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Plesiadapis
Plesiadapis
Plesiadapis
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Plesiadapis
Temporal range: Late Paleocene-Early Eocene[1]
Reconstructed skeleton of P. cookei at the University of Michigan Museum of Natural History
Scientific classification Edit this classification
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: †Plesiadapiformes
Family: †Plesiadapidae
Genus: †Plesiadapis
Gervais, 1877
Type species
†Plesiadapis tricuspidens

Plesiadapis is one of the oldest known primate-like mammal genera which existed about 58-55 million years ago in North America and Europe.[2][3] Plesiadapis means "near-Adapis", which is a reference to the adapiform primate of the Eocene period, Adapis. Plesiadapis tricuspidens, the type specimen, is named after the three cusps present on its upper incisors.

Taxonomy

[edit]
Life restoration at MUSE - Science Museum in Trento

The first discovery of Plesiadapis was made by François Louis Paul Gervaise in 1877, who first discovered Plesiadapis tricuspidens in France. The type specimen is MNHN Crl-16, and is a left mandibular fragment dated to the early Eocene epoch.

This genus probably arose in North America and colonized Europe on a land bridge via Greenland. Thanks to the abundance of the genus and to its rapid evolution, species of Plesiadapis play an important role in the zonation of Late Paleocene continental sediments and in the correlation of faunas on both sides of the Atlantic. Two remarkable skeletons of Plesiadapis, one of them nearly complete, have been found in lake deposits at Menat, France.[2] Although the preservation of the hard parts is poor, these skeletons still show remains of skin and hair as a carbonaceous film—something unique among Paleocene mammals. Details of the bones are better preserved in fossils from Cernay, also in France, where Plesiadapis is one of the most common mammals.

Classification

[edit]

The following are possible shared derived features of Plesiadapiformes: maxillary-frontal contact in orbit, the presence of a suboptic foramen, an ossified external auditory meatus, the absence of a promontory artery, the absence of a stapedial artery, and a strong mastoid tubercle.[4]

P. cookei compared to Notharctus tenebrosus (left), an early true primate. Both come from Eocene Wyoming, though the latter is slightly geologically younger.

Although the placement of the Plesiadapis lineage is still up for debate, the consensus in the 1970s was that they were closest to early tarsier-like primates.[5] Plesiadapiformes have also been proposed as a nonprimate sister group to Eocene-Recent primates. A study done in 1987 linked Plesiadapiformes with adapids and omomyids through nine shared-derived features, six of which are cranial or dental: (1) auditory bulla inflated and formed by the petrosal bone, (2) ectotympanic expanded laterally and fused medially to the wall of the bulla, (3) promontorium centrally positioned in the bulla, and large hypotympanic sinus widely separating promontorium from the basisphenoid, (4) internal carotid entering the bulla posteriolaterally and enclosed in a bony tube, (5) nannopithex fold on the upper molars, and (6) loss of one pair of incisors.[4]

In 2013, a phylogenetic analysis that includes also the basal primate Archicebus positions Plesiadapis firmly outside of the Primates, as a sister group to both Primates and Dermoptera.[6]


Anatomy and remains

[edit]
Restoration

Plesiadapis is one of the most completely known early primatomorphs, with a significant amount of the skeleton known. Though, the skeleton is mostly known from P. gidleyi and the relatively late (derived) P. tricuspidens. The skull is overall reminiscent of a lemur, though it lacks the postorbital bars (a vertical bar bordering the posterior margin of the eye socket). The brain was probably relatively large compared to similarly sized contemporary mammals, namely the arctocyonids. The dental formula is usually 2.1.3.31.1.3.3, with two incisors, one canine, three premolars, and three molars in either half of the upper jaw; and one incisor, one canine, three premolars, and one molar in either half of the lower jaw. The incisors are quite long. Already, Plesiadapis had lost the first premolar from the mammalian common ancestor, but later primatomorphs would lose the second premolar as well. P. dubius consistently lacks the lower second premolar, and about half of P. rex specimens lack it too. P. gidleyi and European Plesiadapis lack the lower canines.[7]

P. cookei cast of a specimen from Wyoming. At the AMNH.

The skeletal adaptations are consistent with a largely arboreal lifestyle in the trees. The sacrum of P. gidleyi is similar to that of the eastern gray squirrel, though lacking the strong spines. The humerus is robust and features a strong S-curve.[7]

Nearly all of what is known about the anatomy of plesiadapiforms comes from fragmentary jaws and teeth, so most definitions of plesiadapiform genera and species are based on dentition. Plesiadapis' dentition shows a functional shift toward grinding and crushing in the cheek teeth as an adaptation towards increasing omnivory and herbivory.[8] The skull of Plesiadapis is relatively broad and flat, with a long snout with rodent-like jaws and teeth and long, gnawing incisors separated by a gap from its molars. Orbits are still directed to the side, unlike the forward-facing eyeballs of modern primates that enable three-dimensional vision.[7] Plesiadapis had mobile limbs that terminated in strongly curved claws, and it sported a long bushy tail which is preserved in the Menat skeletons. The way of life of Plesiadapis has been much debated in the past. Climbing habits could be expected in a relative of the primates, but tree-dwelling animals are rarely found in such high numbers. Based on this and other evidence, some paleontologists have concluded that these animals were mainly living on the ground, like today's marmots and ground squirrels.[2] However, more recent investigations have confirmed that the skeleton of Plesiadapis is that of an adept climber, which can be best compared to tree squirrels or to tree-dwelling marsupials such as possums.[7] The short, robust limbs, the long, laterally compressed claws, and the long, bushy tail indicate that it was an arboreal quadruped. Remains found showed that it had a body mass of around 2.1 kilograms (4.6 lb).[9]

Palaeoecology

[edit]

Large Plesiadapis species in Europe evolved to become increasingly adapted for folivory across the Palaeocene-Eocene boundary, although they were not as adapted for a folivorous diet as Platychoerops daubrei.[10]

References

[edit]
[edit]
Revisions and contributorsEdit on WikipediaRead on Wikipedia

Plesiadapis

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from Grokipedia
Plesiadapis is an extinct genus of early euarchontan mammals belonging to the family Plesiadapidae within the order Plesiadapiformes, often regarded as stem primates or close relatives due to shared traits such as large forward-projecting upper incisors and arboreal adaptations, though lacking forward-facing orbits and relying on claw-based clinging rather than nail-tipped grasping extremities. This genus, characterized by its arboreal adaptations including long claws for clinging to tree trunks and branches, inhabited forested environments across North America and Europe during the late Paleocene (Tiffanian land mammal age) to early Eocene (Wasatchian) epochs, approximately 59 to 54 million years ago. With body masses of about 1.5 to 2.5 kilograms depending on the species, Plesiadapis featured specialized dentition, including tricuspate upper incisors and broad, low-crowned molars suited for an omnivorous or folivorous diet involving seeds, fruits, and foliage. Recent studies reinforce their position as stem primates within Euarchonta. The genus encompasses at least nine recognized species, including P. cookei, P. tricuspidens, P. dubius, P. fodinatus, P. praecursor, P. churchilli, P. anceps, P. gingerichi, and P. simonsi, with fossils primarily recovered from localities in the (such as and ) and the in . These species exhibit morphological variation reflecting dietary and locomotor diversity; for instance, P. cookei shows more specialized folivorous adaptations in its molars, while P. tricuspidens appears more generalized and omnivorous. Postcranial skeletons reveal a quadrupedal arboreal lifestyle, with features like a robust , ovoid radial head, and deep ungual phalanges enabling clambering, claw climbing, and possibly suspensory behaviors on large-diameter supports, akin to modern arboreal squirrels. Plesiadapis played a pivotal role in the early radiation of euarchontans following the Cretaceous-Paleogene extinction, serving as a biostratigraphic marker for late faunas and providing critical evidence for the gradual evolution of traits such as enhanced manual and pedal grasping. Although the genus declined toward the -Eocene boundary, likely due to climatic changes and from emerging euprimates, its well-preserved fossils—including nearly complete skulls, dentitions, and partial skeletons—continue to inform debates on the phylogenetic position of plesiadapiforms as either basal or a to + Dermoptera. Endocranial studies further highlight its primitive brain organization, with low encephalization quotients (0.12–0.34) and exposed , underscoring a transitional morphology between archaic mammals and modern .

Taxonomy

Discovery and Etymology

The genus Plesiadapis was originally described by French paleontologist Paul Gervais in 1877, based on a partial left recovered from Thanetian (late ) deposits in the of . The , Plesiadapis tricuspidens, was named for the distinctive three-cusped morphology of its enlarged lower , a feature prominent in the specimen housed at the Muséum National d'Histoire Naturelle in . The etymology of Plesiadapis combines the Greek prefix "plesio-" (meaning "near" or "almost") with "Adapis," referencing the Eocene adapoid primate genus Adapis first described by in 1822; this nomenclature reflected Gervais's initial perception of Plesiadapis as a close relative to early Eocene known at the time. Early 20th-century paleontological expeditions in , led by figures such as James W. Gidley, uncovered the first Plesiadapis fossils from late formations in and , including the species P. rex named in 1923, thereby establishing the genus's transatlantic distribution between and during the Late . Due to the abundance and rapid morphological evolution of Plesiadapis species, they have since played a key role in biostratigraphic zonation of Late continental deposits in both and , with distinct species assemblages defining local mammalian biozones such as the middle and upper Tiffanian land-mammal ages.

Classification and Phylogeny

_Plesiadapis is the of the Plesiadapidae, placed within the extinct order , which comprises early euarchontan mammals that represent a key group in the radiation of higher mammals following the Cretaceous-Paleogene extinction. The genus includes over 10 recognized species, distributed across late and early Eocene deposits in and , with the being P. tricuspidens. Historically, Plesiadapis was classified in close relation to tarsiers based on shared dental specializations, such as enlarged incisors and reduced premolars, or to dermopterans (flying lemurs) due to similarities in tarsal morphology. The phylogenetic position of , including Plesiadapis, remains debated, with cladistic analyses supporting positions either as stem primates ( to crown ) or as stem euarchontans sister to the clade + Dermoptera () within ; this uncertainty is informed by features such as the auditory bulla, including contributions from the entotympanic in some reconstructions. Key synapomorphies diagnosing Plesiadapis within Plesiadapidae include forward-directed orbits providing enhanced stereoscopic vision and procumbent lower incisors adapted for gnawing, though the genus lacks the complete postorbital bar and other derived cranial closures typical of crown primates. Recent phylogenetic studies have refined understanding of Plesiadapis diversification in , revealing a morphological succession among across the Paleocene-Eocene boundary, with at least two co-occurring forms at sites like Cernay, indicating adaptive trends toward folivory and no direct continuity with Eocene platychoeropines. These findings underscore the role of Plesiadapidae in the post-Cretaceous-Paleogene radiation of , filling ecological niches vacated by non-avian dinosaurs.

Anatomy

Cranial and Dental Features

The skull of Plesiadapis exhibits a lemur-like morphology, characterized by a long, rodent-like snout and large, forward-facing orbits that suggest adaptations for diurnal or crepuscular vision, though the orbits are non-convergent and lack a postorbital bar typical of euprimates. The external auditory meatus is ossified and tube-like, likely of ectotympanic origin, forming part of a small, petrosal-dominated auditory bulla that differs from the more complex bullae in later primates. This cranial configuration, with a posteriorly positioned braincase occupying about 39% of skull length, underscores its primitive euarchontan affinities while highlighting sensory specializations. Dentition in Plesiadapis follows the formula 2.1.3.3/3.1.3.3, featuring enlarged, procumbent lower incisors with a rodent-like morphology suited for gnawing on bark or hard material, and upper incisors with three distinct cusps for precise occlusion. The cheek teeth display molarized premolars and lophodont cusps on the molars, with crests oriented for shearing and grinding, indicative of folivorous tendencies involving tough vegetation. Enamel microstructure includes radial prisms and Hunter-Schreger bands in the incisors for enhanced wear resistance, while cheek tooth enamel shows moderate thickness and prismatic structure to withstand abrasive folivory without rapid occlusion loss. Brain size in Plesiadapis is enlarged relative to other mammals, with an (EQ) of 0.12–0.34, reflecting moderate expansion beyond basal therian levels and comparable to early . The reveals a smooth, domed covering about 22% of the surface, with exposed and downward-shifted olfactory bulbs, suggesting enhanced olfaction and vision over tactile reliance in contemporary non-euarchontans. Recent micro-CT analyses of the oldest known plesiadapiform cranium (Plesiolestes nacimienti, early Torrejonian, ~64 Ma) reveal internal cranial details, including a caudal nasal expansion, intraorbital , and optic configuration, confirming early euarchontan traits such as a petrosal auditory bulla with reduced entotympanic contributions and minimally ossified elements akin to treeshrews. These features, absent in more primitive mammals, highlight transitional sensory anatomy in the lineage leading to crown .

Postcranial Features

Plesiadapis species exhibited body sizes ranging from approximately 0.5 to 2.2 kg, with P. tricuspidens estimated at around 2.1 kg, comparable in scale to a large or small . For instance, P. cookei, one of the larger species, yielded body mass estimates of 1.8–2.1 kg based on measurements from an associated partial . The postcranial skeleton featured elongated forelimbs adapted for arboreal , with mobile joints and strongly curved phalanges terminating in sharp claws rather than , facilitating grasping of tree trunks and branches. Hindlimbs were slightly longer than forelimbs, supporting leaping behaviors among larger supports, as evidenced by femoral morphology with a medially extended lesser for flexion and abduction. A long, bushy tail, inferred from soft-tissue impressions preserving fur patterns and outline at the Menat locality in , likely aided balance during quadrupedal arboreal locomotion. The vertebral column included a flexible lumbar region with six vertebrae, promoting sagittal bending essential for quadrupedal maneuvering in trees, while the scapula possessed a well-developed supraspinous fossa that enhanced shoulder mobility for overhead reaching and climbing. Postcranial preservation is limited but notable, with rare articulated or associated skeletons from —such as the partial skeleton UM 87990 of P. cookei including 43 vertebrae and most limb elements—and from at , where impressions reveal soft-tissue details including claws and tail fur.

Fossil Record

Temporal and Geographic Range

Plesiadapis fossils are known from the late to early Eocene epochs, corresponding to the Tiffanian and Clarkforkian North American Land Mammal Ages (NALMAs) in , with limited extension into the earliest Wasatchian NALMA, spanning approximately 59 to 54 million years ago (Ma). The genus exhibits peak diversity during the Paleocene-Eocene Thermal Maximum (PETM) around 56 Ma, a period of rapid global warming that facilitated faunal turnover and dispersal. In , Plesiadapis originated and achieved its greatest abundance and species richness in the western interior, serving as an index fossil for late strata due to its stratigraphic utility in biozonation schemes based on evolving dental morphology across Tiffanian substages. Geographically, Plesiadapis was initially endemic to , with the earliest and most extensive records from the Fort Union Formation in (e.g., , Hoback Basin) and (e.g., , Bear Creek), as well as the Piceance Creek Basin in and scattered sites in . Dispersal to occurred around 56 Ma via the Thulean connecting to and , marking a biogeographic expansion during the PETM. European occurrences are concentrated in the of (e.g., Cernay, Berru, , Petit-Pâtis formations), with additional finds in (e.g., Hainin, Dormaal), (e.g., Walbeck), and (e.g., at ). The genus' distribution reflects early North American endemism followed by transatlantic colonization, with European populations showing parallel evolutionary trends to North American ones but persisting slightly longer into the early Eocene before , potentially linked to post-PETM and habitat shifts around 54 Ma. This pattern underscores Plesiadapis' role in late biochronology, where species succession aids in correlating continental deposits.

Known Species and Major Finds

The genus Plesiadapis encompasses approximately 10 valid , documented from late to early Eocene deposits in and , though taxonomic debates persist with some synonymies. The , P. tricuspidens, was originally described from Cernay-le-Lee () and represents the earliest named member of the genus, with subsequent referrals from multiple European localities including Walbeck () and Berru () . North American include P. gidleyi from the Tiffanian North American Land Mammal Age (NALMA), known from partial skeletons preserving portions of the postcranial skeleton, and P. churchilli from Tiffanian sites in the () and () . Additional valid are P. fodinatus (Tiffanian, and ), P. cookei (late , and ), P. dubius (Tiffanian-Clarkforkian), P. rex (Tiffanian), P. remensis (late , ), P. walbeckensis, and P. praecursor, with P. gingerichi, P. simonsi, and P. anceps also recognized in various assessments, though P. anceps is sometimes treated as a junior of P. tricuspidens or reassigned based on dental morphology . Major fossil assemblages of Plesiadapis highlight its diversity and provide key insights into its morphology and evolution. The Clark Fork beds in the Bighorn Basin (Wyoming, USA) yield diverse dentition across multiple species, including P. cookei and P. gidleyi, with specimens from the late Paleocene (Cf1-Cf2 zones) preserving isolated teeth and partial jaws that document biochronological transitions . The Menat lagerstätte (Puy-de-Dôme, France), a late Paleocene (approximately 59-60 Ma) maar lake deposit, has produced articulated skeletons of P. cookei with exceptional soft-tissue preservation, including carbonaceous films revealing skin impressions and a long, bushy tail—unique among Paleocene mammals and indicative of arboreal adaptations . Recent discoveries at Petit-Pâtis (Paris Basin, France) from latest Paleocene (MP6a) strata include a new species, P. ploegi, alongside referrals to P. tricuspidens, revealing morphological shifts across the Paleocene-Eocene Thermal Maximum (PETM), such as increased body size (up to 20% larger than earlier conspecifics), enhanced molar cresting for folivory, and greater dental specialization in the lower fourth premolar . Ongoing research continues to refine the fossil record of Plesiadapis. A 2023 study on latest material from French sites, including Petit-Pâtis and nearby quarries, demonstrates evolutionary transitions toward more derived euarchontan traits, such as improved occlusion and simplification, bridging P. tricuspidens with early Eocene forms . Micro-CT analysis of specimen KUVP 9557, the oldest known cranial remain of a plesiadapiform (~62 Ma, Torrejonian NALMA, ), pertains to the related palaechthonid Plesiolestes nacimienti but informs basal euarchontan anatomy relevant to Plesiadapis phylogeny, revealing primitive features like a reduced postorbital process and expanded nasal region . Preservation is predominantly dental, with isolated teeth and jaw fragments comprising over 90% of specimens; rare postcrania, including limb elements suggestive of scansorial locomotion, occur at sites such as Bear Lodge (, USA) and Hainin (), the latter yielding early plesiadapiform isolates that contextualize genus origins .

Paleoecology

Habitat and Environment

Plesiadapis inhabited paratropical forest environments in during the late , primarily within the Fort Union Formation of the Western Interior, characterized by humid subtropical conditions with mean annual temperatures around 20°C and annual precipitation exceeding 130 cm. These settings featured meandering fluvial systems, peat swamps, and low-diversity broadleaf forests dominated by angiosperms such as (e.g., Corylites) and evergreen like Glyptostrobus, supporting arboreal lifestyles amid waterlogged, frost-free floodplains devoid of significant dry seasons or redbed soils. and megafossil records from these deposits indicate mixed deciduous- woodlands with diverse , reflecting a stable, warm, equable climate that fostered the proliferation of early euarchontans. In , Plesiadapis occupied similar subtropical woodland habitats in the , as evidenced by fossils from the Cernay Formation, where continental deposits suggest warm, temperate to humid conditions with forested surroundings influenced by latest warming events. These environments, dated to approximately 56 Ma, featured systems and palynofloras indicative of angiosperm-rich vegetation, paralleling North American paratropical settings but with regional floral variations tied to the Thanetian stage. Coexisting biota included early carnivorans (e.g., early miacids), highlighting biotic responses to pre-PETM climatic fluctuations that expanded wooded areas across . Plesiadapis declined around the Paleocene-Eocene Thermal Maximum (PETM) hyperthermal event around 55.5 Ma, a period of rapid CO₂-driven global warming that elevated temperatures by 5–8°C and promoted the expansion of humid, angiosperm-dominated forests across both continents. This climatic perturbation, linked to massive carbon release, enhanced forested habitats and facilitated faunal dispersals, with Plesiadapis assemblages associated with inaugural appearances of modern mammal orders like (e.g., Paramys) and carnivorans (e.g., early miacids). Taphonomic evidence from lagerstätten such as in reveals volcanic lake deposits influenced by phreatomagmatic activity, preserving riparian zones around forested lakeshores in a warm, humid ecosystem conducive to exceptional fossilization of plants, , and vertebrates.

Diet and Locomotion

Plesiadapis species exhibited a primarily folivorous diet, supplemented by insectivory, as inferred from dental morphology and microwear . Dental microwear patterns on cheek teeth indicate browsing on soft , such as leaves and fruits, with shearing crests facilitating the processing of fibrous material. Stable carbon isotope analyses from associated mammal teeth, including those of plesiadapiforms, reveal a diet dominated by C3 plants typical of forested understories, supporting a herbivorous niche in closed-canopy environments. In the latest Paleocene, French fossil finds from the Paris Basin document an evolutionary trend toward increased herbivory in Plesiadapis, evidenced by molarization of premolars and enhanced crest development on molars, allowing better processing of tougher foliage leading into warming climates associated with the Paleocene-Eocene Thermal Maximum (PETM). Dietary differentiation among species, such as greater folivory in P. cookei and P. russelli compared to P. tricuspidens, suggests niche partitioning with other plesiadapiforms to reduce competition for resources. Locomotion in Plesiadapis was scansorial, combining quadrupedal walking and in arboreal habitats, akin to modern squirrels. Phalangeal curvature and mediolaterally compressed ungual phalanges indicate reliance on claws for traction during vertical clinging and ascent on tree trunks and branches. Postcranial adaptations, including robust limb elements suited to large-diameter supports, further support an arboreal lifestyle, with limited evidence for gliding due to anatomical constraints. Behavioral ecology points to solitary or small-group , consistent with the nocturnal habits and small body size of early euarchontans, enabling efficient exploitation of dispersed arboreal resources without extensive social coordination.

References

  1. https://www.pnas.org/doi/10.1073/pnas.0610579104
  2. https://www.researchgate.net/publication/285776448_Plesiadapiformes
  3. https://pmc.ncbi.nlm.nih.gov/articles/PMC3953834/
  4. https://onlinelibrary.wiley.com/doi/10.1002/evan.21526
  5. https://www.researchgate.net/publication/235625417_Locomotor_adaptations_of_Plesiadapis_tricuspidens_and_Plesiadapis_n_sp_Mammalia_Plesiadapiformes_as_reflected_on_selected_parts_of_the_postcranium
  6. https://www.researchgate.net/publication/30849239_New_North_American_Plesiadapidae_Mammalia_Primates_and_a_Biostratigraphic_Zonation_of_the_Middle_and_Upper_Paleocene
  7. https://paleobiodb.org/classic/basicTaxonInfo?taxon_no=38198
  8. https://pmc.ncbi.nlm.nih.gov/articles/PMC10776232/
  9. https://pmc.ncbi.nlm.nih.gov/articles/PMC1783133/
  10. https://www.researchgate.net/publication/327105076_Evolution_of_plesiadapid_mammals_Eutheria_Euarchonta_Plesiadapiformes_in_Europe_across_the_PaleoceneEocene_boundary_implications_for_phylogeny_biochronology_and_scenarios_of_dispersal
  11. https://onlinelibrary.wiley.com/doi/abs/10.1002/ajpa.21211
  12. https://pubmed.ncbi.nlm.nih.gov/40080934/
  13. https://lsa.umich.edu/paleontology/publications/papers-on-paleontology/skeleton-of-late-paleocene-plesiadapis-cookei--mammalia--euarcho.html
  14. https://lsa.umich.edu/paleontology/news-events/all-news/search-news/new-skeleton-of-plesiadapis.html
  15. https://www.researchgate.net/publication/22043722_Phylogenetic_relationships_of_Plesiadapis_postcranial_evidence
  16. https://deepblue.lib.umich.edu/bitstream/handle/2027.42/48615/ID478.pdf
  17. http://deepblue.lib.umich.edu/bitstream/2027.42/61362/1/SecordPapers35DeepBlue.pdf
  18. https://pubs.geoscienceworld.org/rmg/article-pdf/53/2/75/4574448/05302075.pdf
  19. https://onlinelibrary.wiley.com/doi/10.1111/jbi.12310
  20. https://hal.science/hal-04166484/document
  21. https://paleobiodb.org/classic/basicTaxonInfo?taxon_no=50440
  22. https://www.researchgate.net/publication/325228584_Plesiadapid_mammals_from_the_latest_Paleocene_of_France_offer_new_insights_on_the_evolution_of_Plesiadapis_during_the_Paleocene-Eocene_transition
  23. https://depositsmag.com/2016/08/11/a-palaeocene-lagerstatte-in-france/
  24. https://www.tandfonline.com/doi/abs/10.1080/14772019.2016.1237582
  25. https://terra.rice.edu/department/faculty/morganj/ESCI536/Readings/Wilf-MultiQuarry.pdf
  26. https://npshistory.com/publications/fous/nrr-2015-1004.pdf
  27. https://www.pnas.org/doi/10.1073/pnas.2505795122
  28. https://pmc.ncbi.nlm.nih.gov/articles/PMC3906055/
  29. https://www.researchgate.net/publication/326081432_The_Konservat-Lagerstatte_Menat_Paleocene_France_-_an_overview_and_new_insights
  30. https://doi.org/10.1080/02724634.2018.1460602
  31. https://pubmed.ncbi.nlm.nih.gov/20034012/
  32. https://www.sciencedirect.com/topics/pharmacology-toxicology-and-pharmaceutical-science/forager
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