''Plioviverrops'' is an extinct

genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...

hyaenids Hyenas or hyaenas ( ; from Ancient Greek , ) are feliformia, feliform carnivoran mammals belonging to the Family (biology), family Hyaenidae (). With just four extant species (each in its own genus), it is the fifth-smallest family in the orde ...

that were native to

Europe Europe is a continent located entirely in the Northern Hemisphere and mostly in the Eastern Hemisphere. It is bordered by the Arctic Ocean to the north, the Atlantic Ocean to the west, the Mediterranean Sea to the south, and Asia to the east ...

throughout 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

Early Pliocene Early may refer to: Places in the United States * Early, Iowa, a city * Early, Texas, a city * Early Branch, a stream in Missouri * Early County, Georgia * Fort Early, Georgia, an early 19th century fort Music * Early B, stage name of Jamaican d ...

. ''Plioviverrops'' is unique among most hyaenids in its gradual development towards more hypocarnivorous dentition as opposed to most hyaenids which

evolved Evolution is the change in the heritable Phenotypic trait, characteristics of biological populations over successive generations. It occurs when evolutionary processes such as natural selection and genetic drift act on genetic variation, re ...

to have

hypercarnivorous A hypercarnivore is an animal that has a diet that is more than 70% meat, either via active predation or by scavenging. The remaining non-meat diet may consist of non-animal foods such as fungi, fruits or other plant material. Some extant example ...

dentition.Loddi, Cecilia; Madurell-Malapeira, Joan; Bartolini-Lucenti, Saverio. (2025).
Plioviverrops faventinus reloaded: the last survivor of a successful genus
. ''Bollettino della Società Paleontologica Italiana''. 64(1): 77-105. doi:10.4435/BSPI.2025.06. As ''P. orbignyi'' had especially reduced and simplified dentition, it has been proposed that ''P. orbignyi'' may be ancestral to the genus '' Proteles'', including the modern aardwolf. ''Plioviverrops'' may be a

paraphyletic Paraphyly is a taxonomic term describing a grouping that consists of the grouping's last common ancestor and some but not all of its descendant lineages. The grouping is said to be paraphyletic ''with respect to'' the excluded subgroups. In co ...

group. ''Plioviverrops'' is among the limited number of hyaenid

genera Genus (; : genera ) is a taxonomic rank above species and below family as used in the biological classification of living and fossil organisms as well as viruses. In binomial nomenclature, the genus name forms the first part of the binomial s ...

that avoided extinction during the Miocene-Pliocene boundary, when canids first arrived in

Eurasia Eurasia ( , ) is a continental area on Earth, comprising all of Europe and Asia. According to some geographers, Physical geography, physiographically, Eurasia is a single supercontinent. The concept of Europe and Asia as distinct continents d ...

Characteristics

''Plioviverrops'' were very small hyaenids that weighed only 2 to 7

kilograms The kilogram (also spelled kilogramme) is the base unit of mass in the International System of Units (SI), equal to one thousand grams. It has the unit symbol kg. The word "kilogram" is formed from the combination of the metric prefix kilo- (m ...

.Kargopoulos, Nikolaos. (2019).
Carnivoran Dietary Adaptations: A Multiproxy Study on the Feeding Ecology of the Fossil Carnivorans of Greece
. ''P. orbignyi'' and ''P. faventinus'' were morphologically similar to the modern herpestids, while ''P. guerini'' was more similar to a

viverrid Viverridae is a family (biology), family of small to medium-sized feliform mammals, comprising 14 genera with 33 species. This family was named and first described by John Edward Gray in 1821. Viverrids occur all over Africa, in southern Europe, ...

. ''Plioviverrops'' had hypocarnivorous dentition. Its auditory bullae were primitive in structure and, besides the presence of an alisphenoid canal in ''Plioviverrops'', were similar to those of ''Proteles''.

Species

Up to five species are currently recognized in ''Plioviverrops'' by different authors.

''Plioviverrops orbignyi''

The

type species In International_Code_of_Zoological_Nomenclature, zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the spe ...

, ''P. orbignyi'', lived from the

Vallesian The Vallesian age is a period of geologic time (11.6–9.0 Ma) within the Miocene used more specifically with European Land Mammal Ages. It precedes the Turolian age and follows the Astaracian age. The so-called Vallesian Crisis resulted in th ...

to the

Turolian The Turolian age is a period of geologic time (9.0–5.3 Ma) within the Miocene used more specifically with European Land Mammal Ages. It precedes the Ruscinian age and follows the Vallesian age. The Turolian overlaps the Tortonian and Messinia ...

of the

Late Miocene The Late Miocene (also known as Upper Miocene) is a sub-epoch of the Miocene epoch (geology), Epoch made up of two faunal stage, stages. The Tortonian and Messinian stages comprise the Late Miocene sub-epoch, which lasted from 11.63 Ma (million ye ...

. It has been found in multiple localities in

Greece Greece, officially the Hellenic Republic, is a country in Southeast Europe. Located on the southern tip of the Balkan peninsula, it shares land borders with Albania to the northwest, North Macedonia and Bulgaria to the north, and Turkey to th ...

, including Pikermi, Ravin de la Pluie,

Samos Samos (, also ; , ) is a Greek island in the eastern Aegean Sea, south of Chios, north of Patmos and the Dodecanese archipelago, and off the coast of western Turkey, from which it is separated by the Mycale Strait. It is also a separate reg ...

, and Vathylakkos-2. It had reduced

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 mout ...

and a long m1 molar with a high

metaconid Many different terms have been proposed for features of the tooth crown in mammals. The structures within the molars receive different names according to their position and morphology. This nomenclature was developed by Henry Fairfield Osborn i ...

and an especially low

trigonid The molars or molar teeth are large, flat teeth at the back of the mouth. They are more developed in mammals. They are used primarily to grind food during chewing. The name ''molar'' derives from Latin, ''molaris dens'', meaning "millstone tooth ...

''Plioviverrops faventinus''

''P. faventinus'' is the most recent Plioviverrops and the species with the most hypocarnivorous dentition. It lived during the

Messinian The Messinian is in the geologic timescale the last age or uppermost stage of the Miocene. It spans the time between 7.246 ± 0.005 Ma and 5.333 ± 0.005 Ma (million years ago). It follows the Tortonian and is followed by the Zanclean, the fir ...

of the Late Miocene. ''P. faventinus'' has been discovered in Cava Monticino in

Italy Italy, officially the Italian Republic, is a country in Southern Europe, Southern and Western Europe, Western Europe. It consists of Italian Peninsula, a peninsula that extends into the Mediterranean Sea, with the Alps on its northern land b ...

where hyaenids such as ''Plioviverrops'' and ''

Lycyaena ''Lycyaena'' is an extinct genus of terrestrial carnivore in the family Hyaenidae. ''Lycyaena'' was a cursorial hunting hyaena as opposed to full-time scavenger. It has been suggested by R. F. Ewer that ''Lycyaena'' may be a possible ancestor t ...

'' are the most abundant fossil

carnivores A carnivore , or meat-eater (Latin, ''caro'', genitive ''carnis'', meaning meat or "flesh" and ''vorare'' meaning "to devour"), is an animal or plant whose nutrition and energy requirements are met by consumption of animal tissues (mainly mu ...

. ''P. faventinus'' was the most numerous hyaenid in Cava Monticino.

''Plioviverrops gaudryi''

''P. gaudryi'' lived during the

Middle Miocene The Middle Miocene is a sub-epoch of the Miocene epoch (geology), epoch made up of two Stage (stratigraphy), stages: the Langhian and Serravallian stages. The Middle Miocene is preceded by the Early Miocene. The sub-epoch lasted from 15.97 ± 0. ...

and has been discovered in La Grive-Saint-Alban in

France France, officially the French Republic, is a country located primarily in Western Europe. Overseas France, Its overseas regions and territories include French Guiana in South America, Saint Pierre and Miquelon in the Atlantic Ocean#North Atlan ...

''Plioviverrops gervaisi''

''P. gervaisi'' is among the oldest known hyaenids. It lived during the

Orleanian The Orleanian age is a period of geologic time ( MN 3–5, (mya)), within the Miocene and used more specifically with European Land Mammal Ages. It precedes the Astaracian age and follows the Agenian age. ; Agenian-Early Orleanian migration ...

of the

Early Miocene The Early Miocene (also known as Lower Miocene) is a sub-epoch of the Miocene epoch (geology), Epoch made up of two faunal stage, stages: the Aquitanian age, Aquitanian and Burdigalian stages. The sub-epoch lasted from 23.03 ± 0.05 annum, Ma to ...

and has been discovered in Vieux-Collonges in France. The m1 molar of ''P. gervaisi'' is very primitive in structure and similar to that of another early hyaenid, ''

Protictitherium ''Protictitherium'' ( gr. first striking beast) is an extinct genus of hyena that lived across Europe and Asia during the Middle and Late Miocene, it is often considered to be the first hyena since it contains some of the oldest fossils of the fa ...

intermedium''.

''Plioviverrops guerini''

''P. guerini'' lived during the Turolian of the Late Miocene. It has been found in multiple localities in

Spain Spain, or the Kingdom of Spain, is a country in Southern Europe, Southern and Western Europe with territories in North Africa. Featuring the Punta de Tarifa, southernmost point of continental Europe, it is the largest country in Southern Eur ...

, including Los Mansuetos. ''P. guerini'' is slightly greater in size but otherwise similar to P. gaudryi in characteristics. Galiano et al. (2022) placed this species in the genus ''

Gansuyaena ''Gansuyaena'' is an extinct genus of small hyenas that lived through the Middle to Late Miocene. Among all known fossil hyaenids ''Gansuyaena'' was the most closely related to the extant '' Proteles'' genus which contains the aardwolf, but ''Gan ...

.''

Diet

''Plioviverrops'' was likely an opportunistic

omnivore An omnivore () is an animal that regularly consumes significant quantities of both plant and animal matter. Obtaining energy and nutrients from plant and animal matter, omnivores digest carbohydrates, protein, fat, and fiber, and metabolize t ...

and

insectivore file:Common brown robberfly with prey.jpg, A Asilidae, robber fly eating a hoverfly An insectivore is a carnivore, carnivorous animal or plant which eats insects. An alternative term is entomophage, which can also refer to the Entomophagy ...

. It may have preyed on rodents such as '' Parapodemus'' or ''

Micromys ''Micromys'' is a genus of small rodents in the subfamily Murinae. The genus contains two living species: the widespread Eurasian harvest mouse (''Micromys minutus'') of much of Europe and Asia; and the more restricted Indochinese harvest mouse ( ...

''. It may have consumed birds, insects, reptiles, and other small animals such as '' Schizogalerix''. It may have eaten

fungi A fungus (: fungi , , , or ; or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and mold (fungus), molds, as well as the more familiar mushrooms. These organisms are classified as one ...

and

plant matter Organic matter, organic material or natural organic matter is the large source of carbon-based compounds found within natural and engineered, terrestrial, and aquatic environments. It is matter composed of organic compounds that have come fro ...

such as fruits.

Taxonomy

''Plioviverrops'' was named by Kretzoi in 1938, and assigned to Hyaenidae by Flynn in 1998.J. J. Flynn. 1998. Early Cenozoic Carnivora ("Miacoidea"). In C. M. Janis, K. M. Scott, and L. L. Jacobs (eds.), Evolution of Tertiary Mammals of North America ''

Herpestes ''Herpestes'' is a genus within the mongoose family Herpestidae. Several species in the family are known as slender mongooses. It is the type genus of the family, and comprises 5-6 living species, each with several subspecies. Fossil remains of t ...

guerini'' was described by Villalta Comella & Crusafont Pairó (1945) based on material from Los Mansuetos in Spain. Crusafont Pairó & Petter (1969) reassigned this species to ''Plioviverrops'' as ''Plioviverrops guerini''. Recently, Galiano et al. (2022) included this species in the genus Gansuyaena as ''Gansuyaena guerini''. The reassignment of this species to ''Gansuyaena'' was questioned by Loddi et al. (2025). De Beaumont & Mein (1972) established three subgenera for Plioviverrops, including ''Plioviverrops (Plioviverrops) orbignyi'', ''Plioviverrops (Mesoviverrops) gaudryi'', and ''Plioviverrops (Protoviverrops) gervaisi''. Galiano et al. (2022) elevated these subgenera to genera, reassigning ''P. gaudryi'' to ''Mesoviverrops'' as ''Mesoviverrops gaudryi'' and ''P. gervaisi'' to ''Protoviverrops'' as ''Protoviverrops gervaisi''. Galiano et al. (2022) additionally suggested that ''Plioviverrops'' and ''Mesoviverrops'', but not ''Protoviverrops'', should be included in the hyaenid subfamily '' Protelinae''. De Bonis (1994) described ''Plioviverrops collectus'' from material previously known as ''

Herpestides ''Herpestides'' is an extinct genus of terrestrial carnivore that was endemic to North Africa and Southern Europe during the Early Miocene subepoch (22.4—20 mya) and existed for approximately 2.4 million years. Taxonomy ''Herpestides'' is c ...

collectus'' (De Bonis, 1973) from the early

Burdigalian The Burdigalian is, in the geologic timescale, an age (geology), age or stage (stratigraphy), stage in the early Miocene. It spans the time between 20.43 ± 0.05 annum, Ma and 15.97 ± 0.05 Ma (million years ago). Preceded by the Aquitanian (sta ...

of France, and considered it to be the earliest known hyaenid. Turner et al. (2008) noted that while this material was distinct from ''Herpestides'' there was not enough evidence to support its assignment to Hyaenidae.

References

Prehistoric hyenas Miocene carnivorans Messinian extinctions Cenozoic mammals of Europe Cenozoic mammals of Africa Prehistoric carnivoran genera Tortonian extinctions {{paleo-carnivora-stub