Miomachairodus
Encyclopedia
Miomachairodus was an extinct genus
Genus
In biology, a genus is a low-level taxonomic rank used in the biological classification of living and fossil organisms, which is an example of definition by genus and differentia...

 of large saber-toothed cat
Saber-toothed cat
Saber-toothed cat or Sabre-toothed cat refers to the extinct subfamilies of Machairodontinae , Barbourofelidae , and Nimravidae as well as two families related to marsupials that were found worldwide from the Eocene Epoch to the end of the Pleistocene Epoch ,...

s of the subfamily Machairodontinae
Machairodontinae
Machairodontinae is an extinct carnivoran mammal subfamily of Felidae endemic to Asia, Africa, North America, South America, and Europe from the Miocene to Pleistocene living from c. 23 Ma until c...

, that lived in Europe
Europe
Europe is, by convention, one of the world's seven continents. Comprising the westernmost peninsula of Eurasia, Europe is generally 'divided' from Asia to its east by the watershed divides of the Ural and Caucasus Mountains, the Ural River, the Caspian and Black Seas, and the waterways connecting...

, Asia
Asia
Asia is the world's largest and most populous continent, located primarily in the eastern and northern hemispheres. It covers 8.7% of the Earth's total surface area and with approximately 3.879 billion people, it hosts 60% of the world's current human population...

, Africa
Africa
Africa is the world's second largest and second most populous continent, after Asia. At about 30.2 million km² including adjacent islands, it covers 6% of the Earth's total surface area and 20.4% of the total land area...

, and North America
North America
North America is a continent wholly within the Northern Hemisphere and almost wholly within the Western Hemisphere. It is also considered a northern subcontinent of the Americas...

 during most of the Miocene
Miocene
The Miocene is a geological epoch of the Neogene Period and extends from about . The Miocene was named by Sir Charles Lyell. Its name comes from the Greek words and and means "less recent" because it has 18% fewer modern sea invertebrates than the Pliocene. The Miocene follows the Oligocene...

. It survived until the Late Miocene (early Vallesian
Vallesian
The Vallesian age is a period of geologic time within the Miocene used more specifically with European Land Mammal Ages. It precedes the Turolian age and follows the Astaracian age. The Turolian overlaps the Tortonian and Messinian ages....

).

Miomachairodus coexisted with the barbourofelid Barbourofelis
Barbourofelis
Barbourofelis is an extinct genus of large, mostly carnivorous mammals of the family Barbourofelidae endemic to North America, during the Miocene living from 13.6—5.3 Ma and existed for approximately .-Taxonomy:...

in its North American range.

Fossil record

The only adequate collection of Miomachairodus fossils is the Vallesian
Vallesian
The Vallesian age is a period of geologic time within the Miocene used more specifically with European Land Mammal Ages. It precedes the Turolian age and follows the Astaracian age. The Turolian overlaps the Tortonian and Messinian ages....

-age Bahe Formation
Bahe Formation
The Bahe Formation is a Late Miocene formation in Shaanxi, China. Its coordinates are 33.5N and 109.5E. It has "a complex lithology of predominantly orange-yellow conglomerates, sandstones, tan-yellow sandy mudstones, and tan-red mudstones."...

 in Shaanxi
Shaanxi
' is a province in the central part of Mainland China, and it includes portions of the Loess Plateau on the middle reaches of the Yellow River in addition to the Qinling Mountains across the southern part of this province...

, China
China
Chinese civilization may refer to:* China for more general discussion of the country.* Chinese culture* Greater China, the transnational community of ethnic Chinese.* History of China* Sinosphere, the area historically affected by Chinese culture...

.

A second locality for the discovery of Miomachairodus fossils is Anatolia, Yeni Eskihisar
Yeni Eskihisar
Yeni Eskihisar is a late Miocene formation in Yatağan, Muğla. It is well known for its pollen studies.-Vertebrate paleofauna:*Thalassictis montadai *Proticititherium cingulatum...

. This Turkish
Turkey
Turkey , known officially as the Republic of Turkey , is a Eurasian country located in Western Asia and in East Thrace in Southeastern Europe...

 site is of Miocene age and is well known for its pollen studies.

Though none of their fossil beds have provided Miomachairodus, the region of Greece
Greece
Greece , officially the Hellenic Republic , and historically Hellas or the Republic of Greece in English, is a country in southeastern Europe....

 during the middle Miocene is close to Turkey and displays a similar habitat. Aliveri (Evia island) in Greece shows a rich micromammalian fauna but with few large mammals. The island of Chios is known for its middle Miocene faunas from the beginning of 1940s when the first large mammals have been described from a clay pit near Thymiana village. Of the two Vallesian periods (early, or MN9, and late, or MN10) only the late is represented by the formations of Greece.

Morphology

Miomachairodus was generally small and the canines already began to show the flattening that would soon advance into the distinct "sabre-tooth" design of the more advanced machairodonts. Miomachairodus was not the ancestor to all machairodonts, this title most likely rested with Pseudaelurus
Pseudaelurus
Pseudaelurus is a prehistoric cat that lived in Europe, Asia and North America in the Miocene approximately 20-8 million years ago. It is an ancestor of today's felines and pantherines as well as the extinct machairodont saber-tooths, and is a successor to Proailurus...

. This genus is thought to be ancestral to both machairodontinae and felini, the clades of sabre-toothed and conicle-toothed (modern) felids.

Comparison to Relatives

Sister taxa of Miomachairodus is the more well known Machairodus
Machairodus
Machairodus was a genus of large machairodontine saber-toothed cats that lived in Europe, Asia, Africa and North America during the Miocene through Pleistocene living from 11.6mya—126,000 years ago, existing for approximately .-Species:...

. Of the several species of widespread Machairodus, M. giganteus is probably the most well known. The fossil reccord is well established in its wide range of Eurasia. Most mammals, even the largest, started out small, and Machairodus is no exception. Pseudaelurus ogygia stood at 58 cm at the shoulder and its ancestor Miomachairodus advances little in size, but between Miomachairodus and the it's descandants of the tribe Homotheriini (one of three machairodont tribes; including Machairodus and Homotherium
Homotherium
Homotherium is an extinct genus of machairodontine saber-toothed cats, often termed scimitar cats, endemic to North America, South America, Europe, Asia, and Africa during the Pliocene and Pleistocene epochs , existing for approximately .It first became extinct in Africa some 1.5 million years ago...

), they grow to massive sizes, surpassing the modern lion. Most Machairodus species reach 120 cm shoulder height and Homotherium reaches 110 cm, when compared to the 95 cm shoulder height of the modern bengal tiger
Bengal Tiger
The Bengal tiger is a tiger subspecies native to the Indian subcontinent that in 2010 has been classified as endangered by IUCN...

, Panthera tigris. Machairodus rivals in size among the cats only Smilodon
Smilodon
Smilodon , often called a saber-toothed cat or saber-toothed tiger, is an extinct genus of machairodonts. This saber-toothed cat was endemic to North America and South America, living from near the beginning through the very end of the Pleistocene epoch .-Etymology:The nickname "saber-tooth" refers...

 populator
of South American Pleistocene
Pleistocene
The Pleistocene is the epoch from 2,588,000 to 11,700 years BP that spans the world's recent period of repeated glaciations. The name pleistocene is derived from the Greek and ....

.

The older Pseudaelurus is long-backed and appears lanky. It was arborial and spent a great deal of time climbing. Its long tail was adapted for balancing and it was probably not a successful runner, relying on ambush styles rather than long prusuits. It was a forest dweller that was most likely nocturnal or crepuscular
Crepuscular
Crepuscular animals are those that are active primarily during twilight, that is during dawn and dusk. The word is derived from the Latin word crepusculum, meaning "twilight." Crepuscular is, thus, in contrast with diurnal and nocturnal behavior. Crepuscular animals may also be active on a bright...

 and preyed on birds, small mammals, eggs, and, on the rare occasion, fruits, beetles, insects, or scavenged meals as some modern felids will. Pseudaelurus was not a strict carnivore as later descdants would be, but relied more on a carnivorous diet with some glimpses into an omnivorous past.

The later Machairodus was a massive creature with a tail of medium length that did not reach the ground while standing, large ribcage, stout body, and especially strong forequarters. Its skull was long, canines of moderate length for a machairodont, making it a very good representative of the group. Sexual dimorphisim displayed itself in males that were much larger and, unusual for carnivores, the main killing apparatus, the canines, were shaped differently and were surely not capable of exactly the same method of killing. Male canines were shorter and much more stout, while females were long and thin. The question of the specific usage of the canines is still in limbo, and Machairodus offers a good deal of suggestion as to how they were used, giving support to the usually inadequate claim that they were used for show, not for killing, though the hypothsis is still generally discarded. Machairodus was also in the leagues of the extreme carnivores, or hypercarnivores who could consume little but meat, and generally fresh meat at that, so scavenging was usually out of the range of means of sustinance. Modern species displaying themselves as hypercarnivores are Acinonyx jubatus, the cheetah, and Ursus maritimus, the polar bear. It has been thought that Machairodus hunted in groups, along with Homotherium, who are strangly linked to Mammoth
Mammoth
A mammoth is any species of the extinct genus Mammuthus. These proboscideans are members of Elephantidae, the family of elephants and mammoths, and close relatives of modern elephants. They were often equipped with long curved tusks and, in northern species, a covering of long hair...

 juvenile remains, so whether these machairodonts preyed strictly on mammoth young or scavenged or whether the links are unrelated is uncertain.

Miomachairodus lied between these two, but since their fossil history is wildly incomplete, inferences must be made as to where between these two Miomachairodus lied.

Bahe Formation

Species that Miomachairodus would probably have encountered in this region of China would have certainly been reptiles such as Emydidae
Emydidae
Emydidae, commonly called the pond turtles or marsh turtles, is a family of turtles. Previously, several species of Asian box turtle were classified in the family. However, revised taxonomy has separated them to a different family. Now, Emydidae, with the exception of two species of pond turtle,...

and Testudo
Testudo (genus)
Testudo is a genus of tortoises found in North Africa, western Asia, and Europe. Several species are under threat in the wild, mainly from habitat destruction....

. Both are turtles and Miomachairodus was not suited for killing them in any way. Only species with exceptionally strong jaw bites, such as the modern Crocuta, the spotted hyena
Spotted Hyena
The spotted hyena also known as laughing hyena, is a carnivorous mammal of the family Hyaenidae, of which it is the largest extant member. Though the species' prehistoric range included Eurasia extending from Atlantic Europe to China, it now only occurs in all of Africa south of the Sahara save...

 of Africa, could have torn open the shell to retrieve the meat on the inside. These species would have been left alone. Another unwelcoming creature would have been Erinaceus
Erinaceus
Erinaceus is a genus of European hedgehog from the family of Erinaceidae. There are four main species of Erinaceus. The range is all across Europe, throughout the Middle East, parts of Russia, and extending to northern China...

. Members of this genus still live today, and we generally call them hedgehog
Hedgehog
A hedgehog is any of the spiny mammals of the subfamily Erinaceinae and the order Erinaceomorpha. There are 17 species of hedgehog in five genera, found through parts of Europe, Asia, Africa, and New Zealand . There are no hedgehogs native to Australia, and no living species native to the Americas...

s. Most species can but will not eat them for risk of injury caused by the spines, which are, as with porcupines, modified hair, but nevertheless painful and possibly opening the animal up to infection from an open sore. The rhinocerid
Rhinoceros
Rhinoceros , also known as rhino, is a group of five extant species of odd-toed ungulates in the family Rhinocerotidae. Two of these species are native to Africa and three to southern Asia....

 Dicerorhinus was also located in this region at this time. Members of this species still exist, though may not for much longer: the remaining member is the critically endangered Sumatran Rhinoceros
Sumatran Rhinoceros
The Sumatran Rhinoceros is a member of the family Rhinocerotidae and one of five extant rhinoceroses. It is the only extant species of the genus Dicerorhinus. It is the smallest rhinoceros, although is still a large mammal. This rhino stands high at the shoulder, with a head-and-body length of ...

, or Asian one-horned Rhino as it's technically called. During the Miocene, this genus was likely to be small, but probably still too much of a feat for a lone Miomachairodus, though if it worked in groups as its descendants showed, it might have been able to take the young or the injured. Hipparion
Hipparion
Hipparion is an extinct genus of horse living in North America, Asia, Europe, and Africa during the Miocene through Pleistocene ~23 Mya—781,000 years ago, existing for...

was probably prime prey for Miomachairodus at this locality. It was small and light-footed, and though the morphology of Miomachairodus suggests it did not partake on long chases, it probably used the cover of trees, foliage, and shadows during dusk and at night to creep up on herds of Hipparion and capture one in the short struggle that is typical of felines. The suid Chleuastochoerus was again probably another very good candidate for prey of Miomachairodus. Even modern pigs are primitive in form, and Miocene Cheleuastochoerus looked much like the modern pigs we consume in the modern world in their low forms, two toes per foot, and lack of extreme defense besides tusks that can kill if the predator is too inexperienced or lame to subdue the animal quickly. The giraffid Paleotragus looked much like the modern okapi
Okapi
The okapi , Okapia johnstoni, is a giraffid artiodactyl mammal native to the Ituri Rainforest, located in the northeast of the Democratic Republic of the Congo, in Central Africa...

, Okapia johnstoni. Paleotragus was a common prey for many predators of the Miocene, and its descendant lineage which became the Okapia did not change much and as felines took to the adaptive radiation
Adaptive radiation
In evolutionary biology, adaptive radiation is the evolution of ecological and phenotypic diversity within a rapidly multiplying lineage. Starting with a recent single ancestor, this process results in the speciation and phenotypic adaptation of an array of species exhibiting different...

 of later eras, the lineage became common prey for felines living in the region it inhabited. During the Miocene, there were two species of Paleotragus: P. primaevus, who had no ossicones and was smaller, at just under two meters at the shoulder, and P. germaini who bore the ossicones later giraffe
Giraffe
The giraffe is an African even-toed ungulate mammal, the tallest of all extant land-living animal species, and the largest ruminant...

 and okapie males would bear. Miomachairodus might have been too small to take on Paleotragus alone, but if they acquired the adaptation of grouping, it was likely that this animal, too, fell on their list of prey.

Yeni Eckihisar

The other predators noted here, including the large hyaenids Thalassictis
Thalassictis
Thalassictis is an extinct genus of terrestrial carnivore the family Hyaenidae, endemic to Asia and Southern Europe during the Early Miocene subepoch, 8.7—5.3 mya, existing for approximately .-Taxonomy:...

and Proticititherium, probably meant that here, Miomachairodus was increasingly nocturnal to adoid its duirnal relatives. The fauna was likely widely different, and behavioral aspects might have been noted. Solitary lives could be inferenced because smaller prey must have been taken, and as smaller prey was taken, each individual needed more land to support itself and the population was likely more spread out than it was in the Bahe region. It probably preyed more on amphibians and small reptiles, along with small rodents and birds, not often taking the larger fauna that would have been seen in the Bahe regions.

Greece

Again, though this region never produced Miomachairodus fossils, it is in close proximity with those fossil beds in Turkey that did, so for the purposes of comparison when good fossil records are absent, inference of similar ecosystems and species to some degree may be taken. The fossil beds in Greece during the middle and late Miocene uncovered more hyaenids, such as the large Dinocrocuta
Dinocrocuta
Dinocrocuta is an extinct genus of hyena-like feliform carnivores. It lived in Asia, and Africa, during the Miocene epoch. It had very strong jaws that were able to crush bones. It's estimated to weight roughly 400 kg...

who probably would have forced Miomachairodus, if it existed in this region, back into a nighttime hunting style and avoidance of direct competition. Here, the Miocene primate Mesopithecus
Mesopithecus
Mesopithecus is an extinct genus of Old World monkey that lived in Europe and western Asia 7 to 5 million years ago. It was once thought that it might be an ancestor of the grey langur, but a more recent study suggests that they are more closely related to the snub-nosed monkeys and...

of the family Cercopithecidae, or the old world monkeys, of which there are pleanty of members alive today, from Papio, the baboon
Baboon
Baboons are African and Arabian Old World monkeys belonging to the genus Papio, part of the subfamily Cercopithecinae. There are five species, which are some of the largest non-hominoid members of the primate order; only the mandrill and the drill are larger...

, to Semnopithecus including langurs. These primates would have been ideal for hunting of this semi-arborial forest machairodont, and whether alone or in groups, hiding from Dinocrocuta or dealing well with the large carnivore and playing a more active role as a dominate predator in the ecosystem, it would be likely that Miomachairodus would have hunted Mesopithecus. The rhinocerid Dicerorhinus (who's local fossils might mistakenly been identified, actually being the first appearance of Brachypotherium in Europe) was probably not a good candidate for the lone Miomachairodus, although good prey for Dinocrocuta. The small rodent Pogonomys
Pogonomys
Pogonomys is a genus of rodent in the family Muridae.Species include:* Champion's Tree Mouse * D'Entrecasteaux Archipelago Tree Mouse * Large Tree Mouse...

was present in this region and would likely have been small prey for Miomachairodus. The early hominid Ouranopithecus appeared briefly in this region during the late Miocene and could have been prey for Miomachairodus and Dinocrocuta.

Lifestyle

Since the fossil reccord is so scarce for this species, it's hard to tell exactly what this animal would have acted like or how it might have hunted. With species like Smilodon
Smilodon
Smilodon , often called a saber-toothed cat or saber-toothed tiger, is an extinct genus of machairodonts. This saber-toothed cat was endemic to North America and South America, living from near the beginning through the very end of the Pleistocene epoch .-Etymology:The nickname "saber-tooth" refers...

 fatalis
, it's relatively easy to at least make hypothesis about social life, to infer that the presence of terrible injuries that healed indicated hunting large animals and a strong social structire that cared for the wounded. Species like the large and lean Acinonyx pardinensis can be compared with great accuracy to the modern cheetah, Acinonyx jubatus and their hunting styles and social behavior were probably very much alike. With this old species that had no particularly good fossils or modern descendants, the infernences on its life are at best guesses based on the gueses of more well known extinct relatives such as the Homotheriini tribe.

Behavior

As a felid, it probably experienced some degree of sexual dimorphism, most likely with the males being larger. It would have been territorial and might have sprayed it range with urine
Urine
Urine is a typically sterile liquid by-product of the body that is secreted by the kidneys through a process called urination and excreted through the urethra. Cellular metabolism generates numerous by-products, many rich in nitrogen, that require elimination from the bloodstream...

 like modern cats, telling the marker's age, gender, and if it's a female, estrus cycle. Fighting would have been limited, but unlike many other animals, there would probably have been no elaborate fighting ritual: a fight would consist of blood actually being drawn and injuries that, if one was ganged up on as seen in the modern lion
Lion
The lion is one of the four big cats in the genus Panthera, and a member of the family Felidae. With some males exceeding 250 kg in weight, it is the second-largest living cat after the tiger...

, could be fatal. But these fights, for that reason, would be scarce and would probably be concerned with territorial disputes and mating rights. Biting with the incisors could have been suitable, but the canines were growing more fragile, and a rough jerk from the opponent while being bitten could break a canine off completely, as seen with Smilodon and Machairodus, so this was a risk probably not taken. Vocalizations
Animal communication
Animal communication is any behavior on the part of one animal that has an effect on the current or future behaviour of another animal. The study of animal communication, is sometimes called Zoosemiotics has played an important part in the...

 would have been limited, and the main form of communicating would have been scent left in markers, and possibly visual if two came into contact. Motions of the tail and ears alnog with general body positioning would have been utilized for these purposes.

Breeding

Most feline species have young in a select period of time if the seasons are defined enough, but if this animal lived in a warm, forested environment when food was available year round, it might have had offspring at any time in the year. Modern feline cubs of the larger species, specifically Panthera
Panthera
Panthera is a genus of the family Felidae , which contains four well-known living species: the tiger, the lion, the jaguar, and the leopard. The genus comprises about half of the Pantherinae subfamily, the big cats...

are kicked out between one and two years of age, but there is recent evediance that long teeth that the machairodonts were known for took longer to develop and species such as Barbourofelis
Barbourofelis
Barbourofelis is an extinct genus of large, mostly carnivorous mammals of the family Barbourofelidae endemic to North America, during the Miocene living from 13.6—5.3 Ma and existed for approximately .-Taxonomy:...

(a Barbourofelid, not feline, as it is common mistaken), Smilodon, and especially Machairodus, and cubs might have had to stay with their mothers until three, possibly even four years of age, depending on the individual mother's tolerance or readiness to mate again. Numerous Machairodus skulls are found with mature canines still erupting, and until then, they would be inable to hunt for themselves. If the start of long sabres for Miomachairodus meant a longer time until cubs were independent, they might be staying with their mothers for a full year longer than modern cats.

Hunting

The specialization of the dentition
Dentition
Dentition pertains to the development of teeth and their arrangement in the mouth. In particular, the characteristic arrangement, kind, and number of teeth in a given species at a given age...

 of Miomachairodus would have meant a limited number of prey items. Fewer teeth that were primarily slicing premolars and not crushing molars meant that grass or anything with cellulose
Cellulose
Cellulose is an organic compound with the formula , a polysaccharide consisting of a linear chain of several hundred to over ten thousand β linked D-glucose units....

 would not have been on the menu. The specialization of diet, from omnivore eating mostly meat, to obligatory carnivore eating almost nothing but meat, takes its toll on an individual. Certian machairodont groups crossed the boundary from normal obligatory carnivore, including all of today's cats, to the category of hypercarnivore. This group consists of a handful of the species most dependent on meat, from cheetahs, Acinonyx jubatus, who will eat only fresh kills that they have killed themselves through their unique hunting sequence, to polar bears, Ursus maritimus who eat only seals in the Arctic where most of the year there is no plant material to be eaten. Machairodonts such as the Homotheriini group are placed in this league because their dentition denies them eating nothing but meat, and probably fresh, so scavenging is unlikely for Miomachairodus.

This increasing dependency on meat meant that hunting would occupy more time and an any injury could have led to an inability to hunt and starvation, so the species would have to be careful of such risks. This could be combated, on the other hand, if the species was social, as in Smilodon. Risks would still be risks, but if an animal sprained a wrist, it could still fall back on family members as seen in wolves and occasionally lions in the modern day. Lion prides consist of related females and an unrelated male (or several if they are brothers or cousins) who take over the pride and are periodically kicked out by a new male(s). This might have been the way a Miomachairodus familial group functioned.
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