more hump-like than sail-like, as noted by Stromer in 1915 ( "one might rather think of the existence of a large hump of fat [German:Fettbuckel], to which the [neural spines] gave internal support ")[8]and by Jack Bowman Bailey in 1997.[53]In support of his "buffalo-back" hypothesis, Bailey argued that inSpinosaurus,Ouranosaurus,and other dinosaurs with long neural spines, the spines were relatively shorter and thicker than the spines ofpelycosaurs(which are known to have sails); instead, the dinosaurs' neural spines were similar to the neural spines of extinct hump-backed mammals such asMegaceropsandBison latifrons.[53][55]In 2014,Ibrahimand colleagues instead posited that the spines were covered tightly by skin, similar to acrested chameleon,given their compactness, sharp edges, and likely poorblood flow.[24]

Spinosaurushad a significantly smallerpelvis(hip bone) than that of other giant theropods, with the surface area of theilium(main body of the pelvis) half that of most members of the clade. The hind limbs were short, at just over 25 percent of the total body length, with thetibia(calf bone) being longer than thefemur(thigh bone). Unlike in other theropods, the hallux (or fourth toe) ofSpinosaurustouched the ground, and the phalanges of the toe bones were unusually long and well-built. At their ends were shallow claws that had flat bottoms. This type of footmorphologyis also seen inshorebirds,indicating thatSpinosaurus's feet evolved for walking across unstable substrate and that they may have beenwebbed.[24]

From thecaudalvertebrae of the tail projected significantly elongated, thin neural spines, akin to the condition observed in some other spinosaurids,[23]though to a more extreme degree. Coupled with the also elongatedchevronbones on the underside of the caudals, this resulted in a deep and narrow tail with a paddle or fin-like shape, comparable to the tails ofnewtsandcrocodilians.[56]

Classification

Specimen MNBH EGA1 which may belong toSpinosaurus

Spinosaurusgives its name to the dinosaurfamilySpinosauridae,which includes two subfamilies:Baryonychinaeand Spinosaurinae. Baryonychinae includesBaryonyxfrom southernEnglandandSuchomimusfromNigerin centralAfrica.Spinosaurinae includesSpinosaurus,Siamosaurus,Ichthyovenator,Irritator,Angaturama(which may be synonymous withIrritator),SigilmassasaurusandOxalaia(both of which may be synonymous withSpinosaurus).[14][23]The spinosaurines share unserrated straight teeth that are widely spaced (e.g., 12 on one side of the maxilla), as opposed to the baryonychines, which have serrated curved teeth that are numerous (e.g., 30 on one side of the maxilla).[14][11]

An analysis of Spinosauridae by Arden and colleagues (2018) named the clade Spinosaurini and defined it as all spinosaurids closer toSpinosaurus aegyptiacusthan toIrritator challengeriorOxalaia quilombensis;it also foundSiamosaurus suteethorniandIcthyovenator laosensisto be members of Spinosaurinae.[23]

Phylogeny

The subfamily Spinosaurinae was named by Sereno in 1998, and defined byHoltzand colleagues (2004) as alltaxacloser toSpinosaurus aegyptiacusthan toBaryonyx walkeri.The subfamily Baryonychinae was named byCharig&Milnerin 1986. They erected both the subfamily and the family Baryonychidae for the newly discoveredBaryonyx,before it was referred to Spinosauridae. Their subfamily was defined by Holtz and colleagues in 2004, as the complementary clade of all taxa closer toBaryonyx walkerithan toSpinosaurus aegyptiacus.Examinations by Marcos Sales, Cesar Schultz, and colleagues (2017) indicate that the South American spinosauridsAngaturama,Irritator,andOxalaiawere intermediate between Baronychinae and Spinosaurinae based on their craniodental features and cladistic analysis. This indicates that Baryonychinae may in fact be non-monophyletic. Theircladogramcan be seen below.[21]

Restoration of variousspinosauridsthat did not live in the same time or space
Spinosauridae

The cladogram below depicts the findings of Arden and colleagues (2018):[23]

Spinosauridae

Paleobiology

Function of neural spines

1915 illustration ofS. aegyptiacusdorsal vertebrae

The function of the dinosaur's sail or hump is uncertain; scientists have proposed severalhypothesesincluding heat regulation and display. In addition, such a prominent feature on its back could make it appear even larger than it was, intimidating other animals.[53]

The structure may have been used forthermoregulation.If the structure contained abundant blood vessels, the animal could have used the sail's large surface area to absorb heat. This would imply that the animal was only partly warm-blooded at best and lived in climates where night-time temperatures were cool or low and the sky usually not cloudy. It is also possible that the structure was used to radiate excess heat from the body, rather than to collect it. Large animals, due to the relatively small ratio of surface area of their body compared to the overall volume (Haldane's principle), face far greater problems of dissipating excess heat at higher temperatures than gaining it at lower. Sails of large dinosaurs added considerably to the skin area of their bodies, with minimum increase of volume. Furthermore, if the sail was turned away from the sun, or positioned at a 90 degree angle towards a cooling wind, the animal would quite effectively cool itself in the warm climate of Cretaceous Africa.[57]However, Bailey (1997) was of the opinion that a sail could have absorbed more heat than it radiated. Bailey proposed instead thatSpinosaurusand other dinosaurs with long neural spines had fatty humps on their backs for energy storage, insulation, and shielding from heat.[53]

Many elaborate body structures of modern-day animals serve to attract members of the opposite sex during mating. It is possible that the sail ofSpinosauruswas used for courtship, in a way similar to apeacock's tail. Stromer speculated that the size of the neural spines may have differed between males and females.[8]

Reconstructed skeleton inNational Geographic Museum

Gimsa and colleagues (2015) suggest that the dorsal sail ofSpinosauruswas analogous to the dorsal fins ofsailfishand served a hydrodynamic purpose.[58]Gimsa and others point out that more basal, long-legged spinosaurids had otherwise round or crescent-shaped dorsal sails, whereas inSpinosaurus,the dorsal neural spines formed a shape that was roughly rectangular, similar in shape to the dorsal fins of sailfish. They therefore argue thatSpinosaurusused its dorsal neural sail in the same manner as sailfish, and that it also employed its long narrow tail to stun prey like a modernthresher shark.Sailfish employ their dorsal fins for herding schools of fish into a "bait ball"where they cooperate to trap the fish into a certain area where the sailfish can snatch the fish with their bills. The sail could have possibly reduced yaw rotation by counteracting the lateral force in the direction opposite to the slash as suggested by Gimsa and colleagues (2015).[58]

Spinosaurusanatomy exhibits another feature that may have a modern analogy: its long tail resembled that of the thresher shark, employed to slap the water to herd and stun shoals of fish before devouring them (Oliver and colleagues, 2013). The strategies that sailfish and thresher sharks employ against shoaling fish are more effective when the shoal is first concentrated into a ‘bait ball’ (Helfman, Collette & Facey, 1997; Oliver and colleagues, 2013; Domenici and colleagues, 2014). Since this is difficult for individual predators to achieve, they cooperate in this effort. When herding a shoal of fish or squid, sailfish also raise their sails to make themselves appear larger. When they slash or wipe their bills through shoaling fish by turning their heads, their dorsal sail and fins are outstretched to stabilize their bodies hydrodynamically (Lauder & Drucker, 2004). Domenici and colleagues (2014) postulate that these fin extensions enhance the accuracy of tapping and slashing. The sail can reduce yaw rotation by counteracting the lateral force in the direction opposite to the slash. This means that prey is less likely to recognize the massive trunk as being part of an approaching predator (Marras and colleagues, 2015; Webb & Weihs 2015).[58]

Spinosaurusexhibited the anatomical features required to combine all three hunting strategies: a sail for herding prey more efficiently, as well as flexible tail and neck to slap the water for stunning, injuring or killing prey. The submerged dorsal sail would have provided a strong centreboard-like counterforce for powerful sidewards movements of the strong neck and long tail, as performed by sailfish (Domenici and colleagues, 2014) or thresher sharks (Oliver and colleagues, 2013). While smaller dorsal sails or fins make the dorsal water volume better accessible for slashing, it can be speculated that their smaller stabilization effect makes lateral slashing less efficient (e.g. for thresher sharks). Forming a hydrodynamic fulcrum and hydrodynamically stabilizing the trunk along the dorsoventral axis,Spinosaurus’ sail would also have compensated for the inertia of the lateral neck by tail movements and vice versa not only for predation but also for accelerated swimming. This behavior might also have been one reason forSpinosaurus’ muscular chest and neck reported by Ibrahim and colleagues (2014).[58]

Diet and feeding

Tooth from Morocco in various views
CT scan of possibleSpinosaurussnout NHMUK 16665

It is unclear whetherSpinosauruswas primarily a terrestrial predator or apiscivore,as indicated by its elongated jaws, conical teeth and raised nostrils. The hypothesis of spinosaurs as specialized fish eaters has been suggested before by A. J. Charig and A. C. Milner forBaryonyx.They base this on the anatomical similarity with crocodilians and the presence of digestive acid-etched fish scales in the rib cage of thetype specimen.[59]Large fish are known from the faunas containing other spinosaurids, including theMawsonia,in the mid-Cretaceous of northern Africa and Brazil. Direct evidence for spinosaur diet comes from related European and South American taxa.Baryonyxwas found with fish scales and bones from juvenileIguanodonin its stomach, while a tooth embedded in a South Americanpterosaurbone suggests that spinosaurs occasionally preyed on pterosaurs,[60]butSpinosauruswas likely to have been a generalized and opportunistic predator, possibly a Cretaceous equivalent of largegrizzly bears,being biased toward fishing, though it undoubtedlyscavengedand took many kinds of small or medium-sized prey.[46]

S. aegyptiacushead based on the 2005 reconstruction byCristiano Dal Sasso

In 2009, Dal Sasso and colleagues. reported the results ofX-ray computed tomographyof the MSNM V4047 snout. As theforaminaon the outside all communicated with a space on the inside of the snout, the authors speculated thatSpinosaurushadpressure receptorsinside the space that allowed it to hold its snout at the surface of the water to detect swimming prey species without seeing them.[61]A 2013 study by Andrew R. Cuff and Emily J. Rayfield concluded that bio-mechanical data suggests thatSpinosauruswas not anobligatepiscivore and that its diet was more closely associated with each individual's size. The characteristicrostralmorphology ofSpinosaurusallowed its jaws to resist bending in the vertical direction, but its jaws were poorly adapted with respect to resisting lateral bending compared to other members of this group (Baryonyx) and modern alligators. This suggests thatSpinosauruspreyed more regularly on fish than it did on land animals, although considered predators of the former too.[62]In 2022, Sakamoto estimated thatSpinosaurushad an anterior bite force of 4,829 newtons and a posterior bite force of 11,936 newtons. Based on this estimate, he asserted that the jaws ofSpinosaurusare adapted for generating relatively faster shutting speeds with less muscle input force, indicating that the animal likely killed its prey with fast-snapping jaws rather than slow-crushing bites, a trait commonly observed in animals which have a semi-aquatic feeding habit.[63]

A 2024 paper suggests that Spinosaurus and other spinosaurines in addition to fish also preyed upon small to medium sized terrestrial vertabrates.[5]

Aquatic habits

Restoration ofSpinosaurusattacking thesawskateOnchopristis

A 2010isotope analysisby Romain Amiot and colleagues found thatoxygen isotoperatios of spinosaurid teeth, including teeth ofSpinosaurus,indicatesemiaquaticlifestyles. Isotope ratios from tooth enamel and from other parts ofSpinosaurus(found in Morocco and Tunisia) and of other predators from the same area such asCarcharodontosauruswere compared with isotopic compositions from contemporaneous theropods, turtles, and crocodilians. The study found thatSpinosaurusteeth from five of six sampled localities had oxygen isotope ratios closer to those of turtles and crocodilians when compared with other theropod teeth from the same localities. The authors postulated thatSpinosaurusswitched between terrestrial and aquatic habitats to compete for food with large crocodilians and other large theropods respectively.[64]A 2018 study by Donald Henderson, however, refutes the claim thatSpinosauruswas semiaquatic. By studying the buoyancy in lungs of crocodilians and comparing it to the lung placement inSpinosaurus,it was discovered thatSpinosauruscould not sink or dive below the water surface. It was also capable of keeping its entire head above the water surface while floating, much like other non-aquatic theropods. Furthermore, the study found thatSpinosaurushad to continually paddle its hind legs to prevent itself from tipping over onto its side, something that extant semiaquatic animals do not need to perform. Henderson therefore theorized thatSpinosaurusprobably did not hunt completely submerged in water as previously hypothesized, but instead would have spent much of its time on land or in shallow water.[65][66]

Recent studies of the tail vertebrae ofSpinosaurusrefute Henderson's proposal thatSpinosaurusmainly inhabited areas of land near and in shallow water and was too buoyant to submerge. Studies of the tail, thanks to fossils recovered and analyzed by Ibrahim, Pierce, Lauder, and Sereno and colleagues in 2018 indicate thatSpinosaurushad a keeled tail that was well adapted to propelling the animal through water. The elongated neural spines and chevrons, which run to the end of the tail on both dorsal and ventral sides, indicate thatSpinosauruswas able to swim in a similar manner to modern crocodilians. Through experimentation by Lauder and Pierce, the tail ofSpinosauruswas found to have eight times as much forward thrust as the tails of terrestrial theropods likeCoelophysisandAllosaurus,as well as being twice as efficient at achieving forward thrust. The discovery indicates thatSpinosaurusmay have had a lifestyle comparable to modern alligators and crocodiles, remaining in water for long periods of time while hunting.[56]

Flesh model ofS. aegyptiacusused for a 2022buoyancystudy

David Hone andThomas Holtzpublished a paper in 2021 in which they argue that the anatomy ofSpinosaurusis more consistent with a shoreline generalist lifestyle rather than an active aquatic pursuit predator as suggested by Ibrahim.[67]They highlight the positioning of the nostrils and orbits as one reason why a crocodile-like lifestyle is unlikely: they are ventrally positioned in such a way that the whole head would have to be lifted inefficiently out of the water in order to breathe. Additionally, they argue that the general body shape ofSpinosaurusis poorly adapted for this lifestyle, drawing on the amount of water drag and aquatic instability[66]from the sail, as well as the rigid trunk and seemingly scarcely-muscled tail. Animals like crocodilians require a flexible body in order to move through the water and make sharp turns when chasing prey, and this is directly contradicted by Hone and Holtz's findings.

A 2022 study by Fabbriet al.,made comparisons ofSpinosaurus'bone structure and compared it to that ofBaryonyxandSuchomimus.The study revealed thatSpinosaurusandBaryonyxhad dense bones, which allowed them to dive and pursue prey underwater. Compared to these,Suchomimushad more hollow bones, suggesting it preferred to hunt in shallow water. These findings also suggest that various spinosaurid genera were more ecologically disparate than previously believed, as some were better suited to hunting in subaqueous environments than other, closely related genera.[68][69][70]

In the same year, contradicting the study by Fabbri and colleagues, Sereno and his colleagues suggested thatSpinosauruswas wholly bipedal on land and an unstable, slow moving surface swimmer in deep water. Their results, taken from reconstructing a CT model of the skeleton, and then adding internal air and muscles. Their results, coupled with fossils fromSpinosaurusthat showed it also lived further inland along rivers and lakes, suggest it was a semi-aquatic, ambush piscivore that preferred waterside environments both along the coasts and further inland along rivers and lakes. Simultaneously, they suggested that the large tail fin was probably utilized more for display than swimming, as tails in living animals have the same function when they possess comparably tall neural spines.[50]

A 2024 paper by Myrhvoldet al.also contends that Spinosaurus and Baryonyx were diving persuit predators. Instead they also argue thatSpinosaurusandBaryonyxhunted more likeheronsinstead of diving after prey.[71]Another paper in the same year analyzed the linear measurements of the skull ofSpinosaurus,and concluded that the skull morphology and hunting method ofSpinosauruswould likely be the most similar to those of wading birds likeherons,though the authors noted that they're uncertain how beneficial the skull would have been for the diving pursuit predation method.[72]

Locomotion and posture

Reconstructed skeleton with traditional, long-legged posture

Although traditionally depicted in the scientific community as abiped,Spinosauruswas occasionally depicted in the mid-20th century as an obligate quadruped akin toDimetrodon.[73]Starting in the mid-1970s, it was hypothesizedSpinosauruswas at least an occasionalquadruped,[45][57]bolstered by the discovery ofBaryonyx,a relative with robust arms.[74]Because of the mass of the hypothesized fatty dorsal humps ofSpinosaurus,Bailey (1997) was open to the possibility of a quadrupedal posture,[53]leading to new restorations of it as such.[74]Theropods, including spinosaurids, could not pronate their hands (rotate the forearm so the palm faced the ground),[75]but a resting position on the side of the hand was possible, as shown by fossil prints from an Early Jurassic theropod.[76]The hypothesis thatSpinosaurushad a typical quadrupedal gait since fell out of favor, however it was still believed that spinosaurids may have crouched in a quadrupedal posture, due to biological and physiological constraints.[59][75]

Reconstructed foot, note straight claws and largehallux

The possibility of a quadrupedalSpinosauruswas revived by a 2014 paper byIbrahimand colleagues that described new material of the animal. The paper found that the hind limbs ofSpinosauruswere much shorter than previously believed, and that its center of mass was located in the midpoint of thetorsoregion, as opposed to near the hip as in typical bipedal theropods. It was therefore proposed thatSpinosauruswas poorly adapted for bipedal terrestrial locomotion, and must have been an obligate quadruped on land. The reconstruction used in the study was an extrapolation based on different sized individuals, scaled to what were assumed to be the correct proportions.[24]Paleontologist John Hutchinson of theRoyal Veterinary Collegeof the University of London has expressed skepticism to the new reconstruction, and cautioned that using different specimens can result in inaccurate chimaeras.[77]Scott Hartman also expressed criticism because he believed the legs and the pelvis were inaccurately scaled (27% too short) and didn't match the published lengths.[78]However,Mark Wittonexpressed agreement with the proportions reported in the paper.[79]In their 2015 re-description ofSigilmassasaurus,Evers and colleagues argued thatSigilmassasauruswas in fact a distinct genus fromSpinosaurus,and therefore doubted whether the material assigned toSpinosaurusby Ibrahimet al.should be assigned toSpinosaurusorSigilmassasaurus.[19]In 2018, an analysis by Henderson found thatSpinosaurusprobably was competent at bipedal terrestrial locomotion; the center of mass was instead found to be close to the hips, allowingSpinosaurusto stand upright like other bipedal theropods.[66]

A 2024 article co-authored by Sereno stated that the previous calculations by Sereno that were used to argue quadrupedality forSpinosaurushad erroneously shifted the center of mass in front of the hips. They instead suggested that the dinosaur fit the criteria of being a graviportal (or slow-moving) biped.[80][81]

Ontogeny

An ungual phalanx measuring 21 millimeters (0.83 in) belonging to a very young juvenile cf.S. aegyptiacusindicates that the theropod developed its semiaquatic adaptations at a very young age or at birth and maintained them throughout its life. The specimen, found in 1999 and described by Simone Maganuco and Cristiano Dal Sasso and colleagues, is believed to have come from an animal measuring 1.78 meters (5.8 ft) (assuming it resembled a smaller version of the adult), making it the smallest specimen ofSpinosauruscurrently known.[36]

Palaeopathology

A cf.Spinosaurussp. tooth from the Ifezouane Formation displays enhanced lingual curvature to the tooth's crown, the development of three deep grooves extending from crown root junction in the direction of the crown's apex, an attenuated carina that does not extend apically nor to the base of the tooth, and a wear facet at the tip.[82]

Paleoenvironment

Restoration ofSpinosauruswith contemporaneous animals of theBahariya Formation

The environment inhabited bySpinosaurusis only partially understood, and covers a great deal of what is now northern Africa. The region of AfricaSpinosaurusis preserved in dates from 112 to 93.5 million years ago.[83][6][84]A potential specimen tentatively referred to as cf.Spinosaurushas been found in theCampanianQuseir Formationof Egypt, but no detailed description of the specimen was provided.[85][86]A 1996 study concluded from Moroccan fossils thatSpinosaurus,Carcharodontosaurus,andDeltadromeus"ranged across north Africa during the late Cretaceous (Cenomanian)."[87]ThoseSpinosaurusthat lived in the Bahariya Formation of what is nowEgyptmay have contended with shoreline conditions ontidal flatsand channels, living inmangroveforests alongside similarly large dinosaurian predatorsBahariasaurusandCarcharodontosaurus,thetitanosaursauropodsParalititanandAegyptosaurus,crocodylomorphs,bony and cartilaginous fish, turtles, lizards, andplesiosaurs.[88]In the dry season it might have resorted to preying onpterosaurs.[89]This situation resembles that in the LateJurassicMorrison FormationofNorth America,which boasts up to five theropod genera over 1 metric ton (1.1 short tons) in weight, as well as several smaller genera (Henderson, 1998; Holtzand colleagues,2004). Differences in head shape and body size among the large North African theropods may have been enough to allownichepartitioning as seen among the many different predator species found today in theAfricansavanna(Farlow & Pianka, 2002).

Sculpture based on the 2014 reconstruction, prior to the discovery of the paddle-like tail,Museum of Natural Science,Barcelona

Spinosaurusappeared in the 2001 filmJurassic Park III,replacingTyrannosaurusas the main antagonist.[90]The film's consulting paleontologistJohn R. Hornerwas quoted as saying, "If we base the ferocious factor on the length of the animal, there was nothing that ever lived on this planet that could match this creature [Spinosaurus]. Also my hypothesis is that T-rex was actually a scavenger rather than a killer.Spinosauruswas really the predatory animal. "[91]He has since retracted the statement aboutT. rexbeing a scavenger. In the film,Spinosauruswas portrayed as larger and more powerful thanTyrannosaurus:in a scene depicting a battle between the two resurrected predators,Spinosaurusemerges victorious by snapping theTyrannosaurus'neck.[92]In the fourth film,Jurassic World,there is a nod to this fight where theT. rexsmashes through the skeleton of aSpinosaurusin the climactic fight near the end of the film.[93]The Spinosaurus would appear in many Jurassic Park games most notablyJurassic World Evolution,and itssequel.The sameSpinosaurusfrom the third film returns in the fourth, and fifth season ofJurassic World Camp Cretaceous,this time battling twoT. rex.[94][95]

Spinosaurushas long been depicted in popular books about dinosaurs, although only recently has there been enough information about spinosaurids for an accurate depiction. After an influential 1955 skeletal reconstruction by Lapparent and Lavocat[96]based on a 1936 diagram by Stromer,[97]it has been treated as a generalized upright theropod, with a skull similar to that of other large theropods and a sail on its back, even having four-fingered hands.[74]

In addition to films, action figures, video games, and books,Spinosaurushas been depicted onpostage stampsfrom countries such asAngola,The Gambia,andTanzania.[98][99]

See also

References

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Further reading

  • Glut, D.F. "In search ofSpinosaurus."In:Jurassic classics: a collection of saurian essays and Mesozoic musings,pp. 77–85. Jefferson, NC: McFarland, 2001.ISBN0-7864-0961-4.
  • Nothdurft, W.; and Smith, J.The Lost Dinosaurs of Egypt.New York: Random House, 2002.ISBN0-375-50795-7.
  • A Tribute to Ernst Stromer: Hundred Years of the Discovery ofSpinosaurus aegyptiacus:Saubhik Ghosh
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