Saber-toothed predator

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From top and from left to right, Inostrancevia, Hoplophoneus, Barbourofelis, Smilodon, Machaeroides and Thylacosmilus

A saber-tooth (alternatively spelled sabre-tooth) is any member of various extinct groups of

saber-shaped canine teeth which protruded from the mouth when closed. Saber-toothed mammals have been found almost worldwide from the Eocene epoch to the end of the Pleistocene epoch (42 million years ago – 11,000 years ago).[1][2][3]

One of the best-known genera is the

creodont" genera Machaeroides and Apataelurus; and two extinct lineages of metatherian mammals, the thylacosmilids of Sparassodonta, and deltatheroideans, which are more closely related to marsupials. In this regard, these saber-toothed mammals can be viewed as examples of convergent evolution.[5] This convergence is remarkable due not only to the development of elongated canines, but also a suite of other characteristics, such as a wide gape and bulky forelimbs, which is so consistent that it has been termed the "saber-tooth suite."[6]

Of the feliform lineages, the family Nimravidae is the oldest, entering the landscape around 42 mya and becoming extinct by 7.2 mya. Barbourofelidae entered around 16.9 mya and were extinct by 9 mya. These two would have shared some habitats.

Morphology

The different groups of saber-toothed predators evolved their saber-toothed characteristics entirely independently. They are most known for having maxillary canines which extended down from the mouth when the mouth was closed. Saber-toothed cats were generally more robust than today's cats and were quite bear-like in build. They are believed to have been excellent hunters, taking animals such as sloths, mammoths, and other large prey. Evidence from the numbers found at the La Brea Tar Pits suggests that Smilodon, like modern lions, was a social carnivore.[7]

The first saber-tooths to appear were non-mammalian

synapsids, such as the gorgonopsids; they were one of the first groups of animals within Synapsida to experience the specialization of saber teeth, and many had long canines. Some had two pairs of upper canines with two jutting down from each side, but most had one pair of upper extreme canines. Because of their primitiveness, they are extremely easy to tell from machairodonts. Several defining characteristics are a lack of a coronoid process, many sharp "premolars" more akin to pegs than scissors, and very long skulls. Despite their large canines, however, most gorgonopsians probably lacked the other specializations found in true saber-toothed predator ecomorphs.[8] Two gorgonopsians, Smilesaurus and Inostrancevia
, had exceptionally large canines and may have been closer functional analogues to later sabertooths.

The second appearance is in Deltatheroida, a lineage of Cretaceous metatherians. At least one genus, Lotheridium, possessed long canines, and given both the predatory habits of the clade as well as the generally incomplete material, this may have been a more widespread adaptation.[9]

The third appearance of long canines is Thylacosmilus, which is the most distinctive of the saber-tooth mammals and is also easy to tell apart. It differs from machairodonts in possessing a very prominent flange and a tooth that is triangular in cross section. The root of the canines is more prominent than in machairodonts and a true sagittal crest is absent.

The fourth instance of saber-teeth is from the clade Oxyaenidae. The small and slender Machaeroides bore canines that were thinner than in the average machairodont. Its muzzle was longer and narrower.

The fifth saber-tooth appearance is the ancient feliform (carnivoran) family Nimravidae. Both groups have short skulls with tall sagittal crests, and their general skull shape is very similar. Some have distinctive flanges, and some have none at all, so this confuses the matter further. Machairodonts were almost always bigger, though, and their canines were longer and more stout for the most part, but exceptions do appear.

The sixth appearance is the barbourofelids. These feliform carnivorans are very closely related to actual cats. The best-known barbourofelid is the eponymous Barbourofelis, which differs from most machairodonts by having a much heavier and more stout mandible, smaller orbits, massive and almost knobby flanges, and canines that are farther back. The average machairodont had well-developed incisors, but barbourofelids' were more extreme.

The seventh and last saber-toothed group to evolve were the machairodonts themselves.

Diet

Reconstruction of a Smilodon

The evolution of enlarged canines in Tertiary carnivores was a result of large mammals being the source of prey for saber-toothed predators. The development of the saber-toothed condition appears to represent a shift in function and killing behavior, rather than one in predator-prey relations. Many hypotheses exist concerning saber-tooth killing methods, some of which include attacking soft tissue such as the belly and throat, where biting deep was essential to generate killing blows. The elongated teeth also aided with strikes reaching major blood vessels in these large mammals. However, the precise functional advantage of the saber-tooth's bite, particularly in relation to prey size, is a mystery. A new point-to-point bite model is introduced in the article by Andersson et al., showing that for saber-tooth cats, the depth of the killing bite decreases dramatically with increasing prey size.[10] The extended gape of saber-toothed cats results in a considerable increase in bite depth when biting into prey with a radius of less than 10 cm. For the saber-tooth, this size-reversed functional advantage suggests predation on species within a similar size range to those attacked by present-day carnivorans, rather than "megaherbivores" as previously believed.

A disputing view of the cat's hunting technique and ability is presented by

C. K. Brain in The Hunters or the Hunted?, in which he attributes the cat's prey-killing abilities to its large neck muscles rather than its jaws.[11] Large cats use both the upper and lower jaw to bite down and bring down the prey. The strong bite of the jaw is accredited to the strong temporalis muscle that attach from the skull to the coronoid process of the jaw. The larger the coronoid process, the larger the muscle that attaches there, so the stronger the bite. As C.K. Brain points out, the saber-toothed cats had a greatly reduced coronoid process and therefore a disadvantageously weak bite. The cat did, however, have an enlarged mastoid process, a muscle attachment at the base of the skull, which attaches to neck muscles. According to C.K. Brain, the saber-tooth would use a "downward thrust of the head, powered by the neck muscles" to drive the large upper canines into the prey. This technique was "more efficient than those of true cats".[11]

Biology

The similarity in all these unrelated families involves the convergent evolution of the saber-like canines as a hunting adaptation. Meehan et al.

co-evolution with large prey animals, led the saber-toothed cats of each time period to extinction. As per Van Valkenburgh, the adaptations that made saber-toothed cats successful also made the creatures vulnerable to extinction. In her example, trends toward an increase in size, along with greater specialization, acted as a "macro-evolutionary ratchet": when large prey became scarce or extinct, these creatures would be unable to adapt to smaller prey or consume other sources of food, and would be unable to reduce their size so as to need less food.[13]

More recently, it has been suggested that Thylacosmilus differed radically from its placental counterparts in possessing differently shaped canines and lacking incisors. This suggests that it was not ecologically analogous to other saber-teeth and possibly an entrail specialist.[14] Another study has found that other saber toothed species similarly had diverse lifestyles and that superficial anatomical similarities obscure them.[15]

Phylogeny of feliform saber-tooths

The following cladogram shows the relationships between the feliform saber-tooths, including the Nimravidae, Barbourofelidae and Machairodontinae.[16][17] Saber-toothed groups are marked with background colors.

Nimravidae
Eusmilus

Eusmilus olsontau

Eusmilus villebramarensis

Eusmilus cerebralis

Hoplophoneus dakotensis

Hoplophoneus primaevus

Hoplophoneus occidentalis

Nanosmilus kurténi

Eofelis edwardsii

Nimravus brachyops

("false saber‑toothed cats")

Haplogale media

Barbourofelidae
Barbourofelis

Barbourofelis morrisi

Barbourofelis fricki

Barbourofelis loveorum

Barbourofelis whitfordi

Sansanosmilus vallesiensis

("false saber‑toothed cats")
Felidae

Proailurus lemanensis

"Pseudaelurus"

Pseudaelurus quadridentatus

Machairodontinae
Machairodus

Machairodus aphanistus

Machairodus horribilis

Amphimachairodus

Amphimachairodus kurteni

Amphimachairodus kabir

Amphimachairodus giganteus

Lokotunjailurus emageritus

Homotheriini
Dinobastis

Dinobastis serus

Dinobastis venezuelensis

Xenosmilus hodsonae

Homotherium latidens

(scimitar‑toothed cats)
Metailurini
Dinofelis

Dinofelis petteri

Dinofelis paleoonca

Dinofelis cristata

Metailurus

Metailurus major

Metailurus parvulus

Promegantereon ogygia

Smilodontini
Megantereon

Megantereon whitei

Megantereon cultridens

Megantereon ekidoit

Megantereon hesperus

Smilodon

Smilodon fatalis

Smilodon populator

Smilodon gracilis

(dirk‑toothed cats)
(saber‑toothed cats)
Styriofelis

Styriofelis lorteti

Styriofelis turnauensis

Felinae sensu lato (includes all extant pantherines and felines)

(grade)

Saber-tooth genera

Genus Name Species Appeared
(
BP
)
Died out
(Ma BP)
Regions Canine size
Smilodon 3–5 2.5 0.01 North and South America 17–30 cm
Hoplophoneus 5 33.7 23.8 North and South America
Eusmilus 3 30.5 28 Eurasia and North America
Dinictis 4 40 25 North America
Dinaelurus 1 ? ? North America
Dinailurictis 1 ? ? ?
Eofelis 2 ? ? ?
Nimravidus (Nimravides) 2 ? ? ?
Nimravus (Nimravinus) 6 33.5 20 Europe and North America
Nimraviscus
1 ? ? ?
Pogonodon 2 15 6 Europe and North America
Quercylurus 1 ? ? ?
Archaelurus
1 ? ? ?
Aelurogale
(Ailurictis)
1 ? ? ?
Ictidailurus
1 ? ? ?
Albanosmilus 3 18 3 Eurasia and Africa
Afrosmilus 1 25 10 Africa
Barbourofelis 7 15 3 Eurasia and Africa
Ginsburgsmilus 1 23 10 Africa
Prosansanosmilus 2 18 5 Eurasia and Africa
Sansanosmilus 3 12 3 Eurasia and Africa
Syrtosmilus 1 23 8 Africa
Vampyrictis 1 15 3 Eurasia and Africa
Vishnusmilus 1 ? ? ?
Homotherium 10 3 0.01 Eurasia, Africa, North America, South America
Thylacosmilus 2 10 1.8 South America over 30 cm
Metailurus 9 15 8 Eurasia
Adelphailurus 1 23 5 North America
Paramachairodus
3 20–15 9 Europe
Machairodus 18 15 2 Eurasia, Africa, North America
Miomachairodus 1 13.65 5.33 Eurasia, Africa, North America
Amphimachairodus 4 late Miocene ? Eurasia, North Africa, North America
Hemimachairodus 1 Pleistocene Pleistocene Java
Lokotunjailurus 1 late Miocene late Miocene Africa
Megantereon 8 3 0.5 Eurasia, Africa, North America
Dinofelis 6 5 1.5 Eurasia, Africa, North America
Therailurus
1 5 2 Eurasia, Africa, North America
Pontosmilus
4 20 9 Eurasia
Proailurus 2 30 20 Europe and North America
Pseudaelurus 1 20 10 Europe and North America
Xenosmilus 1 1.7 1 North America
Stenailurus 1 ? ? ?
Epimachairodus 1 ? ? ?
Hemimachairodus 1 ? ? ?
Ischyrosmilus
1 ? ? ?

Saber-tooth taxonomy

All saber-toothed mammals lived between 33.7 million and 9,000 years ago, but the evolutionary lines that led to the various saber-tooth genera started to diverge much earlier. It is thus a

polyphyletic
grouping.

The lineage that led to

nimravids
, before the diversification of the truly feline saber-tooths.

References

  1. ^ "PaleoBiology Database: Smilodon, basic info". Paleodb.org. Retrieved 2012-09-06.
  2. ^ "PaleoBiology Database: Nimravidae, basic info". Paleodb.org. Retrieved 2012-09-06.
  3. ^ "PaleoBiology Database: Barbourofelidae, basic info". Paleodb.org. Retrieved 2012-09-06.
  4. PMID 26893959
    .
  5. ^ Antón, Mauricio (2013). Sabertooth (Life of the Past). Indiana University Press.
  6. S2CID 86749260
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  11. ^
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  17. ^ Piras P, Maiorino L, Teresi L, Meloro C, Lucci F, Kotsakis T, Raia P (2013) Data from: Bite of the cats: relationships between functional integration and mechanical performance as revealed by mandible geometry. Dryad Digital Repository. https://dx.doi.org/10.5061/dryad.kp8t3

Further reading

External links