Enhydriodon Dikikae
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''Enhydriodon dikikae'' is an extinct species in the family
Mustelidae The Mustelidae (; from Latin ''mustela'', weasel) are a family of carnivorous mammals, including weasels, badgers, otters, ferrets, martens, minks and wolverines, among others. Mustelids () are a diverse group and form the largest family in th ...
that existed during the
Miocene The Miocene ( ) is the first epoch (geology), geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and mea ...
and
Pliocene The Pliocene ( ; also Pleiocene) is the epoch in the geologic time scale that extends from 5.333 million to 2.58Ethiopia Ethiopia, , om, Itiyoophiyaa, so, Itoobiya, ti, ኢትዮጵያ, Ítiyop'iya, aa, Itiyoppiya officially the Federal Democratic Republic of Ethiopia, is a landlocked country in the Horn of Africa. It shares borders with Eritrea to the ...
. These fossils were found in the lower basal member of the Hadar Formation, which means they existed more than 3.42 million years ago but the estimated age of the fossils is greater than 3.4 million years old and likely to be closer to 4 million years old. In comparison to most other species in the genus ''Enhydriodon'', ''E. dikikae'' is one of the youngest. It is suggested that ''E. dikikae'' weighed up to 200 kilograms and grew up to 2 meters in length. The fossils evidence suggests that the ''E. dikikae'' was a giant
semiaquatic In biology, semiaquatic can refer to various types of animals that spend part of their time in water, or plants that naturally grow partially submerged in water. Examples are given below. Semiaquatic animals Semi aquatic animals include: * Ve ...
otter and had a jaw structure that suggests it fed on hard foods but this is still undetermined. While related species have also been found in Europe and Asia, the differences in size and dental structures differentiate ''E. dikikae'' from its relatives.


Taxonomy

''Enhydriodon dikikae'' belongs to the order Carnivora, superfamily
Musteloidea Musteloidea is a superfamily of carnivoran mammals united by shared characters of the skull and teeth. Musteloids are the sister group of pinnipeds, the group which includes seals. The Musteloidea consists of the families Ailuridae ( red pandas) ...
, family
Mustelidae The Mustelidae (; from Latin ''mustela'', weasel) are a family of carnivorous mammals, including weasels, badgers, otters, ferrets, martens, minks and wolverines, among others. Mustelids () are a diverse group and form the largest family in th ...
and subfamily Lutrinae. The closest living relatives to this species are the modern otters. Other related animals in the Musteloidea superfamily are from the family, Procyonidae to which animals like
racoons The raccoon ( or , ''Procyon lotor''), sometimes called the common raccoon to distinguish it from other species, is a mammal native to North America. It is the largest of the procyonid family, having a body length of , and a body weight of ...
, olingos and
coatis Coatis, also known as coatimundis (), are members of the family Procyonidae in the genera '' Nasua'' and '' Nasuella''. They are diurnal mammals native to South America, Central America, Mexico, and the southwestern United States. The name ...
belong to. There is some debate over whether ''E. dikikae'' should belong to the genus ''Sivaonyx'' however at this point, it is still classified as being in the genus ''
Enhydriodon ''Enhydriodon'' is an extinct genus of typically large otters that lived in what is now Kenya, Ethiopia, Uganda, Chad, South Africa, Pakistan, and India from the late Miocene up to early Pleistocene. The otter is thought to be a relative of moder ...
''.


Description

Fossils of a partial skull which included a snout, orbit, parts of the lower jaw and most of the teeth that belonged to ''E. dikikae''. According to these fossils, ''E. dikikae'' was similar in size to a bear but resembled the modern otter. It is estimated to have weighed 100–200 kg and to be over 2m in body length. The fossils found in
Ethiopia Ethiopia, , om, Itiyoophiyaa, so, Itoobiya, ti, ኢትዮጵያ, Ítiyop'iya, aa, Itiyoppiya officially the Federal Democratic Republic of Ethiopia, is a landlocked country in the Horn of Africa. It shares borders with Eritrea to the ...
suggest that the species had a large skull, approximately 25 cm with a short, non-prognathic snout. The
dental arch The dental arches are the two arches (crescent arrangements) of teeth, one on each jaw, that together constitute the dentition. In humans and many other species; the superior (maxillary or upper) dental arch is a little larger than the inferior ...
was broad, with long, strong
canines Canine may refer to: Zoology and anatomy * a dog-like Canid animal in the subfamily Caninae ** ''Canis'', a genus including dogs, wolves, coyotes, and jackals ** Dog, the domestic dog * Canine tooth, in mammalian oral anatomy People with the surn ...
grown laterally on the
maxilla The maxilla (plural: ''maxillae'' ) in vertebrates is the upper fixed (not fixed in Neopterygii) bone of the jaw formed from the fusion of two maxillary bones. In humans, the upper jaw includes the hard palate in the front of the mouth. T ...
and large
premolars The premolars, also called premolar teeth, or bicuspids, are transitional teeth located between the canine and molar teeth. In humans, there are two premolars per quadrant in the permanent set of teeth, making eight premolars total in the mouth ...
. The limb-bone fossils measured up to 27 cm in length. ''Enhydriodon dikikae'' differs from other species in the genus ''
Enhydriodon ''Enhydriodon'' is an extinct genus of typically large otters that lived in what is now Kenya, Ethiopia, Uganda, Chad, South Africa, Pakistan, and India from the late Miocene up to early Pleistocene. The otter is thought to be a relative of moder ...
'' in its loss of frontal premolars and a divided lower molar, as well as its overall size. Most of the relatives of ''E. dikikae'' are smaller in size. Unlike its Indian relative, ''Enhydriodon falconeri'' in the larger points in the crown of its teeth. The dental structures of ''E. dikikae'' also differ from its relatives ''Enhydriodon ekecaman'' from Kenya, ''Enhydriodon hendeyi'' from South Africa, and the Enhydriodon africanus from Namibia. The ''E. dikikae'' is also generally larger in size than the other ''Enhydriodon'' species. In comparison to most other ''Enhydriodon'' species, ''E. dikikae'' is relatively young. The fossils of ''E. dikikae'' are likely to be close to 4 million years old, which falls into the mid-Pliocene epoch. While ''Enhydriodon sivalensis'' is estimated to have existed in the late Pliocene which is later than ''E. dikikae'', the rest of the ''Enhydriodon'' species existed in the Early Pliocene, with the earliest fossils of ''Enhydriodon campanii'' from Italy and ''Enhydriodon lluecai'' from Spain dating back to the Late Miocene.


Paleoecology

The ''E. dikikae'' fossils were found in Dikika, Lower Awash, Ethiopia along with aquatic and terrestrial species from that epoch. Carbon and oxygen isotopes of tooth fossils from various other species that existed during the same time as ''E. dikikae'' were used to determine the habitat and food resources available in Pliocene Dikika. The isotopic analyses suggested that the landscape of Dikika during this part of the Pliocene was between grassy woodland and open grassland so habitat resources consisted mostly of woodlands and shrublands, with a shift towards open and arid environments. The oxygen isotopes analysis also indicated that the area was in close proximity to a body of water, most likely a freshwater swamp or shallow lake and that there were various drinking water resources. Studies suggest that there was a decrease in precipitation and increase in temperatures during the Pliocene and the abundance of grassy habitat allowed the fauna to be dominated by terrestrial animals. However, fossils found in the same research site as ''E. dikikae'' and at the same stratigraphical level included aquatic species like
fish Fish are aquatic, craniate, gill-bearing animals that lack limbs with digits. Included in this definition are the living hagfish, lampreys, and cartilaginous and bony fish as well as various extinct related groups. Approximately 95% of ...
, crocodiles, and
hippopotamus The hippopotamus ( ; : hippopotamuses or hippopotami; ''Hippopotamus amphibius''), also called the hippo, common hippopotamus, or river hippopotamus, is a large semiaquatic mammal native to sub-Saharan Africa. It is one of only two extan ...
es as well as terrestrial species. Other fossil evidence suggests that ''E. dikikae'' coexisted with more than one other
Mustelidae The Mustelidae (; from Latin ''mustela'', weasel) are a family of carnivorous mammals, including weasels, badgers, otters, ferrets, martens, minks and wolverines, among others. Mustelids () are a diverse group and form the largest family in th ...
species such as the smaller ''E. ekecaman'' and ''Torolutra''. It is likely that the difference in size between these species forced them to prey on animals of different sizes and therefore they were not in competition with each other. There is no evidence to explain why ''E. dikikae'' went extinct after the
Pliocene The Pliocene ( ; also Pleiocene) is the epoch in the geologic time scale that extends from 5.333 million to 2.58Lars Werdelin Lars Werdelin (born 1955) is a Swedish paleontologist specializing in the evolution of mammalian carnivores. One area of interest has been the evolutionary interaction of carnivores and hominins in Africa. He received his Ph.D. A Doctor of P ...
suggested that the emergence of early
hominins The Hominini form a taxonomic tribe of the subfamily Homininae ("hominines"). Hominini includes the extant genera ''Homo'' (humans) and '' Pan'' (chimpanzees and bonobos) and in standard usage excludes the genus ''Gorilla'' (gorillas). The t ...
may have been a possible cause for the extinction of the large
carnivore A carnivore , or meat-eater (Latin, ''caro'', genitive ''carnis'', meaning meat or "flesh" and ''vorare'' meaning "to devour"), is an animal or plant whose food and energy requirements derive from animal tissues (mainly muscle, fat and other s ...
s that existed at the time due to indirect competition. He suggests that
hominins The Hominini form a taxonomic tribe of the subfamily Homininae ("hominines"). Hominini includes the extant genera ''Homo'' (humans) and '' Pan'' (chimpanzees and bonobos) and in standard usage excludes the genus ''Gorilla'' (gorillas). The t ...
may have competed with these animals for
habitat In ecology, the term habitat summarises the array of resources, physical and biotic factors that are present in an area, such as to support the survival and reproduction of a particular species. A species habitat can be seen as the physical ...
s, forcing a change in some behaviours like
predation Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill th ...
behaviours, and subsequently the effect of the early hominins may have forced these carnivores into extinction.


Paleobiology

The fossils of ''E. dikikae'' had thinner limb bones than aquatic species but weaker than fully
terrestrial Terrestrial refers to things related to land or the planet Earth. Terrestrial may also refer to: * Terrestrial animal, an animal that lives on land opposed to living in water, or sometimes an animal that lives on or near the ground, as opposed to ...
predator Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill th ...
s, suggesting aquatic and non-aquatic adaptations. It is likely that the species spent more time on land than in the water unlike their later relative, ''E. sivalensis'' that were mostly aquatic and lived under boulders or rocks, in dens or under tree trunks along the banks of streams or lakes. The structure of the jaw and teeth of ''E. dikikae'' were used to determine the possible diet of the species. The large teeth along the cheekbone of the skull indicates that their diet may have consisted of hard food. This is supported by the large size of the crushing parts of the teeth that would have been used to break through their hard food. Due to the large size of ''E. dikikae'', it is unlikely that it would have been able to catch fast-moving prey like fish, however
catfish Catfish (or catfishes; order Siluriformes or Nematognathi) are a diverse group of ray-finned fish. Named for their prominent barbels, which resemble a cat's whiskers, catfish range in size and behavior from the three largest species alive ...
were found in shallow waters during the same period of time and the purpose of the jaw structure could be for crushing the skull of the catfish to prevent ''E. dikikae'' from getting hurt. A study suggests that, like their relative species, ''E. falconeri'' and ''E. sivalensis'', ''E. dikikae'' may have fed on bivalves because of a common incisor arch in the three species which would have been used to crush the shells of the bivalves. However, that would mean that the large canines and the larger points in the crown of its teeth would have no purpose in digesting the food and to further disprove this possibility, there have not been many bivalve fossils found at Dikika. Other possible contributions to the diet of ''E. dikikae'' were any prey with a hard
exoskeleton An exoskeleton (from Greek ''éxō'' "outer" and ''skeletós'' "skeleton") is an external skeleton that supports and protects an animal's body, in contrast to an internal skeleton (endoskeleton) in for example, a human. In usage, some of the ...
like young
crocodiles Crocodiles (family Crocodylidae) or true crocodiles are large semiaquatic reptiles that live throughout the tropics in Africa, Asia, the Americas and Australia. The term crocodile is sometimes used even more loosely to include all extant memb ...
,
turtles Turtles are an order of reptiles known as Testudines, characterized by a special shell developed mainly from their ribs. Modern turtles are divided into two major groups, the Pleurodira (side necked turtles) and Cryptodira (hidden necked tur ...
,
ostrich Ostriches are large flightless birds of the genus ''Struthio'' in the order Struthioniformes, part of the infra-class Palaeognathae, a diverse group of flightless birds also known as ratites that includes the emus, rheas, and kiwis. There ...
eggs, or
molluscs Mollusca is the second-largest phylum of invertebrate animals after the Arthropoda, the members of which are known as molluscs or mollusks (). Around 85,000 extant species of molluscs are recognized. The number of fossil species is estim ...
.


References

{{Taxonbar, from=Q60741848 Otters Prehistoric mustelids Miocene mammals of Africa Pliocene mammals of Africa