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Percrocutidae
Percrocutidae
Percrocutidae
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Percrocutidae
Temporal range: Middle Miocene to Late Pliocene
Dinocrocuta gigantea skull cast, Zoological Museum in Copenhagen
Scientific classification Edit this classification
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Carnivora
Superfamily: Herpestoidea
Family: †Percrocutidae
Werdelin & Solounias, 1991
Genera

Percrocutidae is an extinct family of hyena-like feliform carnivorans endemic to Asia, Africa, and Southern Europe from the Middle Miocene through the Pliocene, existing for about 8 million years.[1]

The first percrocutids are known from the middle Miocene of Europe and western Asia and belonged to the genus Percrocuta. Percrocuta already had large premolars, but did not carry such a massive bite as the later form Dinocrocuta, from the later Miocene.[2] Originally, these carnivores were placed with the hyenas in the family Hyaenidae. As of 2022, most scientists considered the Percrocutidae to be a distinct family that evolved their morphology similar to hyenas due convergent evolution,[3] - although they are usually placed sister-taxa/immediate outgroup to Hyaenidae.[4] Sometimes it was placed with the family Stenoplesictidae into the superfamily Stenoplesictoidea. A 2022 study placed Dinocrocuta and Percrocuta as true hyaenids, which if correct would invalidate the family Percrocutidae.[3]

Taxonomy and evolution

[edit]

Taxonomic history

[edit]

Percrocuta was first considered as a side-branch outside of Hyaenidae by Thenius in 1966.[5] It was later named as a different subfamily, Percrocutinae, of Hyaenidae in 1976, and at that time was proposed to include Percrocuta, Adcrocuta eximia, and Allohyaena kadici.[6] Dinocrocuta was elevated from a subgenus to a full genus in 1988.[7]

The family Percrocutidae was formally elevated in 1991, to include the genera Percrocuta, Dinocrocuta, Belbus and Allohyaena.[8]

Later studies have suggested that Belbus and Allohyaena are true hyaenids and not percrocutids.[9]

Classification

[edit]
Family Image Genus Species
†Percrocutidae †Dinocrocuta (Schmidt-Kittler, 1975)
  • †D. algeriensis
  • †D. gigantea
  • †D. salonicae
  • †D. senyureki
†Percrocuta (Kretzoi, 1938)
  • †P. abessalomi
  • †P. carnifex
  • †P. grandis
  • †P. leakeyi
  • †P. miocenica
  • †P. tobieni
  • †P. tungurensis
  • †P. xixiaensis

The list follows McKenna and Bell's Classification of Mammals for prehistoric genera (1997).[10] In contrast to McKenna and Bell's classification, they are not included as a subfamily into the Hyaenidae but as a separate family Percrocutidae.

References

[edit]
Revisions and contributorsEdit on WikipediaRead on Wikipedia

Percrocutidae

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from Grokipedia
Percrocutidae is an extinct family of feliform carnivorans characterized by hyena-like adaptations, including robust skulls and dentition suited for bone-crushing, that lived during the Miocene epoch across the Old World. However, some recent studies propose that percrocutids are actually part of the Hyaenidae family. The family was established to distinguish certain Miocene and Pliocene taxa previously grouped with hyaenids, featuring key morphological traits such as the absence of the second upper and lower molars (M2/m2) and enlarged premolars, particularly P3/p3, which supported a hypercarnivorous or durophagous lifestyle. Fossils indicate they were large predators, with body sizes ranging from medium to very large, often exceeding those of modern spotted hyenas in some species. The primary genera include Percrocuta and , with Percrocuta encompassing species like P. miocenica and P. carnifex, while Dinocrocuta includes D. gigantea; other taxa such as Belbus have been tentatively assigned but remain debated. These animals are recorded from the late Middle Miocene (approximately 13–11 million years ago) through the (Turolian stage, around 8–5 million years ago), with some possible extensions into the early , though the family's diversity peaked in the . Geographically, Percrocutidae fossils are widespread in , from and in the west to and the Siwalik deposits of India-Pakistan in the east, as well as sites in the (e.g., , , ) and eastern Africa (e.g., ). Evolutionary relationships have placed percrocutids as a distinct basal group within Feliformia, potentially sister to or ancestral to Hyaenidae, though a 2022 study argues they are true hyaenids. They exhibit convergent traits with hyenas such as powerful jaws and scavenging behaviors inferred from dental wear patterns. Their decline by the end of the Miocene coincides with the radiation of true hyaenids and increased competition from other carnivorans, leaving no modern descendants. Notable fossil assemblages, such as those from Samos Island in Greece and the Bahe Formation in China, highlight their role as apex predators in diverse Miocene ecosystems.

Taxonomy

Taxonomic history

The genus Percrocuta was first established by Kretzoi in 1938 to accommodate the Hyaena carnifex, originally described by Pilgrim in 1913 from Miocene deposits in the Siwalik Hills of northern and . Early classifications placed Percrocuta within Hyaenidae, reflecting its hyena-like cranial and dental morphology, though some contemporaneous material was occasionally misattributed to creodonts due to limited comparative studies. The Dinocrocuta, described by Colbert in 1935 based on large carnivoran fossils from , was similarly assigned to Hyaenidae, with its robust premolars suggesting a bone-cracking lifestyle akin to later . In a key revision, Thenius (1966) elevated Percrocuta to full generic status and proposed it as a precocious side-branch diverging early from Hyaenidae, emphasizing differences in auditory bulla development and deciduous dentition, such as the distinctive morphology of the fourth deciduous lower premolar (dp4). This work extended to Dinocrocuta, highlighting its hypertrophied carnassials and premolars as adaptations potentially independent of true hyaenid evolution. These features fueled debates on whether percrocutids represented convergent bone-crushers, with the inflated bulla and specialized dp4 cited as evidence of parallel evolution rather than close relation to Hyaenidae. The family Percrocutidae was formally erected by Werdelin and Solounias in 1991 to distinguish these taxa from true hyaenids, based on shared autapomorphies including frontal orbital position, reduced frontal sinuses, and dental specializations for durophagy that differed from the Hyaenidae's carnassial-dominated occlusion. This classification emphasized the family's position within but outside Hyaenidae, attributing similarities like powerful jaws to ecological convergence in carnivoran guilds. Recent analyses have challenged this separation, with Xiong (2022) describing a new species, Percrocuta xixiaensis, from early middle and arguing that the auditory bulla morphology in Percrocuta and aligns closely with the hyaenid pattern, featuring a septate structure unlike that in other feliforms. This evidence, combined with re-evaluations of , suggests percrocutids may warrant subfamily status within Hyaenidae rather than a distinct family, potentially resolving long-standing debates on their phylogenetic affinities through shared basicranial traits.

Classification and phylogeny

Percrocutidae is an extinct family within the order , suborder , and superfamily Feloidea. Phylogenetic analyses based on morphological and craniodental data position Percrocutidae as the to Hyaenidae, with both families sharing middle origins in and convergent adaptations for bone-cracking, such as robust skulls and strong cheek teeth. More recent morphological studies, incorporating detailed cranial and basicranial features, suggest Percrocutidae may instead represent a basal within Hyaenidae or warrant reassignment to status (Percrocutinae), challenging its status as a distinct family. The family indisputably includes the genera Percrocuta and , both characterized by large body sizes and hyena-like adapted for hypercarnivory. Other taxa, such as Belbus and Allohyaena, were historically assigned to Percrocutidae but have been reclassified as true hyaenids based on dental and cranial evidence, leaving them as relative to the core percrocutid genera. Basicranial morphology provides key evidence for the close affinity between percrocutids and hyaenids, with exhibiting a hyaenid-pattern bulla (annular ectotympanic fused to the entotympanic, forming a double-chambered structure), which distinguishes early percrocutids from more distant feliforms like viverrids but aligns them closely with Hyaenidae. This feature supports interpretations of Percrocutidae as either a sister lineage or an early offshoot within the hyaenid radiation.

Anatomy

Cranial and dental features

Percrocutids possessed robust skulls characterized by a low and a prominent , adaptations that supported powerful jaw musculature for carnivorous feeding. In Percrocuta xixiaensis, the cranium is low and flat, with a thin approximately 2 cm high formed by the joined temporal lines at the postorbital constriction, and a broad, flat interorbital region contributing to the overall low profile of the . The is small, extending only anterior to the and not reaching the occipital region, which suggests a lesser degree of specialization for extreme bone-cracking compared to modern hyaenids. The auditory bullae in percrocutids exhibit a hyaenid-like , featuring a caudal-entotympanic sinus as an aeluroid apomorphy, with large, globular structures divided into anterior and posterior chambers by a subhorizontal formed potentially by both ectotympanic and caudal entotympanic elements. In Dinocrocuta gigantea, the bullae are elongated and ovoid, bulging ventrally up to 16 mm below the basicranial plane, with no lateral bony outgrowth of the external acoustic , aligning closely with hyaenid morphology but differing from more primitive forms like Stenoplesictis. Dentition in percrocutids is adapted for hypercarnivory, with large, robust premolars suited for shearing and initial processing of tough tissues. The P3 and p3 are widened anteriorly with distinct anterolingual cusplets and well-developed accessory cusps, while p4 features a high posterolingual shelf and a large anterior cusplet comprising about 30% of its total length; in early species like Percrocuta, these premolars are particularly high-crowned and robust. The , P4 and m1, show specialization with an extremely reduced protocone on P4 and a short, wide m1 positioned higher than the premolars, accompanied by a reduced talonid that is small and bicuspid, facilitating efficient slicing rather than grinding. In later forms such as , molars exhibit bone-crushing capabilities, with the M1 being very short and wide, oriented diagonally to the P4 metastyle and lacking distinct cusps, supported by two roots for enhanced durability. Variations across genera include more enlarged premolars in early Percrocuta species compared to the sectorial dentition in , where enhanced bite force is facilitated by robust zygomatic processes and a backward-shifted , allowing for greater in durophagous feeding.

Postcranial skeleton and body size

The postcranial skeleton of percrocutids is known from fragmentary remains, including limb bones that indicate a robust overall build adapted to a predatory similar to that of modern hyaenids. Forelimbs exhibit strong, robust morphology, as evidenced by the proximal half of a left from senyureki, which features a slender shaft but a more robust structure compared to contemporaneous felids and ursids, with a short lateral coronoid process and deep radial notch suggestive of powerful grappling or digging capabilities. Postcranial remains are limited, with no detailed descriptions of hindlimb proportions available. Body size in Percrocutidae varied significantly through the , with early forms such as Percrocuta species estimated at 63–100 kg, reflecting smaller, more generalized builds in the early to middle . Larger species, exemplified by Dinocrocuta gigantea, had estimated masses of around 200 kg. Some estimates for large individuals suggest up to approximately 300 kg.

Evolutionary history

Origins and temporal range

The Percrocutidae, an extinct family of hyena-like feliform carnivorans, originated during the Middle Miocene, approximately 15–11 million years ago (Ma), in and western , likely evolving from herpestoid ancestors within the broader clade. This emergence reflects a transition from earlier feliform lineages, characterized by adaptations toward durophagous feeding in forested to open woodland environments of the period. Phylogenetic analyses position Percrocutidae as a distinct group possibly basal to or sister with Hyaenidae, sharing traits like robust but differing in cranial proportions. Recent phylogenetic studies (as of 2022) propose that Percrocutidae may represent basal Hyaenidae rather than a distinct family. The family's temporal range spans from the Middle Miocene to the early , roughly 14–3.6 Ma, encompassing a duration of over 10 million years across and . Peak diversity occurred in the , with multiple genera coexisting before a decline in the . Fossil evidence from Vallesian (MN10) faunas, dated to around 11.6–9.7 Ma, documents this diversification phase, highlighting the family's role in early carnivoran guilds. The earliest definitive records of Percrocutidae belong to Percrocuta miocenica, preserved in sediments from approximately 14 Ma sites in (Brajkovac locality, MN6 zone) and western . These hemimandibles and dental fragments indicate an early durophagous niche, with associated faunas including proboscideans and ruminants typical of the Langhian stage. Such finds underscore the family's rapid establishment in Eurasian ecosystems shortly after its origin.

Diversification and extinction

The Percrocutidae exhibited notable diversification in the late Miocene, transitioning from the smaller, more generalized Percrocuta to the larger, specialized Dinocrocuta, which achieved body masses up to 250 kg. This size increase was accompanied by enhanced dental adaptations, including hypertrophied premolars and sectorial carnassials, enabling efficient bone-cracking and meat-processing in open, grassland-dominated habitats. These traits represented convergent evolution with later Hyaenidae, allowing percrocutids to exploit cursorial hunting and scavenging niches amid expanding savannas. Diversification peaked during the Turolian stage (ca. 8–5 Ma), when multiple species coexisted across and , from sites in and to North African localities. This radiation encompassed at least three genera (Percrocuta, , and Belbus), with regional variations reflecting local ecological opportunities, such as the presence of large prey in aridifying landscapes. The family's abundance in faunas underscores its role as a key carnivoran before the turnover. The Percrocutidae became extinct during the , with the youngest records from early sediments (~3.6 Ma) in and , and none surviving into the Pleistocene. Their disappearance likely stemmed from competitive exclusion by advancing Hyaenidae, which outcompeted them in durophagous niches through superior social hunting strategies and broader dietary flexibility. Concurrent climatic cooling and aridification further pressured percrocutid populations by altering habitats and reducing megaherbivore prey bases, facilitating the ecological replacement by modern hyaenids.

Distribution and paleoecology

Geographic range

Percrocutidae fossils are known exclusively from the , with a primary geographic range encompassing , , and during the epoch. In , records are concentrated in regions such as , , , and , reflecting an early presence in the western part of . The family's distribution extended eastward across , including key areas in and , before reaching in the later stages of their temporal range. No fossils have been reported from or other continents, underscoring their restriction to Afro-Eurasian landmasses. The earliest European sites date to the middle Miocene, with notable occurrences at Paşalar in , approximately 14 million years ago (Ma), marking one of the initial appearances of the family in the region. Additional middle Miocene finds come from sites like Brajkovac in , highlighting a Balkan-Anatolian core for early diversification. By the , the range had broadened, as evidenced by abundant material from Pikermi in , a classical locality yielding diverse percrocutid remains, and Spanish sites such as Algezares and Ademuz. These European records suggest an initial Eurasian foothold before broader dispersal. In Asia, percrocutids achieved widespread distribution, with significant expansion into the via the Siwalik Group sediments of and . The Dhok Pathan Formation in , part of the middle Siwaliks, has produced well-preserved specimens, indicating a robust presence during the (around 8–7 Ma). Farther east, the Tunggur Formation in , , preserves fossils from the middle to , while other Chinese localities like Tongxin in extend the record back to the early middle . These Asian sites illustrate a pattern of eastward and southward proliferation from probable Eurasian origins. African records are restricted to the , appearing later than in and suggesting southward dispersal across the or Mediterranean connections. Fossils from Nakali in , dated to approximately 10–9 Ma, represent one of the few well-documented occurrences, with material attributed to large percrocutids. While North African sites, such as those in , have yielded fragmentary evidence, the overall continental footprint remains limited compared to . This latitudinal gradient in distribution aligns with migratory patterns from a Eurasian cradle, facilitating the family's adaptation to diverse environments before their extinction.

Habitat, diet, and behavior

Percrocutidae inhabited a range of environments across and , from seasonal woodlands and forests to more open savannas and steppes, as inferred from associated faunal assemblages at fossil sites. For instance, at the middle Paşalar locality in , the presence of Percrocuta alongside diverse arboreal and browsing mammals suggests a forested with seasonal availability of resources. In contrast, sites like the Linxia Basin in China, where Dinocrocuta gigantea occurs with grassland-adapted taxa such as Chilotherium wimani, indicate open steppe environments. Similarly, the Nakali site in , yielding Percrocuta leakeyi with and other open-country ungulates, points to savanna-like conditions in the Valley. The diet of Percrocutidae was predominantly hypercarnivorous, with a specialization in bone-crushing (durophagy), enabling them to exploit carcasses of large ungulates through scavenging and active predation. Dental features, such as robust premolars and reduced talonids, supported this feeding strategy, allowing efficient processing of bones similar to that seen in later hyaenids. Genera like Percrocuta exhibited transitional durophagous adaptations suited for cracking bones of medium to large prey, while larger species of , such as D. gigantea, combined bone-crushing capabilities with sectorial for slicing flesh, facilitating opportunistic scavenging and . Evidence from a healed bite mark on a Chilotherium wimani , matching the canine dimensions of D. gigantea, confirms predatory attacks on sizable herbivores like rhinocerotids. Behaviorally, Percrocutidae functioned as top carnivores in ecosystems, occupying guilds and likely engaging in both solitary and social hunting strategies depending on and context. Osteological indicators in , including strong forelimbs for grappling, suggest active predatory behavior akin to modern spotted (Crocuta crocuta), with the capacity to subdue large prey such as rhinoceroses during vulnerable periods like calving or drought. Their large body sizes, with D. gigantea estimated at around 200–250 kg, imply potential for pack-like cooperation in or defense, though direct evidence is limited to inferences from size dimorphism and ecological roles. Overall, they transitioned from more generalized carnivory in early forms to specialized bone-crushing niches, influencing community dynamics by reducing competition with felids and amphicyonids.

Genera

Percrocuta

Percrocuta is the of the extinct family Percrocutidae, comprising hyena-like feliform carnivorans known from the of , , and adjacent regions. Named by Kretzoi in 1938, it includes eight species: P. abessalomi, P. carnifex (the , originally described as Hyaena carnifex by Pilgrim in 1913), P. grandis, P. leakeyi, P. miocenica, P. tobieni, P. tungurensis, and P. xixiaensis. These species exhibit medium to large body sizes, with estimated masses of 70–150 kg on average, smaller than the more robust . Their shows durophagous adaptations, including robust premolars for bone-cracking, but with less extreme sectoriality and a more developed P4 protocone compared to later percrocutids, indicating a transitional specialization for scavenging and predation. Fossils of Percrocuta span the middle to (approximately 16–5.3 million years ago), with the earliest records from early middle deposits. Notable specimens include a well-preserved hemimandible of P. miocenica from the middle locality of Brajkovac in , providing insights into mandibular robusticity and morphology. In , Chinese sites yield significant material, such as the complete cranium of P. xixiaensis from the early middle Zhang'enbao Formation at Tongxin, , which reveals details of cranial proportions and bulla structure suggesting close affinities to true hyaenids. Other key Asian fossils encompass isolated teeth and jaw fragments of P. carnifex and P. tungurensis from late horizons in and , respectively. The genus is primarily distributed in and , with records from over 20 middle localities across the , including , , , and . African occurrences are rarer but include P. leakeyi from late sites in , such as Nakali, highlighting a broader paleobiogeographic range during peak diversification. These distributions reflect adaptation to and open habitats where Percrocuta likely exploited carcasses and smaller prey as opportunistic feeders.

Dinocrocuta

Dinocrocuta is an extinct of percrocutid carnivorans, distinguished as the largest members of the family, with body mass estimates reaching up to 300 kg in species like D. gigantea. The genus comprises seven recognized : D. algeriensis, D. gigantea, D. grandis, D. minor, D. robusta, D. salonicae, and D. senyureki. These taxa exhibit advanced adaptations for bone-crushing, including hypertrophied and robust premolars (particularly P₂ and P₄) and a reinforced with increased mechanical strength, enabling them to process large, durable prey items more effectively than basal percrocutids. Such features underscore Dinocrocuta's role as a dominant durophagous predator in late ecosystems. The temporal range of Dinocrocuta extends from the (Vallesian to Turolian stages, approximately 11.6–5.3 million years ago) into the early . Key fossil evidence includes an exceptionally complete of D. gigantea from the Batallones-3 locality in , part of a rich carnivoran assemblage that preserved around 2000 bones representing at least 16 individuals, highlighting the species' abundance in Iberian late Miocene faunas. Additional significant material comprises a well-preserved left from the Markhal area in Pakistan's Dhok Pathan Formation, providing insights into Asian populations of the . Dinocrocuta fossils are distributed across , , and , reflecting its broad paleoecological success. For instance, D. algeriensis is recorded from Algerian sites, D. salonicae from the Axios Valley in , D. senyureki from early deposits in and , and D. gigantea from diverse localities including , , , and . Evidence of active predatory behavior is documented in D. gigantea through osteological traces, such as deep bite marks and associated fractures on a rhinocerotid (Chilotherium wimani) from the of , indicating failed but aggressive attacks on large herbivores. This suggests Dinocrocuta employed or pursuit strategies akin to modern large carnivores, complementing its scavenging habits.

Other genera

Belbus is a minor genus previously assigned to Percrocutidae but now classified within Hyaenidae, known primarily from limited dental remains that reveal a small-bodied form with primitive , including relatively unspecialized and premolars adapted for a mixed carnivorous-scavenging diet. These fossils date to the early to middle and have been recovered from sites in , such as Pikermi in , where they indicate a body size estimated at around 20-30 kg, smaller than the more derived Percrocuta and . The scarcity of postcranial material has led to ongoing debates about its exact phylogenetic position, though it is consistently recognized as a basal hyaenid based on its dental morphology. Allohyaena represents another peripheral genus with percrocutid affinities, characterized by hyaena-like cranial features such as robust jaws and enlarged premolars suggestive of bone-cracking capabilities, though its fossils are fragmentary and primarily consist of isolated teeth and jaw fragments from deposits. Originally described from Asian localities including , it was initially classified within Percrocutidae due to shared traits like the absence of certain molars and frontal orbital positioning. However, subsequent analyses based on additional European material have reclassified it to Hyaenidae, highlighting taxonomic uncertainties stemming from the incomplete nature of the specimens. Several taxa, including indeterminate percrocutid remains from Asian sites, further complicate the family's diversity, with possible additional forms suggested by isolated teeth exhibiting intermediate features between Percrocutidae and early hyaenids. These fragmentary Asian fossils, often from strata in regions like and , underscore ongoing debates on generic validity due to limited diagnostic material, preventing firm assignment beyond the family level.

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